Wisconsin's Biodiversity As a Management Issue

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Wisconsin’s Biodiversity as a Management Issue

A REPORT TO DEPARTMENT OF NATURAL RESOURCES MANAGERS MAY 1995 Wisconsin’s Biodiversity as a Management Issue

A Report to Department of Natural Resources Managers

May 1995

Department of Natural Resources P.O. Box 7921 Madison, Wisconsin 53707 NATURAL RESOURCES BOARD

Herbert F. Behnke, Chair Neal W. Schneider, Secretary James E. Shawano Janesville Tiefenthaler, Jr. Brookﬁeld Trygve A. Solberg, Vice Chair Betty Jo Nelsen Minocqua Shorewood Stephen D. Willett Phillips Mary Jane Nelson Holmen

OFFICE OF THE SECRETARY George E. Meyer, Secretary Ronald L. Semmann, Deputy Secretary Maryann Sumi, Executive Assistant

This report was written for Department of Natural Resources managers to provide them with a context for their work. The Natural Resources Board met on April 27, 1995 to review the report. At this meeting, the Board accepted the Department’s recommendations, as follows: Print and distribute the report “Wisconsin’s Biodiversity as a Management Issue,” and use it to continue an open dialogue with the Department’s partners and customers; Adopt the four strategic recommendations in the revised report (as summarized on pages 7-8 and described on pages 37-39), and use them to guide policy development; and Further analyze the implications of the “possible actions” listed for the seven biological community types in the report. The Department will work with the Board to determine the most appropriate sequence for these analyses and to set priorities for using them to develop policy. In moving to approve the report, the Board also stated that its approval “is made with the understanding that the Natural Resources Board does not endorse, recommend, or sanction the use of this report to judge whether or not a management or regulatory act is appropriate or inappropriate.” Wisconsin’s Biodiversity as a Management Issue

A REPORT TO DEPARTMENT OF NATURAL RESOURCES MANAGERS MAY 1995

PREPARED BY: James Addis, Ronald Eckstein, Anne Forbes (Leader 1994-95), DuWayne Gebken, Richard Henderson, John Kotar, Betty Les (Leader 1993-94), Paul Matthiae, Wendy McCown, Steven Miller, Bruce Moss, David Sample, Michael Staggs, and Kristin Visser

ASSISTED BY: Cathy Bleser, Alan Crossley, June Dobberpuhl, Robert DuBois, Robert Dumke, Eric Epstein, Don Fago, Jonathan Gilbert, Beth Goodman, Mary Hamel, Robert Hay, David Heath, Ralph Hewett, Mark Heyde, Randy Hoffman, Fred Iausly, David Ives, Roy Jacobson, Martin Jennings, Laura Komai (staff assistant), John Lyons, Mark Martin, Thomas Meyer, Thomas Mickelson, William Moorman, Mike Mossman, Richard Narf, Kathy Patnode, Tom Pellett, Tom Ruzycki, Dennis Schenborn, Dave Siebert, William Smith, Jeffrey Smoller, William VanderZouwen, Dreux Watermolen, Wendy Weisensel, and Darrell Zastrow

PREPARED FOR: Division of Resource Management

James Addis (Administrator), Steven Miller (Deputy Administrator), Howard S. Druckenmiller, Robert Dumke, Carl Evert, Charles Higgs, Thomas Hauge, Lee Kernen, Charles Pils, and David Weizenicker

EDITED AND PRODUCED BY: Jeanne Gomoll (graphics and layout), Signe Holtz, Rebecca Isenring, Michelle Jesko, Laura Komai, Betty Les (Leader), and Wendy McCown TABLE OF CONTENTS

2 Executive Summary 72 Southern Forest 116 Grassland Communities 2 Purpose of this Report Communities 116 Description 2 What is Biodiversity? 72 Description 117 Status 3 Ecological Issues 73 Status 122 Actions Causing Concern 4Wisconsin’s Biological 79 Actions Causing Concern 123 Socio-Economic Issues Communities 80 Socio-Economic Issues 123 Potential for Community 7 What This Report Proposes 80 Potential for Community Restoration 9 Some Continuing Policy Restoration 124 Possible Actions Questions 81 Possible Actions 128 Case Study: The Glacial Habitat 84 Case Study: The Baraboo Hills: Restoration Area: An Approach 10 Biodiversity: Issues and Partners Protecting and for Restoration of Grassland Managing in an Ecosystem Communities Implications Context 129 Literature Cited 10 Importance of Biodiversity 85 Literature Cited 12 Historical and Current 130 Wetland Communities Perspectives in Resource Management 88 Oak Savanna 130 Description 18 Ecological Issues Communities 132 Status 30 Addressing Biodiversity Through 88 Description 140 Actions Causing Concern Ecosystem Management 90 Status 142 Socio-Economic Issues 37 Recommendations 92 Actions Causing Concern 142 Potential for Community Restoration 92 Socio-Economic Issues 143 Possible Actions 40 Overview of Wisconsin’s 92 Potential for Community Biological Communities Restoration 145 Case Study: Restoring a Prairie Wetland Landscape in Southern 93 Possible Actions 40 Determinants of Biodiversity Wisconsin 95 Case Study: Kettle Moraine 147 Literature Cited 40 Impact of Glaciation Oak Opening: Natural 41 Impact of Other Natural Community Protection and Disturbance Restoration through Master 150 Aquatic Communities 41 Human-Induced Disturbance Planning 150 Description 43 The Situation Today 96 Literature Cited 162 Past Status 49 Literature Cited 98 Oak and Pine Barrens 164 Past and Present Actions Causing Concern 50 Northern Forest Communities 187 Present Status Communities 98 Description 192 Projected Status 50 Description 102 Status 194 Socio-Economic Issues 50 Status 108 Actions Causing Concern 195 Potential for Community Restoration 62 Actions Causing Concern 108 Socio-Economic Issues 198 Possible Actions 62 Socio-Economic Issues 108 Potential for Community Restoration 202 Case Study: Habitat Restoration 63 Potential for Community Following Dam Removal on the Restoration 109 Possible Actions Milwaukee River at West Bend 64 Possible Actions 111 Case Study: Habitat Conservation Planning: 203 Literature Cited 68 Case Study: Marathon County Species Protection through Forest: Using GIS to Manage Ecosystem Management Forest Interior Habitat 220 Glossary 112 Literature Cited 68 Literature Cited 113 Additional Reading 226 Appendix A: Common and Scientiﬁc Names of Organisms Cited

238 Appendix B: Acknowledgements EXECUTIVE SUMMARY

This report provides a historical perspective on natural resources manage- CHAPTER ment, reviews the range of public values 1 relating to biodiversity, and explores DNR’s role in managing natural resources to conserve biodiversity. It presents ecosystem management as the framework that will help us balance human needs and values Executive with the conservation of biological diversity, and it proposes approaches and tools to help the Department and its partners move more fully into ecosystem manage- Summary ment. The report also offers an overview of the state’s seven major biological communities, describing each, documenting changes that have occurred since the early 1800s, outlining current issues, and suggesting PURPOSE OF THIS REPORT possible actions.

his report presents a Department strategy for the conservation of WHAT IS BIODIVERSITY? biological diversity. It provides Department of Natural Resources Biodiversity is a shortened form of the (DNR) employees with an term “biological diversity.” Simply stated, it overview of the issues associated is the entire spectrum of life forms and the with biodiversity and provides a common many ecological processes that support point of reference for incorporating the them. Biodiversity occurs at four interact- conservation of biodiversity into our ing levels: genetic diversity, species diver- management framework. It will be used as sity, community diversity, and ecosystem a discussion piece for dialogue with the diversity. Genetic diversity is the spectrum public and will be useful for Natural of genetic material carried by all the Resources Board members as they include individuals of a particular species. Species attention to biodiversity in the develop- diversity is the variety of species in a ment of public policy. geographic area, including not only the Our goal is sustainable ecosystems. number of species but also their relative These ecosystems, whether highly modiﬁed abundance and spatial distribution. by humans or largely natural, exhibit A community is an assemblage of ecological characteristics that maintain different plant and animal species, living biological diversity across all land uses. To together in a particular area, at a particular reach this goal we must develop manage- time, in speciﬁc habitats. Communities ment solutions that blend people’s needs usually are named for their dominant plant with nature’s capacity to sustain those species (for example, pine barrens, sedge needs over the long term. These manage- meadows, and oak savannas). Communities ment solutions must be founded in the range in size from less than an acre to perspective that humans are part of, not thousands of acres. Communities are apart from, the global ecosystem. Like all always changing, though often they change species, we depend on a viable biosphere. too slowly for humans to notice in our brief But unlike other species, we have it in our lifetimes. power to destroy the ecosystems on which An ecosystem includes not only we depend. Today’s decisions will have far- biological communities but also the reaching impacts on the choices and quality myriad, continuing interactions of biologi- of life available to us in the future. cal communities with their abiotic (non-

2 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE living) environment, including moisture, nas, and barrens. Euro-American settlement temperature, sunlight, soil, and many other brought many changes to this landscape, physical and chemical factors. Ecosystems, including suppression of fire, large-scale which range in size from minute to millions intensive agriculture, and urban and of acres, exhibit complex linkages among industrial development. Today, Wisconsin’s plants, animals and the physical and landscape is a mosaic of urban areas, farms, chemical environments. Ecosystem diver- commercial and recreational forests, lakes sity is largely determined by the amount and wetlands, and a small amount of land and complexity of these linkages. Ecosys- in protected natural areas. All the natural tems, like biological communities, are in a communities present in the early 1800s constant state of change, called “ecological have been significantly altered in function succession.” Succession is the progressive or size, with some existing today only as change through time of species composi- remnant areas. tion, organic structure, and energy flows Managing Wisconsin’s natural re- throughout an ecosystem. Human activities sources in the context of biodiversity and natural phenomena such as fire and requires that we understand the combina- tornados can alter succession. tion of forces that produced today’s land- Wisconsin is blessed with abundant scape and the effect of human activities on biodiversity. Located at the junction of biological communities and ecosystems. three of North America’s six biotic prov- Although these forces are complex, it is inces—the eastern deciduous forest, the important that we understand them, northern boreal forest, and the temperate support patterns of resource use consistent grasslands—we have a wealth of species with our goal of sustainable ecosystems, and natural communities. Approximately and accept responsibility for the problems 1,800 species of native plants and 657 raised by intensive human use of the species of native vertebrates have been landscape. These problems can be grouped identified in Wisconsin. In addition, there into three major categories for discussion: are thousands of species of nonvascular ecological simplification, fragmentation, plants and invertebrates. The challenge is and environmental pollution. to manage this diversity to conserve Ecological simplification means that Wisconsin’s biological heritage and preserve the interrelationships between organisms future management options. and their environments are reduced in number and complexity. Every organism in an ecosystem has one or more roles to play ECOLOGICAL ISSUES in sustaining that ecosystem. For example, bacteria and fungi cause dead trees to rot, When the glaciers receded from this providing nutrients for plants, which in part of the continent 10,000-12,000 years turn provide food for birds and small ago, humans moved into the area along mammals, which are then eaten by preda- with colonizing plants and animals. Native tors. If these natural processes are inter- Americans managed the landscape using rupted, ecological simplification can occur. fire, agriculture, and harvest of plants and Simplification is caused by loss of habitat, animals. They undoubtedly affected large loss of species in a community, and air and portions of the Wisconsin landscape. When water pollution that affects chemical and European and American settlers moved into physical processes. The addition of non- Wisconsin in the early 1800s, the state’s native species can also simplify biological landscape was characterized by extensive communities and ecosystems, disrupting forests, grasslands, wetlands, and a variety the food chain, destroying habitat for native of other large communities. Fire, often species, displacing native species, and purposefully set by Native Americans, was otherwise upsetting natural processes. a major factor in maintaining many of these Ecological simplification can also result communities, especially grasslands, savan- from land management practices that

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 3 EXECUTIVE SUMMARY

reduce natural variety on the landscape, such as filling wetlands or planting only WISCONSIN’S BIOLOGICAL COMMUNITIES one species of tree. The effects of simplifi- The location and extent of biological cation are often complicated and subtle, communities are determined by environ- reducing the number of species in an area, mental factors, including moisture, tem- and the genetic variety among individuals perature, soils, and climate. Natural factors, of a species. especially the glaciers but also windstorms, Fragmentation is the breaking up of fires, droughts, and floods, shaped large and continuous ecosystems and Wisconsin’s landscape. Human activities, communities into smaller areas surrounded beginning with Native American activities by altered or disturbed areas. Modern and continuing into today’s intensive use of civilization has greatly fragmented the land and water, have also had profound landscape. Farms have been created in the impacts on Wisconsin’s biological commu- middle of forests, and prairies have been nities. plowed for agriculture. Wetlands have been This report profiles seven major filled. Rivers and streams have been biological communities, which represent an dammed, interrupting corridors used for aggregation of the more numerous commu- animal movement and isolating populations nities described by scientists (especially and habitats. Some species, such as white- Curtis) in the 1950s. These seven commu- tailed deer, do well in these altered land- nities are northern forests, southern forests, scapes. However, many species of plants oak savannas, oak and pine barrens, and animals have declined in number as grasslands, wetlands, and aquatic systems. habitats have become too small to allow The term northern forest refers successful reproduction and isolated primarily to location rather than to any populations have lost genetic diversity. specific species composition. Northern Environmental pollution is the forests contain mixed deciduous and human-induced addition of many types of coniferous forests found in a distinct substances to air, land, and water in climatic zone that occurs north of a roughly quantities or at rates that harm organisms, S-shaped transition belt known as the habitats, communities, ecosystems, or “tension zone” that runs from northwest to human health. Water pollution destroys southeast Wisconsin. Early forest surveys aquatic habitats and kills aquatic life indicate that northern forests consisted of a through toxicity, by destroying habitat, or mosaic of young, mature, and “old-growth” by using up dissolved oxygen. Acid rain forests composed of pines, maples, oaks, and air-borne contaminants such as heavy birch, hemlock, and other hardwood and metals and pesticides affect both aquatic conifer species. “Old growth” is defined as a and terrestrial plants and animals. community in which the dominant trees Despite the problems caused by are at or near biological maturity. ecological simplification and fragmentation, The late 19th- and early 20th-century both can be consistent with management loggers cut over virtually the entire north- objectives. Enhancing populations of ern forest. Conditions remaining after certain plant and animal species, providing logging were more favorable to hardwood forest and agricultural products, and than to pine, resulting in limited pine accommodating other human activities are reproduction after logging ceased. Today, obviously important and necessary. The key most areas that were formerly in pine are is to take a landscape-scale view, seeing the now in oak, maple, and aspen, and the age overall mosaic of land and water use in structure of the northern forest is consider- Wisconsin; to recognize the impacts of our ably different than it was before logging proposed actions; to clarify where, when, occurred. Likewise, distribution and and why these actions are desirable; to abundance of animals in the northern know the trade-offs; and to preserve forests have been altered dramatically, with options for future generations. some species declining in numbers and

4 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE others, finding the current forest advanta- Oak savannas are characterized by NORTHERN geous, increasing their populations. open grassland areas interspersed with FOREST COMMUNITIES The major biological issue relating to trees, especially oaks. Savannas, historically the northern forests is that they have been found in southern and western Wisconsin, managed on a stand-by-stand basis with were the gradation between the great little regard for sustaining landscape or prairies and the eastern deciduous forests. regional diversity. The major forest cover The savannas were perpetuated by fire. In SOUTHERN FOREST types are managed largely for harvest at an the early 1800s, Wisconsin had perhaps 5.5 COMMUNITIES economically desirable rotation age, which million acres of oak savanna, virtually all of perpetuates the limited age structure of which has been destroyed for farming and northern forest communities. Fortunately, urban development or has succumbed to there is great potential for maintaining and natural succession as fire has been sup- even enhancing biological diversity in the pressed. Oak savanna is now virtually non- OAK SAVANNA northern forests. We have lost very few existent in Wisconsin, with only a few COMMUNITIES plant or animal species from the area. The remnant areas remaining. key is to use a landscape approach that can Many animal species associated with produce all the successional stages, from savannas have managed to find surrogate young trees to old growth, within large and habitats such as wooded pastures, lawns, small stands in the forest mosaic. and small woodlots. Savanna vegetation has GRASSLANDS Early European observers recognized not fared as well. Many savanna plant COMMUNITIES southern forests (those south of the species are now uncommon and found only tension zone) as distinct from the northern on the fringes of oak woods, brushy areas, types because of the predominance of oaks and lightly grazed pastures. Fortunately, and general absence of conifers. They also oak savanna restoration is possible, through OAK & PINE noted the relative openness or park-like the use of fire and perhaps light grazing. BARRENS appearance, created by the lack of small Oak and pine barrens, like savannas, COMMUNITIES trees and shrubs. There is evidence that depend on fire to maintain their unique these southern forests were shaped by fire character. These communities, which are in the previous 5,000-6,000 years. Begin- found in central and northern Wisconsin ning in the early 1800s, the southern where soils are poor, are characterized by forests were cleared for farming or har- sparse scrub pine or oak scattered among vested for lumber, fuel, and railroad ties. shrubs, brush, and grasses. In the early Fire was also suppressed. As a result, the 1800s, barrens covered about 4.1 million southern forests are today severely frag- acres of Wisconsin. Barrens communities mented into small woodlots. Remaining have been destroyed by agriculture and forest cover is heaviest in the southwest urban development, or have succeeded to coulee region. The large herbivores and forests in the absence of fire. Only a few carnivores originally found in the southern remnant areas remain. As with other forest, including buffalo, elk, and cougar, communities, many of the plant and animal are gone. These species and others were species associated with barrens have unable to survive on increasingly smaller managed to survive, though often in patches of appropriate habitat and were reduced numbers. The potential for restora- also affected by land development practices tion of barrens areas on public and private and over-harvest by settlers. Some bird lands is good if controlled burning and species (notably the passenger pigeon) have cutting are used as management tools. also been lost, though many remain in Wisconsin’s grassland (prairie) reduced numbers. communities, characterized by the absence Forestry practices that reduce frag- of trees and large shrubs and the domi- mentation, increase the use of fire, and nance of grass and forb species, are at the manage the old-growth forests that remain periphery of the extensive North American on public lands are key to restoring biologi- mid-continent grassland biome, which lies cal diversity on southern forests. south and west of the state. These grass-

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 5 EXECUTIVE SUMMARY

lands, which grew up 5,000-6,000 years restored by simply blocking drainage and ago after the glaciers retreated, were allowing water levels to rise; others require maintained by fire and probably by large decades or longer to restore natural func- grazing animals such as buffalo. Prior to tions. Euro-American settlement, Wisconsin had When the glaciers receded, they left about 3.1 million acres of prairies, of which behind a variety of aquatic communities, WETLAND COMMUNITIES almost one million acres were a wet prairie including springs, ponds, lakes, streams, type known as “sedge meadow.” and rivers. Within this grouping is a wide The grassland biome has been de- variety of systems, differing in size, fertility graded throughout its range, generally from (lakes), water temperature (streams), and farming and grazing, but also from urban geographic area. Wisconsin has 620 miles development. Some prairie areas also grew of Great Lakes shoreline, more than 14,000 AQUATIC up into trees and shrubs as fire was con- lakes covering a total of a million acres, and COMMUNITIES trolled. Thus, the prairie community has more than 33,000 miles of rivers and been severely fragmented, with only a few streams, including 1,500 impoundments. remnant areas left. Prairies, along with oak Simplification of many aquatic savannas, are the most endangered natural systems has occurred due to introduction communities in Wisconsin. As a result, an of exotic species of fish such as carp and estimated 15%-20% of the state’s original lamprey, which successfully compete grassland flora is now considered rare here. against many native species, as well as to Grassland mammals and birds adapted the large-scale destruction of shorelines and better, using “surrogate” grasslands such as other habitats. Fragmentation has been pastures for their survival needs. Managed caused by dam construction. Dams block use of fire, removal of trees and shrubs, movement of fish and other aquatic organ- light grazing, and perhaps some crop isms, isolating populations, and sometimes production will aid prairie restoration. resulting in loss of genetic diversity and Populations of grassland mammals and eventual extirpation of species in a portion birds can also be restored by establishing of the river. Dam construction also changes “surrogate” grassland habitat on both water flow and temperatures, resulting in private and public lands. changes in habitat that can lead to extirpa- Wetlands, which are lands on which tion of species. Other activities that create soils or substrate is periodically saturated pollution or cause simplification and with or covered by water, occupied an fragmentation of aquatic systems include estimated ten million acres (nearly one- agricultural and urban development and third of Wisconsin’s land area) in the early resulting runoff, channelization of streams, 1800s. Wetlands have been subject to shoreline development and resulting loss of intense modification, mainly through habitat and spawning areas, and industrial draining and filling for agriculture and and urban development and resulting urban development. Today, about 5.3 effluent and runoff. In addition, some million acres of wetlands remain. Nearly all fisheries management activities such as the remaining wetlands have suffered from indiscriminate stocking have also contrib- the effects of fragmentation and simplifica- uted to disturbance of aquatic communi- tion. ties. Current federal, state, and local Although the abundance of many fish regulations and land acquisition programs species has been greatly altered, most have considerably slowed wetland loss. native fish species are still abundant and However, nonagricultural filling of wet- self-sustaining. This relative health of lands, especially along lake shores, contin- aquatic communities allows us to focus ues to threaten some wetlands. In addition, attention on identifying and restoring the invasion of exotics such as purple specific degraded communities as well as loosestrife pose a threat to wetland ecology. protecting species with declining numbers. Some wetland communities are easily River and stream communities respond

6 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE quickly to habitat protection and restora- the breadth of what needs to be done to tion. Lake communities respond more conserve biodiversity by working on slowly. It is important to shift aquatic public lands alone. community management from a single- species focus to an ecosystem-management Develop innovative and proactive focus. information and education strategies for Department staff and the public regarding biodiversity and its relation to WHAT T HIS REPORT PROPOSES ecosystem management.

DNR’s mission is to conserve, protect, The second type of recommendations and manage both individual species and are possible actions specific to each of natural systems. We have a proud tradition the seven biological community types of leadership in adapting management described and assessed in this report. These techniques based on the cutting edge of are listed at the end of each of the seven knowledge of natural systems. The rapid biological community chapters that com- growth of new knowledge about ecosys- prise the bulk of this report. We call these tems demands that we change the way we “possible actions” because they are consis- view and resolve management problems. tent with ecosystem management but Many Department employees are, and have require more analysis and planning. How been, using ecological principles in formu- priorities are set within this list will be lating their management actions. We need based on ecoregion goals, staff workload, to build on our existing base of knowledge fiscal resources, public input and support, and experience. and legal authority. We will work with our This report, which attempts to bring customers and clients to set priorities and together current knowledge of biodiversity bring recommendations to the Natural and to stimulate thinking on the issue, is a Resources Board for consideration begin- step in this direction. It contains two types ning in the 1995-97 biennium. of recommendations. The first are broad strategic recommendations. These are This report proposes that the best way described generally below and in more to address biodiversity as a manage- detail at the end of the next chapter. We ment issue is to apply the principles recommend that the Department: of ecosystem management to Depart- ment planning and programs. Ecosys- Apply ecosystem management principles tem management is a system to assess, and practices to the Department’s conserve, protect, and restore the programs so that goals and priorities for composition, structure, and function biodiversity can be determined in the of ecosystems, to ensure their context of ecological, socio-economic, sustainability across a range of tempo- and institutional issues. ral and spatial scales, and to provide desired ecological conditions, eco- Build partnerships with other agencies, nomic products, and social benefits. local governments, tribes, the business community, scientists, and interest groups to accomplish common goals for A strategy for applying ecosystem ecosystem management, including management requires at least three impor- specific attention to biological diversity. tant building blocks: Use the ecosystem management decision Build partnerships with private land- model, as described in this report, to owners to accomplish common goals for think through alternatives and make ecosystem management, recognizing decisions. It is a model that requires us that the Department cannot accomplish to propose and evaluate alternative

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 7 EXECUTIVE SUMMARY

actions from their ecological, socio- Use logical steps to conserve biodiversity economic, and institutional (laws, rules, and retain future options, using the best policies) perspectives. This approach information we have now, while con- will help us frame issues in the context tinuously evaluating and improving our of their ecological, social, and economic approach as more information becomes consequences. In doing so, we will be in available. We must make and improve a better position to make decisions that decisions in the face of uncertainty. include human needs and values while Scientists have developed a method, preserving a wide range of options for known as “adaptive management,” to do future generations. this. Adaptive management is a formal, structured approach to dealing with Use ecoregions as the geographic basis uncertainty in natural resource manage- for developing consensus on regional ment, using the experience of manage- goals for program planning. Ecoregions ment as an ongoing, continually improv- are large areas of the state that exhibit ing process. This process will help us similar patterns in potential natural implement ecosystem management at a communities, soils, hydrologic condi- landscape scale, using a strong science tions, landforms, lithology, climate, base and a clear record of why we are natural processes, and resource or land- using particular management practices. use patterns. The ecoregion approach will enable us to set clear and measurable goals for protecting and managing biological communities.

8 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Should we hire employees with differing SOME CONTINUING POLICY QUESTIONS skills than those we now hire if we broaden our concern for biodiversity? Although we have identified key strategic and policy recommendations that What will be the role of surrogate we are ready to implement with our biological communities (e.g., switch- partners and customers, we are aware that grass-dominated grasslands instead of this report will continue to raise important multi-species prairies) or surrogate policy questions over time. These questions processes (e.g., clearcutting instead of affect the balance of interests among a wide burning a forest stand) in meeting our range of organizations and people. We have objectives for biological diversity? not identified all the implications, but we list a number of them below. They relate to To what extent should early 19th- issues of organization, budget, customers, century native plant communities be skills, and management. restored? At what cost? What is biologi- How will including biodiversity as a cally possible and what can be economi- criterion affect the balancing of multiple cally justified? views in DNR decision-making? How should the Department’s environ- How do our traditional customers mental quality programs integrate perceive their interests being repre- ecosystem management and biodiversity sented within ecosystem management concepts into their planning and permit- and attention to biodiversity? ting processes?

Are the present DNR budget structure and associated constraints ﬂexible enough to deal with ecosystem management and biodiversity issues?

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 9 BIODIVERSITY: ISSUES & IMPLICATIONS

pressures on our natural communities. Human population growth, coupled with land development patterns and high per- capita consumption of energy and natural CHAPTER 2 resources, leads to pressure on habitat from development, air and water pollution, and extraction of resources for energy and other uses. All of this can lead to loss of biological diversity. Biodiversity: As human populations grow and our needs and ability to use the environment increase, we will continue to alter ecological systems even though the absolute limits Issues and of ecological systems to absorb human activities are unknown. At the same time, we depend on these systems for clean air and water, food, shelter, and the raw Implications materials that support many of Wisconsin’s by James Addis, Betty Les, Anne Forbes, and Kristin Visser industries. In addition to these beneﬁts, plants have yielded life-saving drugs, and assisted by Robert Dumke, Paul Matthiae, Steven Miller, Dennis studies of animals have provided valuable Schenborn, Wendy Weisensel, and Darrell Zastrow insight into navigation, biochemistry, linguistics, and medicine. Conserving biodiversity will help sustain the ecological systems that we depend on. It will also preserve options for future decision- making. Biodiversity is complicated, occurring IMPORTANCE OF BIODIVERSITY at many different levels. For purposes of study and management, biological diversity iodiversity is a shortened form is usually grouped into four levels: genetic of the term biological diversity1— diversity, species diversity, community diversity, BIODIVERSITY: the spectrum of life forms and and ecosystem diversity (Fig. 1). ISSUES & the ecological processes that Genetic diversity consists of the spec- IMPLICATIONS support and sustain them. trum of genetic material carried by different Biodiversity supports the organisms. Genetic diversity within a integrity of the ecological systems upon population of a plant or animal species has which humans depend. These ecological the potential to change over time, allowing Biological systems (ecosystems) are self-sustaining species to adapt to environmental condi- diversity—or units, and to a certain extent they can tions and retain vigor. Although genetic biodiversity “for absorb disturbance without suffering loss of diversity may be expressed in visible short”—is the function. However, repeated or large-scale characteristics, such as color, size, and human disturbance inevitably changes shape, much is expressed in biochemical spectrum of life ecosystems and can threaten their viability. processes that are hidden from view. forms and the Humans have a profound and con- Individuals within a population carry a ecological tinuing impact on Wisconsin’s ecological variety of genes. If something happens to processes that systems. While some may think of tropical reduce the size or variety of the gene pool, support and rainforests as the only areas where ecosys- then that population’s genetic diversity is sustain them. tems are in danger, continuing human compromised. population growth here at home creates Species diversity results from the variety of species in a geographic area. It 1 Terms in italics are deﬁned in the glossary. includes not only the number of species in

10 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Genetic Diversity Figure 1 The variation in genetic Biological diversity composition of individuals occurs at four within and among species. interrelated levels, (e.g. variation within a adapted from Temple population of rabbits) (1991). Species Diversity The variety of different species found in an area. (e.g. the variety of species found in a prairie) Community and Ecosystem Diversity The variety of physical environments and biotic communities over a landscape. (e.g., the variety of forests, grasslands, wetlands and aquatic systems over a region)

the area but also their relative abundance change—change usually occurs, however, and spatial distribution. Species are the at a rate too slow for humans to note in our most familiar level of diversity because they brief lifetimes. Communities range in size can be classified and counted, and many, from less than an acre (e.g., shaded cliff though not all, are readily visible. Species community) to thousands of acres (e.g., include everything from soil fungi and mesic hardwood forest). The diversity of a insects to eagles and deer, from darters to given community is determined by the muskies, and from mosses and lichens to variety and type of species present, the hemlock and red pine. Every species has a intricacies of their interactions, and the age niche, or a role it plays in a natural commu- and stability of the community. The com- nity, defined by how individuals of a species munity diversity of a landscape is influ- carry out their activities, use resources, and enced by the number of communities occupy space. present, the degree of Understanding the difference among the niche of a single Conserving biodiversity will help sustain communities, and plant or animal the ecological systems that humans how the communities species requires in- are distributed. depend on and preserve a wide range of depth study as well An ecosystem is a as an understanding options for the future. dynamic complex of of the environment in plants, animals, and which the species microorganisms and lives and interacts. their associated non-living environmental A community is an assemblage of components interacting as an ecological species living together in a particular area at unit. An ecosystem takes the biotic commu- a particular time. Communities usually bear nity one step further to encompass interac- the name of their dominant plant species, tions with the abiotic environment, which for example, pine barrens, sedge meadows, includes moisture, temperature, oxygen, and cedar glades. However, the community sunlight, soil, and all the other non-living includes all of the plants living in associa- physical and chemical conditions. The tion with the dominant species plus all of biotic (living) and abiotic (nonliving) the animals present at a given time. Com- environment interact continuously. Often, munities are often perceived as static, but this interaction takes the form of complex they are actually in a constant state of processes that move gases, chemicals, and

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 11 BIODIVERSITY: ISSUES & IMPLICATIONS

minerals in endless cycles such as the three of North America’s six biotic prov- carbon cycle, water cycle, and nutrient inces—the eastern deciduous forest, the cycle. For example, a downed tree will, northern boreal forest, and the temperate through leaching and decomposition, grasslands—we have a wealth of species recycle its nutrients back to the ecosystem and natural communities. Curtis (1959) to be used by other living organisms. The delineated 21 major plant communities for canopy gap created when the tree was Wisconsin, plus 13 lesser communities The interactions downed will let in sunlight, altering condi- restricted to small areas. Approximately that connect tions on the forest floor and providing 1,800 native vascular plant species are opportunities for new plant species to known in Wisconsin, along with 657 microorganisms, become established. While all this is species of vertebrates. In addition, there are plants, and happening, the tree is providing shelter for thousands of species of nonvascular plants animals with the mice and salamanders, food for inverte- and invertebrates. DNR’s challenge is to nonliving brates, and substrate for plants. This tiny work with Wisconsin citizens to conserve environment are all ecosystem exists within a much larger this biological wealth. part of biological, forest ecosystem that might encompass physical, and thousands of square miles. In this larger system are hundreds of species of plants, HISTORICAL AND CURRENT chemical cycles hundreds of animal species, and probably PERSPECTIVES IN RESOURCE that have been thousands of species of microorganisms. MANAGEMENT occurring on Earth Ecosystems are constantly changing in for millions of response to short-term human impacts Throughout the history of natural years. such as timber harvest and naturally caused resources management, decisions have been perturbations such as fire or disease, along based in part on the personal values of with long-term influences such as climatic individuals. Today, each DNR employee has change. a personal history and set of values related Within ecosystems, the processes of to natural resource management. Consider, ecological succession—that is, the progres- for example, a group of managers standing sive changes in species composition, on a hillside looking out over an expanse of organic structure, and energy flows over land below. One person notes the low time—are also constantly at work. Large mounds and plains that indicate glacial ecosystems contain a mosaic of successional topography. Another’s eyes go straight to stages—a forest may have large areas of the creek meander- fully mature trees, but ing through the will also have open scene; this person areas with shrubs, Wisconsin is located where three of North wonders what fish patches of young trees America’s great natural borders join. Here, are present and if growing up after a East meets North and West to create a they get to any size. blowdown, and other wealth of biological diversity. It is DNR’s Another person in vegetative communi- challenge to work with Wisconsin citizens the group spots a ties within the larger to conserve this natural heritage. small plot of millet matrix of the mature growing on the forest. Ecosystems are otherwise fallow land in turn part of the and comments that larger landscape of adjacent and interacting it’s probably a wildlife food patch. Yet ecosystems. Surrounding lands can signifi- another scans the land with binoculars, cantly affect the character of an ecosystem; looking for wild flowers and signs of any therefore, ecosystems must be considered unusual habitat. Some think of this piece of within the context of the broader land- land as potentially something to manage for scape. a natural “product” such as grouse; others Wisconsin is blessed with great in the group think of it primarily as some- biodiversity. Located at the junction of thing to preserve or to restore to its original

12 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE natural condition. If they examined their DEVELOPMENT OF CONSERVATION ETHICS: personal feelings about the land, some 1850S TO 1950S would discover that they view themselves as part of the land. Others would discover Current environmental values and that, while they appreciate and respect the viewpoints are rooted in two different land, they view themselves as separate from schools of thought that developed in the it, as its manager or steward. All of these latter half of the 19th century. These are individuals are natural resource profession- often referred to as the preservation ethic als, and while they have many things in and the conservation ethic. common, they also have obvious differ- The preservation ethic, ﬁrst articu- ences. Most of these differences are due to lated by Ralph Waldo Emerson and Henry training, experiences, and values. A variety David Thoreau in the mid-1800s, and later of values about the environment have espoused by John Muir and others, focused prevailed during different periods of on the spiritual value of nature and viewed history. its preservation as a moral obligation. The

Each DNR employee has a personal relationship to natural resource management. As a group of managers stand on a hillside and look over an expanse of land below, each will see the same scene through the ﬁlters of their personal experiences, traditions, and values. If we come to understand the historical roots of our traditions and values in resource management, we may better understand the perspectives of others and work with diverse views to ﬁnd solutions.

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preservation ethic was a reaction to the ranger school and game warden school in large-scale urban, industrial, and agricul- 1911-1912. The Wisconsin Conservation tural development that seemed to Emerson, Commission and Conservation Department Thoreau, and Muir to be crowding out were created in 1915, pulling together natural environments. This ethic viewed boards and commissions covering parks, nature as having intrinsic value apart from fish, game, forests, and law enforcement. its utilitarian value. Thoreau’s statement These first managers focused their that “in wildness lies the preservation of the work on fish, wildlife, trees, and other world” sums up the preservation viewpoint. resources that had potential for regenera- Although the movement created a tremen- tion. Their job was to manage these highly dous awareness of natural beauty and utilized resources for the “greatest good,” to succeeded in preserving some of our benefit the greatest number of people over nation’s most scenic landscapes, it did little the long term. Conservationists did not to stem the exploitation of species. How- advocate preservation for its own sake. ever, this spiritual valuing of nature re- They believed that careful management mained a strong theme throughout the 20th could produce a sustained flow of benefits century and became entwined with the with an emphasis on valuable species such resource conservation ethic in many ways. as deer or pine trees. The motto of the It is the philosophical foundation for the conservation ethic became “wise use” or way that many managers and citizens relate “use without abuse.” Finely honed profes- to the environment today. In Wisconsin, the sions developed, each specializing in a preservation ethic sparked the establish- particular resource. As the professions We do not need to ment of Wisconsin’s first state park in 1900 became more scientific, they also became choose between and the identification of other areas of more quantitative. This increased technical preservation and natural beauty to be purchased in succeed- orientation plus the passage of time may conservation. ing years. In 1907 the State Park Board was have obscured the basic value lying at the Rather, they each established to oversee this task. And the core of these professions, but the resource nation’s first Natural Areas program began conservation ethic was continuing to exert have a role to play in Wisconsin in 1951. a profound influence. as we create an The prevailing source of values in Aldo Leopold’s land ethic tran- overall set of natural resource management for the past scended both the resource conservation principles for 100 years or so has been the resource ethic and the preservation ethic. The land managing conservation ethic. This ethic grew out of ethic, which Leopold articulated just before resources. the crisis created by the overexploitation of his death in 1948, focused on the nature that occurred during Euro-American interconnectedness of nature and the settlement. As fish and wildlife populations rightness or wrongness of human interac- were decimated and the forests reduced to tion with nature. Leopold’s simple but stumps, the public demanded government compelling statement that “the destruction intervention. As a result, the first conserva- of land, and the living things upon it, is tion agencies were formed. Although these wrong” sums up the basic premise of the first managers had little formal training in land ethic. The land ethic grew out of science or natural resources, the subject Leopold’s personal experience as a forester matter of their work gradually made its way and game manager practicing traditional into colleges and universities as the need techniques of range management, predator for trained managers increased and as the control, and user regulations. The land complexity of the work became evident. ethic does not oppose human use of nature In Wisconsin, the conservation ethic or scientific management of natural sys- was expressed in the establishment of tems; in fact, it assumes both. What matters departments, boards, and commissions to is how we go about our use and manage- protect and manage the state’s resources. In ment of nature. According to the land ethic, 1903, a State Forestry Department was it is in the self-interest of humans to treat established, followed by the first forest the land well since we are part of nature

14 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE and our well-being depends upon it. THE GROWTH OF ENVIRONMENTALISM: THE The notion that we Leopold’s work led to an awareness that 1960S AND 1970S could load the management actions have far-reaching and earth with often unpredicted consequences and that For natural resource professionals, the the natural world is far more complex and environmental movement of the 1960s chemicals and interrelated than scientists had previously added new values to the preservation ethic, wastes with no realized. This thinking was gradually resource conservation ethic, and land ethic. consequences integrated into the developing science of This new set of values has been termed the gave way to a ecology and called for a new consciousness environmental protection ethic. This more sober, regarding our relationship to the land. ethic views the environment as a set of realistic view of physical systems that must be maintained limits and harm THE T URNING POINT: SILENT SPRING in a healthy, functional state. Regulating pollutants going into systems, monitoring and paved the way The decades since the emergence of movement of pollutants within systems, for the Leopold’s land ethic have seen a variety of and predicting their impact are prime environmental landmark events concerning the environ- concerns for natural resource managers protection ethic. ment and natural resources. Foremost working in environmental protection. among these was the publication in 1962 of Concern for species was not left out of this Rachel Carson’s book Silent Spring. This period; in fact, the reasons for the decline book raised the specter of a soundless of species was now seen in the larger spring—a spring without insects, inverte- context of environmental damage. The brates, frogs, birds, and other animals—and Endangered Species Act of 1973 provided a brought home the meaning of the effects of legal means to conserve the ecosystems pesticides in complex food webs. Its which support endangered species and publication marked an end to a period of threatened species. This act also gave innocence for the expression to American public. Leopold’s adage that The notion that we “The land ethic simply enlarges the “the first rule of could load the earth boundaries of the community to include intelligent tinkering is with chemicals and to save all the parts.” the soils, waters, plants, animals, or wastes with no Multiple values, consequences gave collectively: the land.” (Aldo Leopold, The often appearing as way to a more sober, Sand County Almanac) incompatible, were realistic view of simultaneously limits and harm. As developing among more evidence of citizens. For example, one value held that damage surfaced, public pressure to we must protect the environment for future regulate pollution increased. This new generations. Another held that we must public concern focused on the environment protect the environment primarily for as a whole rather than the species orienta- present economic benefits. Individuals with tion that had dominated decades earlier. In this latter viewpoint often saw human 1967 Wisconsin became the first state to qualities as separate and “at the top of” ban the pesticide DDT. At the national nature, giving people special rights and level, comprehensive environmental responsibilities. An opposing viewpoint legislation (Environmental Policy Act, saw nature as having value in its own right Clean Water Act, Clean Air Act) was put in and violation of nature as immoral; humans place. This legislation added a new dimen- were seen as just one part of nature, sion to natural resource management and perhaps a small and inconsequential part in led to the creation of new environmental the total scheme of things. The result was protection professions. lively and intense public debate, but no clear or unified vision developed to guide policy decisions. The first Earth Day, led by

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Wisconsin U.S. Senator Gaylord Nelson, flected these differences. For example, the was held in 1970. This event drew many Division of Resource Management was In 1967 the people into the debate and widened the organized into separate bureaus for man- Executive Branch base of support for environmental protec- agement of fisheries, wildlife, endangered Reorganization Act tion. resources, forestry, and parks, with research During this period, single-species and property management functions brought together management was still the rule in resource providing support. The Division for Envi- closely related management, but the strong Aldo Leopold ronmental Quality was organized around traditional tradition in Wisconsin was also being felt. the major abiotic components of the conservation tasks For example, wildlife and fisheries manag- environment (air, water, land), emphasizing and newly ers who were aware of the value of wet- their management and regulation, mostly emerging lands for a wide spectrum of species (not through permit control. environmental just target species like ducks or northern pike that the public normally associated EMERGENCE OF CONCERN FOR BIOLOGICAL protection with wetlands) broadened projects to DIVERSITY: 1980S-PRESENT responsibilities to benefit the system as a whole. Acquisition create the of land adjoining trout streams in the The next major step in the evolution Wisconsin Central Sands counties not only protected of natural resource management occurred Department of the stream habitat and allowed fishing in the 1980s, when public concern for loss of natural spaces and rapidly increasing Natural Resources. access but also protected the bottomland vegetation adjacent to the stream. Land scientific knowledge about the acquisition for watershed protection also interconnectedness of all the pieces of began in this era. ecosystems merged to produce both public Although Wisconsin had regulated and scientific interest in managing re- pollution as early as 1927, when the State sources with the goal of conserving biologi- Committee on Water Pollution was created, cal diversity. This increased concern for the 1960s marked the beginning of a biological diversity cannot be attributed to tremendous increase in programs devoted any given person or group. Indeed, the to the environment. Much of this was due thoughts of many people from around the to the formation of the Wisconsin Depart- world—scientists, managers, philosophers, ment of Natural Resources, which occurred and the public—contributed to its develop- in 1967 when the Executive Branch Reor- ment. Scientists have come to understand ganization Act, developed under the that some concepts—especially the idea guidance of the that ecosystems reach a “steady state” or Kellett Commission, “climax” condition— became law. This law do not provide an brought together The conservation of biological diversity accurate picture. This closely related became the next major step in the correction to an traditional conserva- evolution of natural resource established assumption tasks and newly management. tion has led to emerging environ- questioning some mental protection established manage- responsibilities. It merged the Department ment principles and activities. These of Resource Development, air pollution changing concepts of management place functions of the State Health Board, and the new and challenging demands on DNR Wisconsin Conservation Department to employees and the agency as a whole. form a single state agency. Throughout 1993 and 1994, discus- The merging of these programs also sions with DNR staff, county forest admin- meant a merging of administrators and istrators, leaders of groups representing managers with different historical influ- business, environmentalists, hunting and ences, values, and approaches to problem- fishing organizations, academics, and solving. The new DNR organization re- members of the public have shown us that

16 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Wisconsin residents do indeed hold a wide DNR’S ROLE IN CONSERVING BIODIVERSITY Wisconsin citizens range of opinions about biodiversity. Most hold a wide range people we spoke with do agree on a By creating the Department of Natural of opinions about definition of biodiversity similar to the one Resources, the Wisconsin Legislature used in this report. Most feel that recognized the need for an integrated the conservation of biodiversity is a serious issue, brought to approach to protecting, conserving, and biodiversity. Most public attention by a combination of both enhancing Wisconsin’s environment and people who shared natural resources. The legislature further advances in scientific knowledge and their thinking with recognized that the needs of traditional public concern for loss of wild and natural us agreed on a places. Some believe that changes in conservation programs and environmental basic definition of management must be made, but that we programs were closely interrelated and that have time to think through changes and forming a single agency would provide biodiversity similar make incremental changes. Others feel that better management and greater public to the one in this resource managers have not adequately benefit. Thus, the Department’s mission report. This considered biodiversity in their decisions, statement reflects this holistic approach: common ground and that there is an urgent need to revise Protect and enhance our Natural Re- will be important management practices and to set aside large sources—our air, land and water; our as we seek to biodiversity reserves to protect ecosystems. wildlife, fish and forests. integrate ecology, Still others are concerned that biodiversity economics, and is too complex a concept on which to base Provide a clean environment and a full management actions. They may want to range of outdoor opportunities. human values in manage to conserve biodiversity but are resource unsure what actually needs to happen “on Insure the right of all Wisconsin citizens management the ground.” Some people we’ve spoken to use and enjoy these resources in their policy and with feel that concern for biodiversity is work and leisure. practice. just a fad, others think of it as the newest environmental buzzword, and some said And in cooperation with all our citizens that resource management professionals to consider the future and those who have been managing for biodiversity all will follow us. along. Those with whom we spoke do agree Managing to conserve biological that ignoring issues until inflexible posi- diversity helps the Department carry out tions have been staked out is not the way to this mission. Although biodiversity was not handle biodiversity issues in Wisconsin. a common term until recently, the Conser- Factual data analysis together with open- vation Act of the early 1920s provides the minded dialogue among citizens of diverse direction required for our present-day perspectives will be required to develop a response to biodiversity as a management consensus for policies that integrate ecol- issue. Chapter 23.09 of the Wisconsin ogy, economics, and human values. Statutes states that “the purpose of this Biodiversity and economic health must not (conservation) section is to provide an be seen as conflicting; they must be inte- adequate and flexible system for the grated into rational resource policies that protection, development, and use of forests, will enhance the lives of present and future fish and game, lakes, streams, plant life, generations. The practice of ecosystem flowers, and other outdoor resources in this management, as described later in this state.” The balance of protection, develop- chapter, will play an important role in ment, and use of our natural resources bringing this integration about. demands that we conserve the long-term functional health of ecosystems, including their biological diversity.

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the 17th century, they found a much ECOLOGICAL ISSUES smaller human population than had existed two hundred years before. The result was Plants, animals, and humans colo- that much of the area may have become nized Wisconsin as the glaciers receded more “natural” due to less human manage- 10,000-12,000 years ago. The earliest ment and use. archeological evidence of human presence The first Euro-American settlers, in Wisconsin dates from 11,000 years ago. arriving in Wisconsin in the 1830s and Thus, the biotic communities that took 1840s, found a landscape characterized by hold in Wisconsin after the glaciers receded extensive forests, grasslands, wetlands, and were influenced by human activity from the a variety of other biotic communities. beginning. The size of the human popula- While the species of plants and animals the tion in Wisconsin in the millennia before Europeans found here had adapted to European contact is a subject of specula- Wisconsin’s soils and climate over thou- tion. At one time the settlement of Cahokia, sands of years, they had done so in the in southern Illinois, supported a population presence of humans who were continually of perhaps 30,000 individuals. We can affecting the landscape to the extent assume that pre-Columbian Wisconsin also allowed by their population size and their had a large human population, although technology. Europeans brought technolo- there is no evidence that communities the gies of the industrial age that began more size of Cahokia were located here. intensive manipulation of the environment. The native populations managed the They also introduced, both purposefully landscape to produce food, fiber, fuel, and and accidentally, many non-native plants other needs. We do not know how much and animals to compete with the native land might have been in agriculture, but species, often resulting in broad changes in there is archeological evidence of irrigation, ecosystem composition, structure, and raised beds, and other intensive farming function. practices. Farming and the use of fire for Today, Wisconsin’s landscape reflects a agricultural clearing and for managing high degree of human use. It is a mosaic of animal and plant populations were possibly urban areas (cities, towns, suburbs), the most significant factors affecting natural production areas (farms, mines, industries, succession of plant communities in Wis- commercial forests), multiple-use areas consin. In northern Wisconsin, native (parks, lakes, public forests), and protected populations hunted and fished intensively, natural areas (conservancy, wilderness). and impacted the northern forests through This patchwork bears little resemblance to gathering firewood, creating clearings for the landscape the native populations knew, settlements, favoring plants useful for or to the one the first European explorers medicine and food through cultivation and saw. management, and using forest materials for Managing Wisconsin’s natural re- tools and shelter. sources to conserve biodiversity requires When Europeans landed in the New that we understand how today’s patterns of World, this picture changed dramatically. land and resource use were created from Native populations lacked immunity from these earlier landscapes and how human such diseases as smallpox, influenza, activities and natural processes continue to measles, venereal diseases, and the com- produce those patterns (Fig. 2). The mon cold. Beginning in 1492, disease activities and processes of particular spread along trade routes even to tribes that concern in relation to biodiversity can be had no direct contact with Europeans. grouped into three major categories: Throughout North, Central, and South ecological simplification, fragmentation, and America, native populations declined environmental pollution. While simplification dramatically due to disease epidemics. and fragmentation result from both natural When Europeans arrived in Wisconsin in

18 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Future Pattern of Cities, Land Use Towns Parks, Suburbs Lakes, Wetlands, Forests Farms, Corridors, Industries Riverways, Figure 2 Natural Visualization of current Presettlement Areas and potential future Landscape land use patterns superimposed on the presettlement landscape. and human-caused events, human actions Although this report focuses on tend to alter the ecosystem at an increased natural vegetation and natural communi- rate and at a greater magnitude than natural ties, we also need to consider urban and events do. Certain species, communities, agrarian systems as we continue to move and ecosystems are not doing well in forward to ecosystem management, because Wisconsin because of these human-caused humans are an integral part of the environ- changes. In contrast, populations of gener- ment. Our goal is sustainable ecosystems— alist species that are well adapted to the whether highly modified by humans or simplified or fragmented environment are largely natural—that maintain ecological flourishing. Environmental pollution, which composition, structure, and function and is almost entirely a maintain genetic, result of human species, community, activity, also poses Our goal is sustainable ecosystems in the and ecosystem formidable threats to context of today’s landscape. Knowledge diversity across all biodiversity. Al- of the presettlement vegetation will help land uses. though the effects of For natural us recognize the ecological potential of an pollution on ecosys- communities, early tems are often similar area and answer questions about how, 19th century vegeta- to those caused by how much, and where biodiversity can be tion can be used as a simplification and restored. guide for restoration fragmentation, the that provides valu- causes and mecha- able insights into the nisms of pollution are very different. ecological potential of an area. Although Understanding simplification, fragmenta- recreating this former state would be an tion, and environmental pollution will help unrealistic goal given current conditions us understand the ecological trade-offs and (except in some natural reserves), it is consequences of our decisions and will important to understand what was here in help define the institutional, social, and the era prior to Euro-American settlement. economic choices that will be made in the This period represents a time when scien- future to guide DNR’s activities in relation tific biological observation began, when to biodiversity. native plant communities were less dis-

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turbed by management activities and fairly What follows is an overview of undisturbed by the many species of non- ecological simpliﬁcation, fragmentation, native plants and animals. This information and environmental pollution, with an can guide us as to what elements of emphasis on explaining what these con- biodiversity can be protected and restored, cepts mean and how they impact in what levels of abundance, and in what biodiversity. The concept of scale provides a geographic areas, helping us reach our goal foundation for understanding how to deal of sustainable ecosystems in the context of with these issues (see inset). today’s landscape.

SCALE Scale is the relative amount or For ecological purposes, the degree of something, often expressed in amount of detail with which an ecosys- terms of a progressive classification as to tem can be described for management size, complexity, or importance. In planning is determined by the spatial management of natural resources, scale and temporal scales. Due to time and is often used to describe the scope of a resource constraints, we are often able to management action—whether site- provide more detail at smaller scales specific, local, regional, or statewide in than at larger scales. We often speak of space, and annual, seasonal, or succes- this situation using the term resolution, sional in time—and the degree to which i.e., as having a high degree of resolution the management action alters the at small scales and a low degree of existing plant and animal communities. resolution at large scales. For example, Thus, when the concept of scale is in an endangered plant inventory of a applied to ecosystems, it has both spatial very small plot, we may be able to and temporal meanings. Spatial scale is thoroughly sample the plot inch-by- used to describe the geographic size of a inch. An inventory of a large area would community or ecosystem (Fig. 3). This be done at a lower degree of resolution, size can range from a microsite such as perhaps by running transects at intervals the underside of a leaf on the forest across the area. The former sampling floor, to the entire forest, to the larger approach gives us a lot of information landscape. The biosphere, including about a very small area, and the latter earth, its enveloping air mass, and all its gives us less detail but includes a wider biota can be thought of as the largest geographic area and a larger amount of scale from a biological point of view. total information on plant distribution. Temporal scale describes the time re- The desired spatial scale for overall quired to complete a life history event or ecosystem management planning is the an ecological process, such as the a landscape. A landscape is an area com- series of successional stages (Fig. 4). For posed of interacting ecosystems that are life history events such as life cycles, related due to underlying geology, temporal scale can vary from a few hours landforms, soils, climate, biota, and for certain microbes and insects to human influences. Broad management thousands of years for ecosystems. goals will be set at this scale and will Ecological processes can vary from a few relate to relatively large geographic areas, seconds for individual biochemical using the information collected with a reactions to decades for forest regenera- low degree of resolution, or less detail, tion. When tied to geologic changes, as described above. Management of temporal scale reaches millions of years. specific sites within the broad landscape

20 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE will occur based on goals set at the would include an analysis of the ecologi- landscape scale. Information with a high cal conditions of all the streams and degree of resolution will be collected at watersheds on the Wisconsin shoreline specific sites as needed to check the of Lake Michigan and/or Superior. The accuracy of goals set on the landscape analysis would indicate how the overall scale or to fine-tune management plans management plans for these streams for specific sites. address statewide issues of biodiversity Landscape-scale management as well as other important related issues encourages us to approach problems such as recreation and water quality. and projects using the broadest scale Biodiversity is maintained by the with ecological meaning. Thus, the presence of an array of communities and geographic area or region included in species occurring within ecosystems any particular analysis will be deter- which are intact and sustainable, that is, mined by our knowledge of the breadth they usually contain a wide range of of the interconnections among the biotic species and natural processes. The communities involved. For example, a appropriate scale for management must proposal to create a new Natural Area in be considered and deliberated along Wisconsin for the protection of with other considerations if biodiversity biodiversity would include a series of is to be preserved or enhanced. If we are considerations—among these are the not aware of the concept of scale in size and quality of adjacent buffer areas planning a proposed action or do not needed to protect ecological integrity on understand the implications of our the site; the relative importance of the choice, we run the risk of developing site to biodiversity within a statewide inappropriate plans and prescriptions. view of community and ecosystem Worse, we can unknowingly change the status; and concerns such as the trans- community or ecosystem involved. Figure 3 port of pollutants or the condition of These choices are complex, for decisions Examples of spatial migratory bird habitats on continental or that favor increasing diversity at a given scales can be observed with the inter-continental scales. Or, a proposal scale may decrease diversity at other “nesting” of small to acquire land to support an anadro- scales. areas, such as a local mous sport fishery on the Great Lakes site, within progressively larger geographic units.

Local Site Watershed Landscape Ecoregion

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N. flying squirrel

Cerulean warbler

Four-toed salamander

Rose-breasted grosbeak

Indigo bunting

Deer mouse

Copes gray tree frog

Eastern cottontail

Gray catbird

Massasauga rattlesnake

Meadow vole

Savannah sparrow

Six-lined race runner

White-tailed deer

Grasses and Forbs Shrubs and Saplings Young Forest Mature Forest Old Growth

Figure 4 ECOLOGICAL SIMPLIFICATION COMPOSITION Examples of temporal Ecological simplification means that scale can be observed Composition refers to the fundamen- with the succession of the interrelationships between organisms tal elements of natural systems—the a southern Wisconsin and their environments are reduced in specific organisms or groups of organisms grassland to a forest. number and complexity. Simplification can The composition of that a unit area or geographic area contains. be caused by habitat loss, non-native plant and animal At the statewide level it includes ecosys- communities change species encroachment, air and water tems, communities, species, and their along with the pollution, and many other factors. Al- landscape. Adapted genetic composition. Thus, an ecological though the effects of simplification are with permission from system simplified in terms of composition material produced by complicated and often subtle, they are often might have reduced numbers of species the Minnesota discussed in terms of their impact on the Department of Natural present or a limited gene pool for a rem- three major attributes of ecosystems: Resources. nant or isolated species. composition, structure, and function (Fig. 5). The most radical impacts on composition occur when there is total destruction of the biotic, abiotic, spatial, or temporal needs of species. The conversion of native

22 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE prairies, savannas, wetlands, and southern Thus, the presence of non-native forests to agriculture and urban develop- species within terrestrial and aquatic ment since Euro-American settlement are communities and ecosystems often leads to among the most conspicuous examples. displacement of native species and change The introduction of exotic species in ecosystem function. Displacement is provides another usually not one for example. Purple one; one exotic species loosestrife, known to When ecosystems are simplified, we may can displace many exist in restricted areas observe a reduction in species numbers, native species. Because of Wisconsin’s wilds their gene pools, the physical structure of exotics are generally for over 40 years, has their habitats, or the complexity of life- introduced without spread across three- sustaining processes such as food webs consideration for quarters of the state in or water cycles. natural biological and ten years, seriously ecological controls, simplifying many once they “escape into wetlands. Garlic the wild” they some- mustard has invaded southern Wisconsin times prove very successful in competing forests, displacing native forbs and depen- against native species. Today, an estimated dent fauna. Common buckthorn and 22% of Wisconsin’s 2,300 vascular plant Japanese honeysuckle have invaded south- species are non-native species. ern mesic and dry mesic forests, while glossy buckthorn has invaded wet forests and bogs. All three have displaced forbs STRUCTURE and shrubs, and in bogs even established Structure refers to the pattern or trees are being lost to competition. In some physical organization of an area; it is used southern Wisconsin oak forests, buckthorn to define physical appearance both verti- and honeysuckle encroachment has begun cally and horizontally. Vertical stratification to significantly reduce oak regeneration. is readily visible in a mesic hardwood Similarly, the rusty crayfish, introduced as forest, where one group of species occupies sport-fishing bait, has spread to a large the canopy layer, another group the number of inland lakes, displacing native subcanopy, another the sapling layer, and Figure 5 crayfish and disrupting entire plant com- so forth, down through shrubs, tall herbs, The three major munities. The result has been an adverse short herbs, and ground cover (surface) attributes of ecosys- impact on other fauna such as fish, inverte- plants. Horizontal variation occurs across tems: composition, brates, and zooplankton that are dependent the length and breadth of any community structure, and function. upon the aquatic macrophytes consumed Ecological simplifica- or, at a larger scale, across sub-regional and tion occurs in relation by the rusty crayfish. regional landscapes. Canopy gaps, forest to all three.

Litterfall

Plants Detritus

Soil

Composition Structure Function The make-up of an ecological unit, including The pattern or physical organization of an area. The roles that the living and nonliving the speciﬁc organisms or groups of organisms It has both vertical and horizontal components. components of ecosystems fulﬁll in driving the in a particular area. processes that sustain ecosystems.

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openings, seasonal ponds, savanna alternat- tem or landscape scale, vegetation controls ing with prairie, riffles alternating with the community environment, is the primary slow water, and backwaters adjoining main source of energy for other organisms within streams are all examples of horizontal the community, and is the principal source variation. The location of vegetation in a of minerals and chemical compounds horizontal plane is related to the slope, necessary to sustain animal life. Animals are exposure, soil, proximity to other plants of the main consumers, with primary con- the same or different species, and dispersal sumers eating plants and secondary con- A simplified mechanisms. Vegetative debris—fallen trees sumers eating other animals. Still other ecosystem has and limbs, standing dead trees, and leaf plants and animals along with fungi and little of the and forb litter—is also part of a bacteria perform essential functions as structural community’s horizontal and vertical struc- decomposers and transformers of waste ture. products and detritus, converting dead complexity that Animals follow a pattern of vertical material back into elements essential to creates diverse and horizontal organization similar to plant growth. Each individual plant and habitat plants. For example, birds and inverte- animal has a functional role in the support opportunities as a brates that occupy the canopy of a forest of other species and the community as a basis for community differ from those that occupy whole. ecosystem the midlevel, many of which differ from Increased diversity and functional those residing on the forest floor. Similarly, complexity generally provide resilience to function. the fauna associated with a lake’s emergent ecosystems. Conversely, for a given ecosys- vegetation differ from the animals associ- tem, reduced biological diversity may result ated with the floating or submerged vegeta- in less resilience. In less biologically diverse tion. systems minor changes in energy flow or A simplified ecosystem has little of the population structure produce major structural complexity that creates diverse changes in energy transfer and populations. habitat opportunities as a basis for ecosys- Unpredictable and chaotic changes may tem function. For example, a forested area occur. A community will cease to be part of being managed on short rotation for even- a viable ecosystem if there is significant age, single-species trees is a structurally functional loss. The Lake Winnebago simplified system. Because the tree species system provides a good example. An are all of similar age and are cut before increase in water level in the system and other layers of vegetation can become well other factors led to a severe reduction in established, there is little vertical or hori- aquatic plant populations beginning in the zontal layering. Animal species inhabiting late 1800s. Wind and wave action across the area would likewise reflect lower these large expanses of water prevented diversity than would be found in a forest macrophyte reestablishment by increasing with trees of various ages. turbidity, eroding the shoreline, and uprooting plants. Populations of invertebrates, fish, and waterfowl have fluctuated FUNCTION through the years with a general trend Function refers to the roles biotic and toward decreasing numbers. Numerous abiotic components of ecosystems (e.g., attempts to manage the water level have producers, consumers, digestors, trans- met with limited success due to the great formers, water, minerals, and micro- functional losses to the system. A system- climates) fulfill in driving the ecological wide approach is now being taken through processes (e.g., water and carbon cycles, DNR’s Lake Winnebago Comprehensive mineral and nutrient cycles) that sustain Management Plan and offers a much better the ecosystem. Every naturally occurring chance for improvement. organism within an ecosystem has one or more roles in sustaining the dynamics of that ecosystem. For example, on an ecosys-

24 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE FRAGMENTATION gradually blend one into another. This “patchiness” may often protect plants from The natural variation of biological catastrophes such as disease, insect out- communities across a landscape, often breaks, and fire. referred to as “natural patchiness,” has Natural landscapes gradually merge always been a normal part of the environ- one habitat type with another or leave ment. At the time of Euro-American corridors or ways for animals to move and settlement, the natural landscape of Wis- still stay in their preferred habitat. Frag- consin was broken up into wetlands, mentation is the breaking up of large and prairies, forests, lakes, and streams, all continuous ecosystems, communities, and occurring in numerous patches of varying habitats into smaller areas surrounded by sizes. Some species, such as prairie chick- altered or disturbed land or aquatic sub- ens, thrived only on very large patches of strate. Fragmentation eliminates corridors suitable habitat. Others were more success- and causes the abrupt meeting of different ful at the interface edge between plant habitat types. In Wisconsin human activity communities and has greatly frag- took advantage of two mented the land- or more habitat In order to observe fragmentation of scape, producing types—for example, biological communities or ecosystems, changes that are the white-tailed deer we look at the pattern of fragments on different from natural which uses forest, the landscape, their sizes and proximity landscape heteroge- brushland, and neity or patchiness. to one another, and the types of edge prairie edges. For example, during Figure 6 Many animal that define them. Euro-American species need a high Changes in a wooded settlement the area of Cadiz degree of “patchiness” northern forests were Township, Green because their life requirements are met by logged and many areas were burned, County, Wisconsin, using different habitats at different times. during the period of leaving only scattered “islands” of forest Euro-American Others, such as grassland birds and interior remaining. These disturbances in the north settlement. The forest songbird species, are favored by occurred within a 50 year time period. shaded areas relatively large, continuous habitats of represent land After that time, the land-use pattern remaining in, or similar vegetation. More subtle differences remained “undeveloped” in character and reverting to, forest. in soil, microclimate, moisture level, slope, the land progressively grew back to forest. This fragmented and aspect permit plants to thrive in one As agriculture and urbanization grew in landscape is likely to area and not another. For example, the exhibit effects from southern Wisconsin, the southern forests, changes in the amount northern forest ecosystem includes numer- prairies, and wetlands were broken into of edge, reduced size ous communities representing a range of increasingly smaller and more isolated of fragments, and successional stages as well as natural isolation of fragments. fragments that remain today. Roads, sewer, From Curtis (1956) patches of oak, aspen, balsam fir, and tag and utility corridors; dams; residential, with permission of the alder, which under natural conditions commercial, and industrial development; University of Chicago Press.

1831 1882 1902 1950

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and land clearing and conversion continue In the north, communities and to contribute to fragmentation in both the ecosystems historically dependent on fire, north and the south. Some traditional e.g., the oak and pine barrens, have been resource management practices have also fragmented and diminished in size by forest contributed to fragmentation. encroachment. Populations of species such Many species continue to do well in as sharp-tailed grouse, which depend on these artificially segmented landscapes. these open areas, continue to decline across Some, such as white-tailed deer, are even their historic range due to loss of barrens more successful than they were historically. and other open habitat. However, many species of plants and animals lose ground as a result of increased fragmentation. Fragmentation generally ISOLATION results in three types of change: fragment Isolation of habitat patches occurs as size reduction, increased isolation of the landscape becomes progressively fragments, and increased edge-to-interior fragmented. Areas of the same type become ratios (Fig. 6). isolated, not only by distance but by hostile intervening environment, putting plants and animals without adaptations for long- SIZE EFFECTS range movement at a severe disadvantage. As the size of a particular fragment The inability of a species to move between becomes smaller and smaller, more and habitat patches leads to loss of genetic more species disappear from it. For ex- viability and diversity and can ultimately ample, very large blocks of prairie histori- lead to elimination of that species within cally contained more than 400 vascular that fragmented habitat. As the intervening plant species and a multitude of animal environment becomes increasingly hostile species including microfauna, insects, even the more mobile species have their herptiles, birds, and mammals. As the movement between habitat islands prairies were broken up, large ungulates thwarted. Today, many community and such as bison and large predators such as ecosystem fragments are so far apart and so wolves quickly disappeared. As fragmenta- reduced in size that many animals fail to tion continued, some plant species disap- maintain populations. The communities peared and many others became rare. Now become closed systems subject to cata- many are in serious decline. strophic change from events such as The size effects of fragmentation are disease, drought, wind storm, or floods. particularly noticeable in the southern and Some of the most prominent examples west-central two-thirds of the state. Here of isolation resulting from habitat or major ecosystems (grasslands, savannas, community fragmentation can be found in and southern forest) were reduced in size what is left of the prairie ecosystem. This and severely fragmented by agriculture and ecosystem is now severely reduced to small by urban and rural residential develop- isolated fragments scattered about in a “sea” ment. Many remnants of these once- of agricultural and other lands inhospitable expansive ecosystems are represented by to many of the prairie species, especially community fragments too small to support invertebrates and plants. Recent survey viable populations of many species. For work in Illinois suggests that at least 15% example, most species of the grassland bird of what is left of prairie insects are now guild are in decline, as are reptiles and restricted to prairie remnants. These amphibians. The ornate box turtle requires remnants range in size from two to 1,000 a minimum of 250 acres of prime sand acres; most are in the 2-40 acres in size. As barrens habitat to sustain a viable popula- much as 30%-40% of our prairie plants tion, but only three or four areas of this may also be remnant-restricted. Without habitat size and quality remain. connecting corridors or “stepping-stones” of close enough proximity and large

26 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE enough size, many of these species popula- forests, many exotics gain entry to interiors tions will remain permanently isolated and by first getting established in the distur- thus subject to inbreeding, continued bance zone associated with human-caused decline in numbers, and eventual elimina- disruption. Interior edge, which is more tion from that patch of habitat. common in the north, is caused by logging, Aquatic ecosystems are also subject to agriculture, blowdowns from wind storms, isolation. All of Wisconsin’s large rivers, fire, and residential and commercial most of its medium-sized rivers, and many development, and takes on the same form smaller streams have been fragmented by and effect as exterior edge. Area- and edge- dams. Fragmentation causes streams to sensitive interior species are especially become a series of modified ecosystems vulnerable to interior edge conditions. which no longer represent the native Corridors for roads, power transmis- ecosystem in structure, function, and sion lines, and pipelines create linear edge composition. Lake shores have also been throughout the north, while in the south, fragmented by sand blankets and vegeta- these corridors sometimes bisect woodlots, tion removal associated with shoreline wetlands, and grasslands. These corridors developments. Dams prevent fish from are havens for edge species and allow for reaching upstream spawning grounds, but penetration of species into otherwise there are other, more subtle effects of dams. continuous communities and ecosystems. For example, damming frequently isolates mollusks from the fish that host their larval ENVIRONMENTAL POLLUTION stages; mollusks unable to complete their life cycle because of this isolation are Environmental pollution is the eventually eliminated from the stream. For human-induced addition of many types of other species, populations are diminished substances to air, land, and water in when individuals succumb to siltation and quantities and/or at rates that harm organ- other effects of damming. isms, habitats, communities, ecosystems, or human health. Examples are nutrients, Pollution may heavy metals, organic compounds, and change the EDGE EFFECTS sediments. Pollution may change the physical, chemical, physical, chemical, or biological character- Edge effects occur near the interface of or biological istics of air, water, or land, thus affecting two or more different habitat types. Edge the health, survival, or activities of living characteristics of effects are beneficial for many plant and organisms in a variety of detrimental ways, air, water, or animal species, since edge allows them to including impacts on genetic, species, land—affecting the take advantage of two or more habitat types community, and ecosystem diversity. for their survival. However, many other health, survival, or Any Department policies relating to species are negatively affected by too much activities of biological diversity need to consider the edge. The concentration of many species organisms and effects of pollution and the efforts required near edges causes increased competition, contributing to the to manage resources that have been ad- predation, and parasitism. For example, versely affected by pollution. The following forces that simplify waterfowl nesting near the edges of grassy examples illustrate some of these effects as and fragment fields experience high nest predation, as do they relate to water, air, and land resources. communities and some songbirds nesting near forest-field edges. Or, some plants may disappear from ecosystems. previously suitable interior habitat when a ADVERSE IMPACTS TO SURFACE new edge changes the micro-climate. As AND GROUND WATER SYSTEMS community or ecosystem islands get smaller or more disturbed, they become Poorly managed construction sites less and less viable for interior plants and and bare fields allow soil to wash off the animals. In effect they become all edge. land in runoff. This sediment can smother Encroachment of exotic species is gravel riffles in a stream, destroying the closely associated with edge dynamics. In habitat for aquatic invertebrates and

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 27 BIODIVERSITY: ISSUES & IMPLICATIONS

spawning sites for some fish. Excessive areas. In fact, some species of trees simply organic waste flowing into a lake or stream cannot grow in areas with severe air pollu- uses up dissolved oxygen as it decays, tion. For example, high levels of ozone in which can kill aquatic life either through the air of southeastern Wisconsin are direct toxicity, destruction of habitat or known to limit the growth of several food supplies, or by elimination of the species of trees. Even at moderate levels of dissolved oxygen needed by aquatic plants air pollution, some individuals within a and animals. Phosphorus and other nutri- population are genetically more sensitive to ents flowing off fertilized lawns and crop- air pollution and will be eliminated from land into waterways may allow growth of the population, resulting in simplification excess algae and other aquatic plants. of the gene pool. This is a good example of When large amounts of this vegetation die, reduction in genetic diversity. Neither the decomposing bacteria use up dissolved short-term nor the long-term implications oxygen, killing fish and other aquatic life. of this simplification is understood at this Direct discharge of industrial effluent time. and sewage, which contain organic resi- Acid deposition from air-carried dues, chemicals, and sediments, can limit pollutants may change water chemistry in dissolved oxygen in the receiving water, some lakes, which in turn can change the otherwise change water chemistry, alter diversity and abundance of aquatic organ- habitat, and kill organisms. Some chemicals isms and communities. Acid deposition present in industrial and municipal efflu- may change the pH of a waterbody, which ent, such as dioxin, have been shown to can encourage the release of mercury cause diseases, suppress the immune already present in sediments or substrates. systems of a variety of species, harm This process may enhance bioaccumulation reproductive capability, and produce of mercury, which accumulates in organ- genetic defects in offspring. The tempera- isms at the top of the aquatic food chain, ture of wastewater may also change normal affecting their health, survival, or offspring. aquatic temperature gradients and disrupt There is limited but increasing the life cycles of some aquatic plants and evidence that mammals, birds, and other animals. organisms are also adversely affected by Pollution of ground water, caused by inhaling airborne pollutants such as events such as gasoline leaking from pesticides, heavy metals, and organic underground storage tanks, nutrients chemicals. If not directly toxic within an leaching from inadequate septic systems, or adult organism’s life span, these substances pesticides washing off farm fields, poses a may be toxic to an organism’s progeny by direct human health hazard when it reaches causing birth defects, depressing the household water supplies. Contaminated immune system, or changing the structure ground water can also flow into streams of DNA. and lakes, creating the same pollution On a global scale, a build-up of effects as effluent that has directly entered carbon dioxide in the atmosphere from surface water. fossil fuel combustion may eventually affect climate and dramatically change ecosystems by causing global warming. Also, the ADVERSE IMPACTS FROM release of chlorofluorocarbons and similar AIR-CARRIED POLLUTANTS chemicals also depletes the ozone layer in There is increasing evidence that the earth’s stratosphere, thus exposing chronic exposure to certain levels of air living things to harmful levels of ultraviolet pollution impedes the long-term survival radiation with potentially dangerous global and vigor of many species of plants. Trees implications. in heavily polluted urban areas, for example, have a much shorter life span than trees of the same species in less polluted

28 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE identify. First, staff will need the tools to ADVERSE IMPACTS TO LAND-BASED SYSTEMS determine the appropriate spatial and Many land-based activities fragment temporal scales as they plan and conduct or simplify ecosystems through pollution, their management activities. In the past, we either by direct effects such as an oil spill or have been most comfortable managing through secondary impacts such as soil individual DNR properties on a short time erosion from poor farming practices. Spills frame (ten years or less), a scale at which of hazardous materials can affect local areas we are able to observe immediate and by smothering animals or interfering with obvious impacts, obtain the most informa- their movement. Improper disposal of tion, and provide the most certainty. In the hazardous wastes can result in local con- future, we will be managing at a larger centrations of metals or organic compounds scale, considering entire landscapes and that harm organisms and ecosystems. much longer time frames, with less obvious Pesticides and herbicides can kill nontarget immediate impacts. species, changing the species composition One important tool that will help us in the area and weakening the ecosystems think and plan on the landscape scale will in which these organisms lived. be the delineation of ecoregions. Ecoregions Land-based pollution also impacts are large areas of the state that exhibit other systems, most often surface and similar patterns in potential natural com- ground water systems. For example, an munities, soils, hydrologic conditions, improperly functioning landfill may landforms, lithology, climate, and natural contaminate nearby soil and harm plants processes. Ecoregions provide a focus for and animals living in the immediate area. resource assessment and inventory of biotic However, leachate from the same landfill and abiotic elements. We need to determine may also enter the ground water and the most useful boundaries for ecoregions contaminate lakes and streams miles away. within the state, and develop goals and management strategies for them. These will IMPLICATIONS OF ECOLOGICAL ISSUES give us the framework needed to choose our priorities and focus our resources on Our current understanding of ecosys- carefully selected programs and projects. tems and, specifically, the implications of We will also need data management ecological simplification, fragmentation, techniques such as computerized Geo- and pollution present considerable oppor- graphic Information Systems (GIS) to compile tunities and challenges to the Department’s information on ecosystems and landscapes management programs. Present-day man- and to design process-oriented manage- agement strategies consider biological ment approaches. These and other tools, diversity mostly in a peripheral sense. such as a statewide aquatic and terrestrial Although awareness is increasing, overall inventory, will help us collect and manage program planning is not consistently based the extensive amount of information on the principles of ecosystem manage- needed to make decisions at a landscape ment. It is these principles that will allow scale. us to address biodiversity within the The issues of ecological simplification, context of ecological, socio-economic, and fragmentation, and pollution are not institutional concerns. These principles and distinct issues that can be debated or their application to Department programs weighed in isolation from each other. Like are fully discussed in the next section, ecosystems themselves, these issues are “Addressing Biodiversity through Ecosys- often interrelated and complex. Ecosystem tem Management.” management focuses on evaluating the While the main implication of the cumulative impact of proposed actions at ecological issues is the need to implement the landscape level. At the same time, ecosystem management, there are a number fragmentation and simplification may not of related implications that are important to always be bad. For example, the creation of

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 29 BIODIVERSITY: ISSUES & IMPLICATIONS

edge between habitats to enhance populations of game species is desirable in certain ADDRESSING BIODIVERSITY T HROUGH locations within the landscape. The impor- ECOSYSTEM MANAGEMENT tant thing is to recognize the complex impacts of our proposed actions (how WHAT IS ECOSYSTEM MANAGEMENT? much edge is desired and where should it To understand what the conservation This report be?), to clarify why they are desirable, to of biodiversity will mean to the proposes that the know the trade-offs, and to try to under- Department’s management system, we must best way to stand their impacts on ecosystem understand ecosystem management as the sustainability and, especially, on our address philosophy and process we will be using in options for the future. biodiversity as a the future. Ecosystem management is a Because our understanding of the system to assess, conserve, protect, and management issue ecological issues is constantly increasing, restore the composition, structure, and is to apply the we can use our current understanding to function of ecosystems, to ensure their principles of make decisions to implement now and then sustainability across a range of temporal ecosystem adapt as we learn more. One approach that and spatial scales and to provide desired we will need to explore is currently known management to ecological conditions, economic products, as adaptive management. Adaptive manage- and social benefits. Department ment considers many alternate manage- planning and The above definition reflects the ment scenarios developed collaboratively complexity that the ecosystem management programs. by scientists, managers, and policy makers. “umbrella” is meant to encompass. What Ecosystem Models are then developed to predict the does it really mean? In simpler terms, management is a results from each management alternative, ecosystem management blends human system to assess, management prescriptions are subsequently needs and values with ecosystem capability chosen, and monitoring programs are conserve, protect, and sustainability (Fig. 7). This blending of designed to scientifically test whether the and restore the multiple perspectives is essential to making management alternative does indeed wise choices about how resources will be composition, accomplish the predicted results. In this managed to result in an agreed-upon structure, and way, the management practice enhances the pattern of land uses. This pattern will be a function of “institutional memory” that documents our mosaic of cities, towns, and suburbs; parks, ecosystems, to decision-making process while continually lakes, wetlands, and forests; farms and improving the science base for our manage- ensure their industries; natural areas and reserves; and ment practices and advancing our knowl- corridors and riverways. sustainability edge of ecosystem functions. across a range of The landscape pattern that is desired Because the focus of this report is on will be defined over time as we work with temporal and the management of public lands, we do not multiple interests and partners in decision- spatial scales, and propose specific recommendations for how making. This report proposes that we make to provide desired the Department’s regulatory programs a commitment to the principles and ecological might change to address the conservation processes of ecosystem management and of biodiversity. However, because pollution conditions, fully develop within that commitment the can seriously affect the biodiversity of a economic goals and criteria that will be needed to variety of aquatic and terrestrial communi- conserve biodiversity as an essential products, and ties, in the long run we must consider how element of ecosystem sustainability. social benefits. the science of biodiversity can be incorpo- In listing these principles and pro- rated into the regulatory and technical cesses, it may be helpful to think of them in assistance work of DNR’s environmental the following two categories: Ecosystem quality programs. Approaches and Critical Thinking Skills.

30 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Ecosystem Human Needs Wisconsin’s Landscape, Management and Values Desired Continental, and Applies Tools Landscape Global Contexts societal resource integration teams needs future cities, towns, suburbs adaptive management generations parks, lakes, wetlands, public involvement and forests farms, industries natural areas TO BLEND ➤ Ecosystem corridors, riverways Sustainability ➤ biological diversity air, WITHIN land, and water quality

TO HELP SHAPE ➤

goals. Set goals within ecoregions to Figure 7 ECOSYSTEM APPROACHES meet a wide variety of diverse ecological The processes and Management goals should be set and and socio-economic needs, including components of the conservation of biodiversity. ecosystem manage- action taken using an ecosystem management. ment approach that integrates staff and resources across programs and disciplines. Manage to preserve ecological compo- Current statutory charges were developed sition, structure, and function. Con- to manage single species or small assem- sider not only immediate impacts but blages of economically important species. also the dynamics of long-term changes However, biological diversity is not, and induced by proposed management should not be treated as, a separate organi- actions. zational subprogram. Rather it will be included within the entire range of issues Manage at a landscape scale. Deter- that DNR managers must consider when mine both spatial and temporal scales they carry out the agency mission. appropriate to a problem or project. Assess the impacts of decisions made at Determine ecoregion boundaries and any one scale on both smaller and larger use them to develop management scales.

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Incorporate a transdisciplinary per- environment. Continue to bring diverse spective. Develop and support a diverse interests together to discuss and resolve staff with a working knowledge of a the issues. Work to anticipate problems, wide range of disciplines and a willing- manage conflict, and avoid the kinds of ness to integrate those disciplines in confrontations between conflicting innovative ways. New ways of working interests that have occurred in other together will transcend the limitations states. posed by traditionally separate disciplines. Managers working in integrated Act on a vision of biodiversity that is teams will form the foundation for the grounded in scientific fact, clarified way we “do” ecosystem management. values, and hard-nosed realism. View projects that aim to conserve Find ways to take action without biodiversity with the same scientific “knowing all the answers.” Use a rigor that we view those to develop the management approach that can be landscape for other human needs. Be applied now while allowing us to responsive to the role that values play in continually increase our understanding exploring alternative solutions to of ecosystems and adapt our practices as problems and in selecting final ap- we learn more. Used within an ecosys- proaches. tem management approach, the adaptive We won’t be able to implement all of management approach described earlier the recommendations and possible actions can help us do this in a formal and in this report at once. We will need to set structured way. realistic priorities and clear course of action that provides a roadmap for DNR staff and our partners and clients. CRITICAL T HINKING SKILLS Employees and the public must be USING ECOSYSTEM MANAGEMENT encouraged to take responsibility for ROUTINELY examining their own knowledge and beliefs. Commitments to solutions that treat We will be expected to apply the problems, not just symptoms must also be principles of ecosystem management on a encouraged. This will require that we foster daily basis. Some aspects of ecosystem critical thinking skills. Critical thinking is a management are familiar to DNR managers, process of reflection and analysis that many of whom have been thinking in terms involves the identification of assumptions of ecosystems and integrated approaches and known facts, the exploration of alterna- for many years. Many of the procedures tives, and the integration of new under- required to conserve biodiversity are standings into thought and behavior already included in our “toolboxes,” and patterns. For example, in order to support others need to be invented. Whether the critical thinking, we must: ideas are familiar or new, all managers will have questions about how to use ecosystem Provide DNR employees and the management principles in daily work and public with training and awareness how to meld these comprehensive manage- opportunities. Prepare staff to lead the ment approaches with traditional manage- public dialogue that will produce a clear ment activities. They will ask questions and widely accepted understanding of like: “How much restoration can we do?,” biodiversity and decisions that we can “How should we view active management collectively support and implement in practices like fish-stocking and our day-to-day living. clearcutting?,” “How can we reconcile biodiversity with user demands?,” “How Lead the discussion clarifying public does ecosystem management consider the values related to biodiversity and the needs of humans as well as the needs of all

32 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE other species?” The answer to each question The ecosystem management decision The ecosystem depends on the specific circumstances or model shown in Figure 8 provides a management context in which it is asked. For example, a framework that requires us to approach decision model decision to restore a damaged community decision-making from different perspec- is appropriate in some situations but tives. By examining several alternatives presents a inappropriate (or impossible) in others. relative to their consequences in ecological, structured Similarly, a decision to stock fish may fulfill socio-economic, and institutional contexts approach to the a user demand at the expense of native in the PLAN phase, the model holds that search for species, or it may replace a missing preda- we will make wiser management decisions. solutions that tor and restore balance to a lake’s ecosys- Whatever alternative we decide to imple- integrate tem. The appropriateness of any management (DO) is then monitored (CHECKED) ment practice or environmental decision for its actual results in all three contexts, ecological, socio- depends on its context. There are no easy and we revise our management (ACT) economic, and answers or “cookbook” formulas for according to those results. Our success as institutional management practices that will always resource stewards is a function of our concerns. conserve biodiversity. Rather, we must use a ability to understand, analyze, and integrate management model that brings perspec- alternatives across all three of these con- tives and knowledge from different disci- texts. plines together in an integrated search for Figure 8 alternatives. An ecosystem management decision model.

Plan Check

Ecological Ecological

Analyze Socio- Socio- Economic Do Economic Alternatives Decisions, Actions Institutional Institutional

Adapt Act

Success Complete

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 33 BIODIVERSITY: ISSUES & IMPLICATIONS

THE ECOSYSTEM MANAGEMENT DECISION MODEL

The ecosystem management decision model is based on an examination of questions and considerations within each of three contexts: the ecological, socio-economic, and instituional:

THE ECOLOGICAL CONTEXT Defining and subsequently managing in an ecological context is both an art and a science. As our understanding of ecological concepts grows, as our body of ecological theory and scientific evidence is enriched through time, and as our measures of the impact of various decisions on the ecosystem become more predictable and precise, our ability to identify and make informed ecological choices will increase. Depending on the ecosystem and the management issue, a variety of considerations are used to establish the ecological context. The process of determining the ecological context begins with a definition of scale, followed by an assessment of the system’s capability and function.

SCALE FUNCTION Question: At what spatial and temporal scales should the Question: How can we improve or protect the system’s decision be defined? function? How well is it working now? Some considerations: Some considerations: ✓ ✓ size of the affected system and its parts interrelationships between the abiotic and biotic components—missing components (composition, ✓ current inventory of what is there now, including structure) measures of biological ✓ diversity connectivity ✓ ✓ species with critical needs or special status fragmentation ✓ ✓ presence of ecological gradients and corridors resilience ✓ ✓ patterns of community distribution gene flow ✓ ✓ existing patterns of disturbance or change energy and nutrient flow ✓ ✓ current management practices, land use, and land food webs ownership

CAPABILITY Question: What is the system’s past, current and potential future ecological capability; where is the landscape headed? Some considerations: ✓ natural and potential capability (e.g., presettlement vegetation as one indicator) ✓ history of successional stages ✓ potential for recovery, enhancement, or expansion ✓ potential to be self-sustaining

34 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE THE ECOSYSTEM MANAGEMENT DECISION MODEL, CONT’D

THE SOCIO-ECONOMIC CONTEXT The Department also manages in a socio-economic context that reﬂects the varied and sometimes conﬂicting needs and values of people. Our mission requires us to be responsive to society, yet it recognizes that our decisions also help to shape society’s values. Furthermore, our actions have direct impacts on the economy at the local and state levels, and beyond. Our decisions are evaluated within this socio-economic context, and society determines our success or failure. As we work with stakeholders to understand their views and values, we also need to understand the impact of management activities on local business, land values, and other economic factors, including a view of long-term economic perspectives. We can understand the socio-economic context by knowing the stakeholders and the range of economic issues they represent.

SCALE ECONOMIC IMPACTS Question: At what spatial and temporal scale should the Question: What are the opportunities and threats for decision be deﬁned? What is the magnitude of social and business and employment? economic effects? Some considerations: Some considerations: ✓ direct and indirect impacts on local and state ✓ number of people affected, directly and indirectly businesses ✓ cost and funding for the project ✓ potential new businesses created ✓ time period for completion ✓ impact on current and projected property values ✓ time period of social or economic impacts ✓ impact on employment ✓ scope of social or economic impacts (local, state, ✓ sources of raw materials or other resources for regional, national, international) business ✓ relationship to transportation or utility networks ✓ relationship to national and international economy LAND USE Question: What are the past, present, and potential land uses? STAKEHOLDER INTERESTS Some considerations: Question: Who are the stakeholders and how can they ✓ current and previous land uses best be involved? What are their various perspectives, needs, and values? ✓ projected changes in land uses ✓ alternative future land uses Some considerations: ✓ ✓ indirect effects on adjacent or regional land uses major and minor stakeholders (public and private) ✓ ✓ land ownership patterns role of elected ofﬁcials ✓ public involvement and information strategies ✓ relationship of stakeholders to DNR and to each other ✓ opportunities for partnerships with stakeholders

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 35 BIODIVERSITY: ISSUES & IMPLICATIONS

THE ECOSYSTEM MANAGEMENT DECISION MODEL, CONT’D

THE INSTITUTIONAL CONTEXT As an institution, the Department operates within a matrix of complex institutional relationships. First, the Department’s actions must be based on sound legal authority. This authority is defined by state constitution, state and federal statutes, administrative rules, and court decisions. Second, Department actions are also affected by internal policies and by budgets, staffing, and various authorities granted to the agency. Management actions that do not fit within this external or internal institutional framework or authority are either not feasible or require changes in our laws, codes, or mission. And third, the Department has many opportunities to create cooperative agreements and partnerships with other public and private institutions, as well as with individuals, in order to meet mutual goals for resource protection, restoration, or enhancement.

LEGAL FRAMEWORK INTERAGENCY COOPERATION AND PRIVATE PARTNERSHIPS Question: What institutional support or constraints Question: What kinds of relationships with other public come from outside DNR? What is DNR’s legal relation- agencies or with private interests are needed? ship to local, state, and federal governments? Some considerations: Some considerations: ✓ existing and potential partners and cooperators ✓ legal authority (statutes and administrative rules) ✓ institutional constraints of partners and coopera- ✓ legal constraints tors, such as their ability to enter into long-term ✓ processes for obtaining decisions from appropri- agreements ate authority (local government, legislature, ✓ innovative approaches to cooperation and partner- federal agencies) ship ✓ need for new legal authorities or changes in existing laws ✓ state budget development, federal grants, etc.

INTERNAL DNR POLICIES: Question: What internal policies or procedures support or hinder a proposed action? Some considerations: ✓ strategic plans ✓ budget and stafﬁng ✓ manual codes and handbooks

36 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Is this model all that new? Many parts of the model are not new, in that they RECOMMENDATIONS represent the way many managers have Implementing ecosystem management thought and acted for years. The Depart- in Wisconsin requires concrete actions. The ment has conducted a number of planning following recommendations are the product efforts that are based on watersheds or of review by and dialogue with internal other ecological units and have dealt with a DNR staff and external partner agencies, range of socio-economic and institutional scientists, interest groups, and citizens. The issues. Examples are the Green Bay Reme- recommendations are put forth to begin the dial Action Plan, the Winnebago Pool process of working with the Natural Integrated Resource Management Plan, the Resources Board and all our partners and Milwaukee River Integrated Resource customers to outline the details of specific Management Plan, and the Mississippi actions needed. As we move forward, we River Strategic Plan. There are other will need much more discussion, both examples cited as case studies within the internally and with the public. following chapters on Wisconsin’s biological communities. These include projects Apply ecosystem management prin- related to the Baraboo Hills, Marathon ciples and practices to the Department’s County Forest, Habitat Restoration Area programs so that goals and priorities for (formerly Glacial HRA), Southern Unit biodiversity can be determined in the Kettle Moraine State Forest, and Patrick context of ecological, socio-economic, and Lake wetland mitigation. institutional issues. What is new is putting all the parts together and using them throughout the Use the ecosystem management decision Department as a decision framework. Also model, as described in this report, to new is the importance of considering propose and evaluate alternative actions multiple levels of scale in all decisions. from the ecological, socio-economic, While some managers already do this, we and institutional perspectives. need to work together to do it consistently, with attention to all contexts of the ecosys- Develop and use ecoregion goals and tem-management decision model. objectives to design and implement Success lies in carefully considering geographically-based management all three context—ecological, socio-eco- guidelines. These procedures will nomic, and institutional—and finding an provide the criteria needed for: alternative that is acceptable in all three. We cannot only consider the ecological ✓ land acquisition priorities context in making management decisions. The “correct” ecological decision may be ✓ new and revised master plans for impossible or unworkable given the state-owned property realities of our society, the economy, and ✓ priorities for restoration of biological the institutions in which we work. Simi- communities larly, the “best” economic decision may be ecologically disastrous. Our hope is to ✓ use of appropriate genotypes in provide an open public process in which restoration and management social, economic, and ecological perspec- ✓ management of populations of tives are evaluated and weighed early on— troublesome non-native species before positions become hard and fast. It is, in reality, a search for acceptable alterna- ✓ goals for consumptive use, such as tives that help preserve long-term options harvest of forest products, fish, and and improve the quality of our everyday wildlife lives and the lives of the generations to come.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 37 BIODIVERSITY: ISSUES & IMPLICATIONS

Continue to develop a Geographic Build partnerships with other agen- Information System (GIS) as a major cies, local governments, tribes, the busi- management tool needed to implement ness community, scientists, and other ecosystem management and to establish interest groups to accomplish common and evaluate ecoregion goals and goals for ecosystem management, includ- objectives. Train staff to use GIS to ing specific attention to biodiversity. analyze and monitor regional landscapes. Seek innovative ways to work with partners to set landscape-scale goals that Continue to develop and use a statewide cross ownership boundaries. resource inventory of the status and distribution of aquatic and terrestrial Continue to participate in joint efforts species, biological communities, and such as the 1994 Wisconsin Forest other attributes within the ecoregions of Accord (a memorandum of understand- Wisconsin. ing to adopt uniform criteria to describe, evaluate, and share critical ecological Use the biennial budget guidance to information among private landowners address management priorities and goals and nonprofit, county, state, and federal for ecosystem management. agencies) and the Interagency Coopera- tion on Ecosystem Management or Review existing laws and procedures to ICEM (a multi-agency resolution form- analyze their consistency with the ing a consortium of 20 Midwestern state principles of ecosystem management. and federal agencies, including Modify them where necessary. This will Wisconsin’s DNR, working together to be a formidable task, since our proce- coordinate ecosystem management dures are embodied in a multitude of activities). handbooks and guidelines. However, these procedures must be consistent and Take advantage of the knowledge and must work for, not against, ecosystem capability of the scientific community in management and the conservation of reviewing how we apply scientific biological diversity. knowledge to our management strategies. Monitor and respond to ecological, socio-economic and institutional issues Seek input from business and economic as they develop. Examples of current interests to develop strategies to imple- issues that arise in the ecological context ment ecosystem management. include the role our management actions may play in ongoing fragmentation and Work with hunting and fishing groups simplification, the need for prescribed to use the principles of ecosystem burning as a management tool, and the management to improve the quality of desire to manage for the range of hunting and fishing in Wisconsin. successional stages of each major community type. Encourage and support efforts at the national level to screen proposed Employ both research and experimental introductions of non-native species. management to develop new management approaches and to refine existing ones. These efforts should be directed at expanding our understanding of ecosystems and clarifying options for future management.

38 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Build partnerships with private Design approaches to outreach and landowners to accomplish common goals training to increase understanding of for ecosystem management, recognizing ecosystem management and to clarify that the Department cannot accomplish the variety of professional and personal the breadth of what needs to be done to values regarding biodiversity. conserve biodiversity by working on public lands alone. Provide support to employees to increase their skill in bringing diverse Establish coalitions and partnerships groups of people together to discuss and with private landowners to protect and resolve issues related to ecosystem restore biological diversity. management and biodiversity.

Provide technical assistance, economic Set priorities for implementing the incentives, and education to encourage possible actions speciﬁc to each of private landowners to participate in the Wisconsin’s seven biological community conservation of biological diversity. types described and assessed in the remaining chapters of this report. These Develop innovative and proactive possible actions are described in detail at information and education strategies for the end of each of the seven biological Department staff and the public regarding community chapters. We call these “pos- biodiversity and its relation to ecosystem sible actions” because they are consistent management. with ecosystem management but require more analysis and planning. The following Create ecosystem management demon- possible actions are consistent with ecosys- stration areas in each ecoregion that tem management, but require more analysis invite hands-on participation and and discussion. How priorities are set illustrate applied ecosystem manage- within this list will be based on ecoregion ment and biodiversity concepts. goals, staff workload, ﬁscal resources, public input and support, and legal authority. We will work with our customers and clients to set priorities and bring recommendations to the Natural Resources Board for consideration beginning in the 1995-97 biennium.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 39 OVERVIEW OF WI BIOLOGICAL COMMUNITIES

IMPACT OF GLACIATION

The greatest historical event impacting Wisconsin vegetation occurred 10,000- CHAPTER 3 60,000 years ago when Wisconsin was invaded by continental ice sheets. These glaciers transformed Wisconsin’s landforms and vegetation. Vegetation was scoured away and mountains were planed down in Overview of all except the southwestern region of the state, leaving a rolling plain covered by a layer of glacial till. Remnants of Wisconsin’s earlier topography are visible in hard rock Wisconsin’s outcrops such as Rib Mountain and the Baraboo Hills. Each interglacial period, including the present one, was revegetated through migration, which occurred through Biological range extension and seed dispersal to favorable habitats. Migrants originated from communities centered as far away as the Communities Ozarks, Pennsylvania, Texas and other areas. Some of the tree species now in by Betty Les Wisconsin had glacial refuges in the Bureau of Endangered Resources southern Appalachians and the eastern coastal plain. Although only a portion of Department of Natural Resources species were able to perpetuate themselves over the long term, what is now Wisconsin regained tremendous floristic diversity The location and through migration and colonization follow- DETERMINANTS OF BIODIVERSITY ing glaciation. An additional component of extent of plant floristic diversity is derived from the relict communities he location and extent of plant species occurring in the Driftless Area of and the animals communities and the animals southwestern Wisconsin. These species associated with associated with them are deter- pre-date glacial activity and often exist mined by environmental gradi- them are nowhere else in the state. ents of moisture, temperature, determined by The glaciers also had a tremendous soil type, and climate. They are influence on Wisconsin surface waters. environmental also shaped by historical events, migration, Glacial deposits dammed rivers and gradients of and natural and human-induced distur- scoured out lakebeds creating large water moisture, bance. The most pronounced environmen- bodies such as the Great Lakes and Lake temperature, soil tal gradient in Wisconsin is located in a Winnebago. Some small water bodies were type, and climate. narrow band that runs from northwestern created by the numerous pits or depres- to southeastern Wisconsin. This band has They are also sions in the glacial till, which filled with been termed the tension zone (Fig. 9). shaped by groundwater. In the north, most are found Many species of plants and animals reach in sandy, pitted outwash and were formed historical events, the limit of their ranges in this zone. In when buried ice blocks melted after retreat migration, and Wisconsin, the tension zone delineates the of the glacial ice. The nature of these natural and northern forest, including the boreal aquatic features was determined, in large human-induced element, from the southern forest and part, by their landscape position. For prairies. Although climate is a major reason disturbance. example, the landscape position of lakes in for the tension zone, soil type and other the groundwater and surface flow system factors also play a role. determined their basic water chemistry.

40 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Basic water chemistry, in turn, influenced Bayfield and continues to influence the sensitivity of Douglas Ashland waters to eutrophication and such present- Iron day concerns as mercury toxicity and acid Vilas Sawyer rain. Landscape position has also influ- Washburn Price Forest Florence enced the species present in a given lake Oneida Burnett Polk Marinette because it determines how connected that Barron Rusk Lincoln particular lake is to other lakes and sources Langlade Taylor Oconto of colonization. Thus, the glaciers created a Chippewa St. Croix Dunn template, or backdrop, on which present- Marathon Menominee Clark Shawano day waters have developed. Pierce Eau Claire Door Pepin Wood Portage Waupaca Buffalo Outagamie Brown Jackson Kewaunee

IMPACT OF OTHER NATURAL Trempealeau Juneau Adams Waushara Winnebago Manitowoc Monroe

DISTURBANCE La Crosse Calumet Marquette Green Lake Sheboygan

Other types of natural disturbance Vernon Fond Du Lac Sauk Columbia Dodge Richland Washing- have also inﬂuenced Wisconsin plant and ton Crawford Ozaukee animal communities. Chief among these Dane have been windstorms, lightning-induced Iowa Jefferson Waukesha

fire, and droughts, which were often a Grant Milwaukee Green Rock factor in severe fires. Floods have also Lafayette Walworth Racine Kenosha influenced the natural communities, but to a lesser degree. Historically, all of these factors individually glacier probably Figure 9 and in combination created extensive Location of the tension have impacted the The greatest historical event impacting floodplains and zone, adapted from landscape. Climate Wisconsin vegetation occurred 10,000- terraces in south- Curtis (1959). and temperature 60,000 years ago when Wisconsin was central Wisconsin. changes have greatly The presence of invaded by continental ice sheets. These influenced the prairie species in the significance of these glaciers transformed Wisconsin’s pollen record of this disturbances. landforms and vegetation. Vegetation was period, plus an Windstorms scoured away and mountains were planed increase in charcoal, frequently produced down in all except the southwestern indicates an inter- disturbance that spersion of dry varied in significance region of the state, leaving a rolling plain covered by a layer of glacial till . . . . The periods and fire. The from a local to frequency, combina- glaciers also had a tremendous influence landscape scale. A tion, and size of these recent example of this on Wisconsin surface waters. disturbances contin- is on the Flambeau ues to influence the River State Forest, mosaic of the natural communities. where much of the old growth was destroyed in a minimum of five different windstorms between 1949 and 1977. The HUMAN-INDUCED DISTURBANCE most significant of these storms occurred in the downburst of 1977, when hundreds of Human-induced disturbance had a thousands of acres across the state were profound impact on Wisconsin plant and disturbed. A much earlier example of animal communities, exceeded only by the climatic influence occurred during the impact of glaciation. For centuries before warming period that followed the last Euro-American settlement, Native Ameri- glaciation. Floodwater from the melting cans lived in the area now known as

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 41 OVERVIEW OF WI BIOLOGICAL COMMUNITIES

Although the Wisconsin. Although the size of their created in the mid-1800s, when the U.S. landscape was populations and the extent to which they Geological Survey’s land survey of Wiscon- used the land are undocumented, both are sin was completed. This was some 350 affected as Native probably far greater than once thought. years after contact, well after disease Americans used Scientists estimate North American popula- introduced by European explorers had resources and tions at 3.8 million or more at the time of decimated Native American populations. interacted with the European contact (Denevan 1992). The Vegetation maps based on the land survey land, changes Wisconsin region no doubt supported large records show a diversity of natural commu- reflected a level of Native American populations due to its nities including extensive forests and abundant natural resources. wetlands plus the fire-dependent grassland, human use many Fire was one of the Native Americans’ barrens, and savanna communities (Fig. times less intensive primary tools for managing these resources. 10). These communities are commonly than present-day Fire was used to concentrate game for referred to as Wisconsin’s presettlement use. Urbanization, hunting, increase game habitat, and clear vegetation. This convention was adopted highway paths for travel. Also, natural fires went for our report. construction, and largely unsuppressed. The result was Euro-American settlement marked the the host of other development of extensive communities— beginning of a simplification of Wisconsin’s prairies, savannas, barrens, and oak wood- landscape and a decrease in biodiversity. In developments lands—that were fire-dependent and, in the absence of fire, the prairie, savanna, associated with fact, a product of fire. Approximately 40%- barrens, and oak woodland communities modern life have 45% of Wisconsin’s land surface was gradually filled in with shrubs and trees. produced covered by these fire-dependent communi- When settlers realized the depth and tremendous ties prior to Euro-American settlement richness of the prairie and savanna soils, changes in (Curtis 1959). The more mesic northern these areas were cleared for agriculture and and southern forests were fire resistant, but grazing, leaving only traces of the original Wisconsin’s their composition and structure were plant communities. Forested areas were landscape. probably altered to some extent through either cleared for farming or cut for timber intentional management by Native Ameri- as the need for building material surged at can populations. the turn of the century. Except for pockets The nature of the plant communities of forest, northern Wisconsin was com- Nineteenth century land surveying, a pen prior to European contact is unknown, pletely cut over. Slash timber left on the drawing from the although scientists continue to piece ground fueled unnaturally severe fires that collections of the State together a description based on the record further denuded the land and at times Historical Society of left by pollen, sediment, and explorers’ damaged the soil. The forest was slow to Wisconsin. Illustration courtesy of Kenneth journals. The first systematic record of regenerate itself, and when it did, it was Lange. Wisconsin’s vegetation communities was very different from the earlier forests. Although the landscape was altered as Native Americans used resources and interacted with the land, changes reflected a level of human use many times less intensive than present-day use. Urbaniza- tion, highway construction, and the host of other developments associated with modern life have produced tremendous changes in Wisconsin’s landscape. Fragmentation and simplification of plant communities and pollution have accompanied these changes. Impacts of these disturbances are discussed in more detail in the “Ecological Issues” section of this report.

42 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE These seven communities represent THE SITUATION T ODAY an aggregation of the more numerous communities described by Curtis (1959) Today, Wisconsin has forest cover (Table 1). Curtis’ system of classifying roughly equal to that in place at Euro- vegetation was chosen as the framework for American settlement, but it is very different this report because it was designed specifi- in age structure and species composition. cally for Wisconsin Barrens, savannas, and has stood the and grasslands exist test of time. Recent but only in scattered Today’s remaining biological communities interest in locations. Postglacial biodiversity and lakes and rivers have provide valuable sources of genetic and ecosystem manage- remained relatively species diversity. Our challenge is to ment has spurred the constant in number retain the range of diversity still present development of a and surface area, but and, where possible, regain diversity number of regional we have lost about through restoration. and national systems one-half of our for classifying wetlands along with communities and the seasonal ponds associated with them. ecosystems. One such system, the National Most of what we have left of prairies, Hierarchy of Ecological Units, has been savannas, and certain wetlands (e.g., sedge adopted by the Department’s Division of meadows) is the result of managed use of Resource Management as the standard for fire, since these community types are fire- its work (U.S. Dep. Agriculture 1993). The dependent. Curtis system and others such as Kotar et Thus, today Wisconsin still has a great al. (1988) will nest within this hierarchy, deal of biological diversity, but it has also which is designed to stratify the Earth into lost a great deal of diversity (Fig. 11). All of progressively smaller areas of increasingly today’s communities provide valuable uniform ecological potentials. Thus, it can sources of genetic and species diversity. be used at multiple geographic scales Our challenge is to retain the range of ranging from a single site in our state to an diversity still present and, where possible, area that spans several states or the entire regain diversity through restoration. In nation. Maps depicting ecoregions and going about this work, we must strive to various other ecological units will be measure diversity in ways other than species developed to assist in setting management richness. Understanding and measuring goals and objectives. diversity at a functional level will put us in In this report, each of the aggregated a better position to predict the impact of communities is described and compared to various actions on plant and animal com- its presettlement status (for an overview of munities. how rare each of the communities has The following sections of this report become, see the global ranks in Table 1). profile Wisconsin’s seven major biological After status, actions causing concerns and communities: socio-economic issues related to conserva- northern forests tion of each community are discussed, and the potential for restoring the community southern forests to a sustainable, functional state is assessed. oak and pine barrens Finally, possible actions are listed for managing and restoring each community. oak savannas Note that some of Curtis’s communities are grasslands discussed in more than one of the aggregated communities due to overlap in wetlands composition, structure, or function. Al- aquatic systems though the communities are categorized

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 43 OVERVIEW OF WI BIOLOGICAL COMMUNITIES

Table 1. Wisconsin plant communities. Compiled by R. Henderson based on Curtis (1959) and the Wisconsin Natural Heritage Inventory.

Aggregations Curtis (1959) Communities Aggregations Curtis (1959) Communities Used in this Report and their Global Rank* Used in this Report and their Global Rank*

Northern Forest Boreal Forest (G3)Grassland Bracken-Grassland (G3) Northern Dry Forest (G3) Sand Barrens Northern Dry-Mesic Forest (G4) Dry Prairie (G3) Northern Mesic Forest (G4) Sand Prairie** Northern Wet-Mesic Forest (G3)Dry Lime Prairie** Northern Wet Forest+(G4) Dry-Mesic Prairie (G3) Mesic Prairie (G2) Wet-Mesic Prairie (G2) Wet Prairie+ (G3) Calcareous Fen+ (G3) Southern Sedge Meadow+ (G3) Northern Sedge Meadow+ (G4) Southern Forest Southern Dry Forest (G4) Aquatic Emergent Aquatic (G4) Dry Oak Woodland** Submergent Aquatic Southern Dry-Mesic Forest (G4) Lake Beach Mesic Oak Woodland** Southern Mesic Forest (G3) Southern Wet-Mesic Forest Southern Wet Forest+ Oak Savanna Oak Opening (G1) Wetland Open Bog (G4) Dry Oak Opening** Alder Thicket (G4) Dry-Mesic Oak Opening** Shrub Carr Mesic Oak Opening** Northern Wet Forest (G4) Wet-Mesic Oak Opening Southern Wet Forest Wet Oak Opening** Wet Prairie (G3) Cedar Glade Calcareous Fen (G3) Southern Sedge Meadow (G3) Northern Sedge Meadow (G4) Oak/Pine Barren Oak Barrens (G2) Minor Misc. Exposed Cliff Pine Barrens (G3) (not covered Shaded Cliff in the report) Lake Dune

* Rank reflects global rarity. Community classification is not standardized for the nation. Thus, not all of the communities have ranks, and some of the ranks had to be adapted to Wisconsin communities based on criteria for similar communities elsewhere. Also, some relatively low-ranked forest communities may be rare in some seral stages (i.e., specific occurrences of the community may be highly ranked). Ranks appearing in italics are considered tentative at this time. G1 = Critically imperiled globally because of extreme rarity (5 or fewer occurrences or very few remaining individuals or acres) or because of some factor(s) making it especially vulnerable to extinction. G2 = Imperiled globally because of rarity (6 to 20 occurrences or few remaining individuals or acres) or because of some factor(s) making it very vulnerable to extinction throughout its range. G3 = Either very rare and local throughout its range or found locally (even abundantly at some of its locations) in a restricted range (e.g., a single state or physiographic region) or because of other factors making it vulnerable to extinction throughout its range; in terms of occurrences, in the range of 21 to 100. G4 = Apparently globally secure, though it may be quite rare in parts of its range, especially at the periphery. G5 = Demonstrably secure globally, though it may be quite rare in parts of its range, especially at the periphery. ** Postulated communities. Due to their rarity, these communities cannot be rigorously quantified at this time. + Also covered to some extent in the Wetland Communities.

44 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Figure 10 (overleaf) Vegetation cover of Wisconsin in the mid-1800s, compiled from U.S. General Land Office Notes by Robert W. Finley, 1976. This map is based on the original land survey of Wisconsin carried out in the mid- 1800s by the U.S. General Land Office. The purpose of the original survey was to establish the township-range-section grid for Wisconsin. For each section and quarter-section point, nearby trees were selected as bearing trees and their diameters and distances from the corner were recorded. In treeless areas, the crew built a mound of earth at the corner point and recorded that the point was in an open area. Surveyors were also required to describe the timber and agricultural value of the land, its topography, and water bodies and to provide a general description and map for each township (Lange, 1990). These records were used by Robert W. Finley of the University of Wisconsin Extension to reconstruct the vegetation patterns present at the time of the survey. Dr. Finley completed his work in 1976. This map and others like it are useful in helping people visualize the general location, extent, and diversity of vegetation present in the last century. Finley’s map was originally designed and prepared by the Cartographic Laboratory, University of Wisconsin-Madison and was subsequently digitized by the University. The digital version presented in this report was further modified and enhanced by staff of the Department’s Bureau of Information Management, GEO Services Section.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 45 OVERVIEW OF WI BIOLOGICAL COMMUNITIES

Figure 10 Vegetation cover of Wisconsin in the mid- 1800s, compiled from U.S. General Land Office Notes by Robert W. Finley, 1976.

46 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Figure 11 Land use and land cover for Wisconsin, compiled from high- altitude aerial photography taken from 1971-81.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 47 OVERVIEW OF WI BIOLOGICAL COMMUNITIES

Figure 11 (overleaf) Land use and land cover for Wisconsin, compiled from high-altitude aerial photography taken from 1971-81. This map is based on data from the U.S. Geological Survey Land Use Data Analysis Program. Data were interpreted from 1:58,000 scale color infrared and 1:80,000 panchromatic aerial photography from the National High Altitude Photography Program. Photographs were acquired in the years 1971-81. The map is useful in helping people visualize the current land cover for Wisconsin and for assessing the magnitude of change over the past 100+ years. Although this map and the companion map on mid-1800s vegetation cover (Fig. 10) are based on very different types of data and technology, broad comparisons of cover types during the two time periods can be made. Figure 11 was produced by the Bureau of Information Management, GEO Services Section.

48 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE according to the plants they support, the numerous and functionally important faunal element of each community is also animal groups (e.g., insects) are the least discussed to the extent possible with documented or understood. Thus, animal existing information. Some of the most coverage is inadequate but points the way for future work.

In discussing restoration, it is important to note that we do not envision restoring communities to conditions prior to Euro-American settlement. This would clearly be an unrealistic goal. The presettlement status of each community is an important indicator of site potential and serves primarily as a guide and benchmark in our restoration efforts. Our desired state is a more diverse landscape, considering all four levels of diversity (genetic, species, community, and ecosystem level) across all land uses.

LITERATURE CITED

Curtis, John T. 1959. The vegetation of Lange, Kenneth I. 1990. A postglacial Wisconsin: an ordination of plant vegetational history of Sauk County and communities. Univ. of Wisconsin Press. Caledonia Township, Columbia County, Madison, Wisconsin. 657 pp. south central Wisconsin. Wis. Dep. Denevan, William M. 1992. The pristine Nat. Resour. Tech. Bull. No. 168. 40 myth: the landscape of the Americas in pp. 1492. Annals of the Assoc. of Amer. United States Department of Agriculture. Geographers. 82(3). pp 369-385. 1993. National hierarchical framework Kotar, J., J.A. Kovach, and C.T. Locey. 1988. of ecological units. Washington D.C. 19 Field guide to forest habitat types of pp. northern Wisconsin. Univ. Wis.- Madison Dep. of For. and Wis. Dep. Nat. Resour. 212 pp.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 49 NORTHERN FOREST COMMUNITIES

with dominance generally being shared by only two of three. Throughout the region, mature stands on the medium to rich soils CHAPTER 4 (loams and silt loams) are dominated by various mixtures of five or six principal species: sugar maple, basswood, hemlock, yellow birch, white ash, and American beech. Red oak and red maple are the most common minor associates. In presettlement Northern times, white pine was also a common associate in these northern hardwood forests. The poorer soils (sands and loamy Forest sands) are generally dominated by mixtures of pines (jack, red, and white), aspen, white birch, red maple, and red oak. Extensive wetland forests are also common to this Communities region. These can be divided into two basic types: conifer swamps (black spruce/ by John Kotar tamarack and white cedar) and hardwood Department of Forestry swamps (black ash, red maple, and elm). These broadly described forest types, University of Wisconsin-Madison based upon dominant vegetation, only and begin to reflect the total biological diversity Ronald Eckstein of forest communities of the region. A North Central District system of ecological classification of forest Department of Natural Resources communities and sites on which they occur is necessary. Such a system has been developed for forests in northern Wiscon- sin (Kotar et al. 1988) (Table 2) and will be completed for the rest of the state by 1994.

DESCRIPTION STATUS The term northern he term northern forest, as forest, as applied applied in Wisconsin, is prima- PAST in Wisconsin, is rily a geographic designation and primarily a does not in itself imply any POST-GLACIAL ENVIRONMENT geographic specific species composition. In broad terms, it may be character- designation and The entire area of present-day north- ized as a region of mixed deciduous and ern forest has been affected by Pleistocene does not in itself coniferous forests that represent one of the glaciation. Several major glaciations and imply any specific two distinct climatic zones in Wisconsin as countless minor ice advances and reces- species separated by a loosely defined S-shaped sions have created a complex pattern of ice composition . . . . It transition belt known as the “tension zone.” and meltwater-influenced deposits. Some of may be The region north of this zone is generally these deposits were subsequently covered called the northern forest (see Fig. 9). characterized as a by wind-blown silty or loamy material Forest communities, present and region of mixed called loess. This tremendously complex historic, display considerable diversity in array of deposits formed the parent material deciduous and composition of dominant species. About 30 from which present soils have developed coniferous forests tree species occur in the northern forest as and are, in fact, still in the process of occurring north of a whole, although fewer than ten are development. Although soil scientists the tension zone. usually found in any given community, recognize several hundred soil classes

50 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Table 2 Curtis (1959) Kotar et al. (1988) An ecological Communities Habitat Types classification system for the northern forest. Northern Dry Forest Acer-Quercus/Vaccinium Quercus-Acer/Epigaea Quercus/Gaultheria-Ceanothus Northern Dry-Mesic Forest Pinus/Maianthemum-Vaccinium Pinus/Amphicarpa Quercus/Amphicarpa The dominant Acer/Vaccinium-Desmodium vegetation of the Acer/Vaccinium-Viburnum northern forest Acer-Quercus/Viburnum Acer/Athyrium only begins to reflect the total Northern Mesic Forest Acer/Viola-Osmorhiza Acer/Hydrophyllum biological diversity Acer/Caulophyllum-Circaea of the region. Acer-Tsuga/Dryopteris Acer-Fagus/Dryopteris Acer-Tsuga/Maianthemum Northern Wet-Mesic Forest Tsuga/Maianthemum-Coptis within the region of the northern forest, the Outwash plains and terraces. These present soils can be grouped into four deposits are similar to those of the pitted The northern forest broad categories based on the mode of outwash, often sandier than pitted landscape. We see a glacial deposition of parent material. These outwash, but the terrain is flat or only matrix of forest with aquatic features are: gently sloping. Unless modified by a imbedded. The blanket of loess, pitted outwashes and Ground moraines or till plains, consist- continuous forest is a outwash plains form the droughtiest and mosaic of old-growth ing of assorted material including least fertile soils of the region. stands of hemlock boulders, gravel, and sand but usually (Plum-Star Lakes also containing considerable amounts of These four basic types of glacial Hemlocks State Natural Area is silt and clay. Soils developed from till are deposits form a moisture-nutrient gradient, between the lakes), usually the most productive. which is the strongest factor controlling the wildlife openings, and establishment of invading plant species. adjacent stands that have been harvested End moraines and recessional moraines. Plants themselves exert considerable (foreground). Photo by These deposits are also composed of till, influence on soil development. Even Michael J. Mossman. but are usually coarser textured than are ground moraines, and they form more rugged topography. The resulting soils are somewhat droughtier and have lower nutrient content than do soils derived from ground moraines.

Pitted outwash. These meltwater- deposited sands and gravels contain depressions (pits) that often have steep slopes and may be ﬁlled with water. Several large areas in northern Wiscon- sin are dominated by this type of landform (e.g., Burnett, Washburn, Vilas, and Oneida counties).

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 51 NORTHERN FOREST COMMUNITIES

though the original parent material has 1944). Members of the Alleghenian prov- been modified to varying degrees since the ince are more or less distributed through- last glaciation (10,000 to 60,000 years ago, out the southern and northern forests, depending on location), the distinctions while the boreal species are more prevalent due to parent material persist. in the northern half of the state and prairie Present community composition is the species more prevalent in the southern half. result of environmental influence (soil and There is evidence that many species climate) and various historical factors. It is are still extending their ranges; conse- not meaningful to speak of “original quently, floristic stability on the geologic vegetation” without reference to some scale may not be reached for some time specific time period. Many plant species even if the climate remains stable. This fact found in Wisconsin today may have been is often overlooked, especially in North present before the Pleistocene, but not America where the presettlement condition necessarily in present locations and in of the vegetation is often presumed to have present combinations. Paleoecologists have been in a relatively static climax state. This determined that present Wisconsin vegeta- change, however, is so slow that most of tion consists of elements from three distinct the changes seen in the past 150 years can floristic provinces: the Boreal, the Prairie, be attributed to the influences of Euro- and the Alleghenian (Hulten 1937, Cain American settlement.

Figure 12 Presettlement forests of Wisconsin, adapted from Finley (1976) as modiﬁed by Kotar (1990).

MILES

010203040

Northern Forests Mainly Coniferous Boreal (white spruce, balsam fir, tamarack, white cedar, white birch, aspen) Pine (white and red pine) Jack pine, scrub oak forest and barrens Mixed Coniferous-Deciduous Forests (mesic) Hemlock, sugar maple, yellow birch, white and red pine Sugar maple, yellow birch, white and red pine Beech, hemlock, sugar maple, yellow birch, white and red pine Southern Forests Mesic Sugar maple, basswood, red-white-black oak

Beech, sugar maple, basswood, red-white-black oak Dry-mesic and Xeric Forests and Prairies Oak forest (red, white, black and bur oaks)

Oak openings (bur, white and black oaks)

52 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE forest communities—they only identified COMPOSITION OF PRESETTLEMENT FORESTS individual trees—Finley constructed The exact nature of the floristic and abstract communities based on dominant structural composition and the geographic tree species. He organized the data into 11 variation of the northern forest in forest community types, seven of which presettlement times has never been de- represent the upland forests of northern scribed, and it probably will never be Wisconsin. Figure 12 is a redrawn and known with certainty. However, descrip- simplified version of Finley’s large and tions and occurrences of prominent forest complex map. It shows the primary distri- types, at least in terms of tree species bution of the six major forest types of composition, were recorded by numerous northern Wisconsin plus three southern early observers (e.g., Knapp 1871, forest types. Numerous scattered fragments Chamberlin 1877, Warden 1881). These of depicted types were deleted. The six early observers had already recognized the northern forest types are described as tension zone, without using the term, and follows: consistently described four to six forest 1. Boreal forest—white spruce, balsam fir, types occurring north of the zone: (1) pine tamarack, white cedar, white birch, forests, composed of aspen. This forest white pine and red type occurred in a pine mixtures with limited area of the The exact floristic and structural no hardwoods; (2) extreme northern part mixtures of hemlock, composition of the northern forest in of the state, near Lake sugar maple, and presettlement times has never been Superior. Most yellow birch, with, to described, and it probably will never be ecologists today agree the east, beech and known with certainty. that this community large white pine; (3) type, although scrub pines and resembling the boreal scrub oaks; (4) forests of Canada, is a hardwoods without conifers—mainly sugar distinct geographic variant of its north- maple, yellow birch, basswood and some- ern namesake. times a mixture of red oak and white oak; (5) swamp forests composed of spruce, fir, 2. Pine forest—dominated by white pine tamarack, and white cedar; and (6) oak and red pine. Contrary to the common openings or savanna (only south of the belief that most of northern Wisconsin tension zone). was covered by extensive pure stands of The best information on the composi- white pine and red pine, this forest was tion of the northern forest during the extremely limited even before Euro- earliest period of Euro-American settlement American settlement. The most exten- comes from the records of the rectangular sive block occurred in Vilas and Oneida survey of public lands (General Land Office counties. Surveys). These surveys contain a systematic record of the kinds and sizes of trees 3. Jack pine, scrub oak forests, and used as witnesses for lines and corners, as barrens. This is a loosely described type well as more or less detailed accounts of characterized by mixtures of poor- vegetational changes encountered. Finley quality trees or poorly stocked stands of (1976) produced a map of the jack pine, pin oak or bur oak, or some- presettlement vegetation of Wisconsin times red oak. Mixtures of red pine and based on survey records contained in 671 white pine, red maple, aspen, and white volumes of surveyor notebooks. These birch were often included. Figure 12 records describe 54,000 square-mile shows three principal areas of occur- sections and 110,000 linear miles of rence: Washburn, Burnett, and Douglas traverse. Although surveyors did not record

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 53 NORTHERN FOREST COMMUNITIES

counties in the northwest; Marinette does not follow that presettlement forests County in the northeast; and Juneau, were unaffected by perturbation and were Adams, and Jackson counties in the stable and in balance with regional climate. central part of the state. Of the six presettlement forest types described above, none can be explained 4. Hemlock, sugar maple, and yellow without invoking some form of environ- birch, with mixtures of white pine mental disturbance. The three northern and red pine. This was the largest and hardwood types (4, 5, and 6 above) could perhaps the most characteristic forest presumably self-perpetuate without the aid formation in northern Wisconsin. It is of disturbance, because sugar maple, also sometimes referred to as the “hem- hemlock, beech, and to some degree yellow lock-northern hardwood” or simply the birch are shade tolerant. However, the “northern hardwood” forest. presence of shade-intolerant white pine and especially red pine in these communities 5. Sugar maple and yellow birch, with a could not be explained without a distur- Simply because mixture of red pine and white pine. bance factor. present forest This type represents the southern and Although fires occurred less fre- western transition of the preceding type. quently in mesic hardwood stands than communities are Absence of hemlock, which reaches the they did in coniferous forests, many fire- known to be western limit of its natural range in scarred trees and stumps predating the largely the result of these regions, is its main characteristic. logging era were observed by early survey- human-caused Although this community designation ors. However, severe and extensive fires disturbances, it may appear arbitrary, the abrupt termi- were probably not very common in north- does not follow nation of the range of hemlock, a species ern Wisconsin. The main evidence for this that presettlement which is ubiquitous eastward to the is a very low occurrence of aspen and birch Atlantic coast, suggests a significant stands among presettlement forests. In the forests were climatic shift. Lake States, such stands are almost always unaffected by associated with fires. Finley’s map shows perturbation and 6. Beech, hemlock, sugar maple, and only widely scattered, small patches of this were stable and in yellow birch, with a mixture of red type and almost none within the northern balance with pine and white pine. American beech is hardwood-pine regions. There is also regional climate. another tree species that reaches the evidence that extensive windthrows in western limit of its range in Wisconsin. hardwood stands were even more common. Just as in the case of hemlock, climatic Often the majority of stumps from old- influence is presumed to control the growth pines are found on mounds or range of beech, although the role of knolls in stands that have a characteristic calcareous soils, climate, and incomplete kettle-knoll microtopography caused by the post-Pleistocene migration have been uprooting action of winds. In fact, numer- suggested as additional factors (Davis et ous studies have shown that disturbances al. 1986). have been occurring in somewhat cyclic fashion in all terrestrial ecosystems (Heinselman 1973, Lorimer et al. 1988). FACTORS CONTROLLING THE DYNAMICS OF PRESETTLEMENT FORESTS CLIMAX AND OLD GROWTH Before we examine the present status Much unnecessary confusion exists of the northern forest complex, we must today regarding these two terms. The consider the factors controlling the compo- concept of climax vegetation and in fact the sition and perpetuation of presettlement entire concept of succession, as originally forests. Simply because present forest defined by Clements and other early 20th communities are known to be largely the century ecologists, have been seriously result of human-caused disturbances, it questioned in recent years (Christensen and

54 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Peet 1984, Christensen 1987). Community development after a disturbance leads toward a more or less definable climax community controlled by regional climate (climatic climax). However, in many areas, due to topographic or soil influences, a regionally uniform climatic climax cannot be attained, except perhaps on a geologic time scale. Such terms as “topographic,” “edaphic,” or “topoedaphic climax” are commonly used to refer to the presumed “terminal” communities on such sites. Thus, in northern Wisconsin the climatic climax of sugar maple, hemlock, and beech can be expected only on more mesic, nutrient-rich soils. The droughtier and less thought of as structural and functional Old-growth community fertile sandy soils simply do not support of yellow birch and parts of larger landscapes. However, most hemlock. A recent tip- these demanding species; instead these soils studies have focused on vertebrate species up creates gap and are colonized by a number of shade- and vascular plants. Habitat needs of allows light to release intolerant or pioneer species. Because all of invertebrates and lower plants in relation to sugar maple seedlings. Photo by Michael J. the pioneer species on old growth are Mossman. sandy soils are shade- largely unknown. intolerant, they are Old-growth ecosystems may best be The alliance Lobarion incapable of replacing thought of as structural and functional pulmonaria, an themselves through association of rare parts of larger landscapes. advance regeneration, lichens, grows as is the case with primarily in old- mesic forest species. growth forests of Which, if any, species can be considered to northern hardwoods (Thompson 1990). represent the edaphic climax on the poorest The early survey records suggest that Table 3 soils is still not clear. Perhaps the answer to presettlement forests consisted of a mixture Generalized age this question is only of academic interest; of young, mature, and old forests. Old characteristics of old sooner or later a disturbance inevitably growth. Precise age forests were common in many areas, but varies with site-specific initiates a new cycle. successional processes were evident conditions. Based on “Old growth” is a much simpler (Lorimer and Frelich 1992). representative sites in concept than is climax vegetation. In some north-central Wisconsin. Compiled cases old growth may also be climax (e.g., a by R. Eckstein. 300-year-old sugar maple—hemlock community without a mixture of white pine), but most often it is simply a community with dominant trees at or near biologi- Age (years) cal maturity (Table 3). However, studies show that very old stands possess ecologi- Old Cover Type Individual cal properties that differ significantly from Cover Type Growth Begins Deteriorates Tree Longevity those of immature stands of the same floristic composition (Lorimer and Frelich 1992). However, although old growth Aspen 60 80 150 appears to provide optimal habitat for some Northern red oak 100 160 250 species of plants and animals, to date no White/red pine 130 200 400 vertebrate species have been shown to be obligate inhabitants of old growth. Thus, Northern hardwood 150 — 350 the old-growth ecosystems may best be Hemlock-yellow birch 150 — 500

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 55 NORTHERN FOREST COMMUNITIES

tionally, most of the reproduction was also eliminated. Hemlock was removed in a later wave of logging when the tanning industry, which used hemlock bark, was developed. Hardwoods were harvested last, after railroads and, later, logging roads were built. Both clearcutting and high-grading (i.e., cutting only the most valuable trees) were practiced. Because most hardwoods have less stringent requirements for germination and seedling establishment than do the pines and hemlock, and in addition possess sprouting ability, species such as sugar maple, beech, basswood, yellow birch, and ash were seldom eliminated Old-growth pine forest. An old tip-up creates THE LOGGING AND EURO-AMERICAN SETTLEMENT from a site unless there were repeated ﬁres. the coarse, woody However, the species composition of new ERA debris characteristic of stands was often severely altered. High- old-growth pine communities. Dunn Between the mid-1800s and early grading consistently favored sugar maple Lake Pines State 1900s Wisconsin forests were almost and beech, whereas clearcutting usually Natural Area. Photo by entirely cut over. The impact of logging and resulted in more mixed stands. Signe Holtz. associated activities was widespread and A large portion of presettlement forest varied. Space here does not permit a was later cleared for agriculture. Many comprehensive treatment of the ecological cleared lands proved unsuitable for farming consequences. Only those factors most and were abandoned. This was especially responsible for the differences between true for areas with sandier soils that origi- presettlement and current forest conditions nally supported conifers. Many of these are highlighted. lands were later planted back to trees, but Early logging concentrated on white often without regard for site potential and pine and, to some degree, on red pine. species compatibility. The successful Scattered trees as well as pure stands were farming that remains in northern Wiscon- The stump of an old, harvested wherever they were found. This sin is largely conﬁned to sites formerly large white pine (>36" dbh) within an even- had the immediate impact of virtually occupied by high-quality mesic hardwoods. aged stand of young eliminating the white pine seed source from sugar maples. Photo the northern hardwood complex. Because PRESENT by Michael J. slash was burned intentionally or uninten- Mossman. VEGETATION Both the species composition and relative proportion of presettlement forest types have been greatly altered by humans. The mixed coniferous-deciduous types have, with a few exceptions (e.g., the Menominee Indian Reservation), lost their coniferous component. Hemlock occurs sporadically in second-growth hardwood stands, but white pine is virtually absent in many areas and shows no signs of regeneration, even where suitable seedbed is created by natural or human-caused disturbance. The necessary supply of seed simply does not exist.

56 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE The relative importance of hardwood species has also changed signiﬁcantly in Bayfield many stands. While sugar maple has Douglas Ashland retained its dominant position, yellow birch Iron is much less common than it once was. On Vilas the other hand, basswood and white ash Washburn Sawyer Price Forest Florence are now usually the most important associ- Oneida Burnett ates of sugar maple, although they were Polk Marinette Barron Rusk seldom listed as such by early surveyors. Lincoln Langlade Taylor Oconto Most of the presettlement white pine Chippewa St. Croix Dunn forests (pure or mixed with red pine) are Marathon Menominee Clark today occupied by mixtures of red oak, red Shawano Pierce Eau Claire maple, white birch, and aspen, although Door Pepin Wood Portage Waupaca white pine is showing a remarkable come- Buffalo Outagamie Brown Jackson

back in many areas. Kewaunee Trempealeau By far the largest change has occurred Juneau Adams Waushara Winnebago Manitowoc Monroe in the distribution of the aspen-birch type. La Crosse Calumet Marquette Green While scarcely present on Finley’s map of Lake Sheboygan Vernon Fond Du Lac presettlement vegetation, today it repre- Sauk Columbia Dodge Richland Washing- sents the largest single forest cover type in ton Crawford Ozaukee the state. Much of it extends over the Dane landscape previously occupied by mesic Iowa Jefferson Waukesha hardwoods, indicating that the post-logging Grant Milwaukee Green Rock Lafayette Walworth Racine fires also occurred in these communities. MILES Kenosha Considering the northern forest 010203040 region as a whole, the overall species industry land ranges from less than 1% to Figure 13 richness of plants and animals does not 25% of total land area in northern forest appear to be threatened. Probably few if The continuous, counties. Of commercial forest land, maple- extensive forest of any species of flora have been lost, al- birch makes up 31%, aspen 29%, elm-ash- northern Wisconsin, though relative abundance of many has adapted from soft- maple 7%, been greatly altered. McCaffery and Creed paper birch 5%, oak Figure 13 shows the (1969). 5%, and balsam fir extent of the largely Considering the northern forest region as 5%.3 White cedar, continuous northern a whole, the overall species richness of black spruce, white forest in 1992. The plants and animals does not appear to be spruce, white pine, northern forest threatened. Probably few if any species of red pine, and jack includes parts of 19 pine forest types each counties. The total flora have been lost, although relative make up less than 5% land area in forest abundance of many has been greatly of commercial forest cover ranges from altered. land. Non-stocked 59% to 93% on a land makes up 1% of countywide basis. commercial forest land (Spencer et al. There are 8.3 million acres of com- 1988). mercial forest land in the northern forest.2 The northern forest is characterized Public land totals 3.5 million acres and by a sapling and pole-sized forest. Seed- ranges from 17% to 56% of the total land lings-saplings range from 6% of commercial in northern forest counties. The forest forest land in Menominee County to 38% industry owns 987 thousand acres; forest of commercial forest land in Oneida 2 Commercial land is defined as land producing or capable of producing crops of industrial wood 3 Maple-birch is a loosely labeled type that and not withdrawn from timber production (Spencer includes sugar maple, yellow birch, basswood, white et al. 1988). ash, and hemlock.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 57 NORTHERN FOREST COMMUNITIES

County. Pole timber makes up 49% of inferior genetic stock, row planting, fur- commercial forest land. Sawtimber stands rowing, destruction of humus layer, and do not predominate. This reflects the elimination of ground vegetation caused continual rebuilding and maturation of ecological problems. stands from the cut-over and burned-over The growing-stock volume of nearly conditions at the turn of the century. It also every tree species increased between 1968 reflects the domination of pole and small and 1983, except for elm, hemlock, and sawtimber-size material for timber products yellow birch, for which volumes declined. as the primary objective of forest manage- An average of 21.6 million board feet of ment on several million acres of forest land. hemlock and 14.7 million board feet of Sawtimber area and volume decreased from yellow birch were removed annually from 1936 to 1956, then began increasing from commercial forest land in northern Wiscon- 1956 to 1983 (Stone and Thorne 1961, sin. These tree species exhibit very low Spencer and Thorne 1972, Smith 1986). regeneration rates (Raile 1985). Pine plantations cover 355,000 acres Between 1964 and 1983, 12% of in northern Wisconsin. Red pine makes up commercial forest land was harvested. Of Table 4 61% of the total, and jack pine makes up the 12% harvested, 72% had a partial cut Changes in the relative 22% of the total (Smith 1986). Between and 28% was clearcut. All past and current abundance and 1956 and 1968, 500,000 acres were logging practices change forest communi- distribution of selected planted in all of Wisconsin. Most acres ties. In addition, introduced insects and wildlife in Wisconsin’s were planted to red pine (Spencer and diseases, such as dutch elm disease and white northern forests: 1850Ð1994. Compiled Thorne 1972), but red pine was often pine blister rust, have significantly altered the by R. Eckstein. inappropriate for the site. In addition, composition of post-settlement forests.

Relative Abundance Distribution

Mid- Early Mid- Mid- Early Mid- Species 1800s 1900s 1900s 1994 1800s 1900s 1900s 1994

White-tailed deer Low Low Abundant Common Clumpy Clumpy Continuous Continuous Coyote Low Common Abundant Common Clumpy Clumpy Continuous Continuous Bobcat Low Low Common Rare Clumpy Clumpy Continuous Clumpy Moose Low Rare Gone Rare Clumpy Isolated Gone Isolated Snowshoe hare Low Common Abundant Low Clumpy Continuous Continuous Continuous Timber wolf Common Common Gone Rare Continuous Continuous Gone Clumpy Fisher Common Rare Gone Common Continuous Isolated Gone Continuous Pine marten Abundant Rare Gone Rare Continuous Isolated Gone Isolated Elk, wolverine Low Gone Gone Gone Clumpy Gone Gone Gone Bald eagle, osprey Common Common Low Common Common Continuous Clumpy Continuous Ruffed grouse Low Common Abundant Common Clumpy Continuous Continuous Continuous Woodcock Low Common Abundant Common Clumpy Clumpy Continuous Clumpy Sharp-tailed grouse Low Abundant Common Rare Clumpy Continuous Clumpy Isolated Beaver Common Rare Low Abundant Continuous Isolated Clumpy Continuous Grassland birds Rare Common Common Rare Isolated Continuous Clumpy Isolated Interior forest birds Abundant Rare Rare Common Common Continuous Clumpy Continuous Young-forest birds Rare Common Common Common Isolated Clumpy Continuous Continuous

58 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE ANIMALS Benyus et al. (1992) compiled a list of 389 vertebrate species present in the northern forests of Michigan, Minnesota, and Wisconsin. The list included birds (71%), mammals (17%), and reptiles/ amphibians (12%). Fifty-three percent of these species were uncommon, 37% common, and 10% occasional. These species used all kinds of habitats in all successional stages. Of the forest species, 49% used mature forest and 40% used young forest. Thirty-three species were classified as highly versatile in habitat use, while 204 species had intermediate versatil- ity and 152 species were restricted to specific habitats. agriculture provided favorable habitat. In The fisher was once The distribution and abundance of extirpated from the recent years, despite maturing forests, northern forest but has animals in the northern forest have badgers have become established in low recovered after changed dramatically (Table 4). Among numbers throughout the northern forest. reintroduction. DNR mammals, unregulated commercial hunting Presettlement white-tailed deer photo and trapping as well as dramatic habitat populations ranged from five to 15 deer per changes has resulted in extirpation of elk, square mile (Dahlberg and Guettinger wolverine, woodland caribou, Canada lynx, 1956, Habeck and Curtis 1959). Deer fisher, pine marten, moose, eastern cougar, occurred at very low numbers between and eastern timber wolf. In recent years 1900 and 1915 but then began to increase Lack of large fisher, pine marten, and eastern timber wolf (Swift 1946). Abundant favorable habitat blocks of wild land have been reestablished, and eastern cougar caused populations of white-tailed deer and with low human and moose occur in very low numbers. snowshoe hare to grow to very high num- Lack of large blocks of wild land with bers. Snowshoe hare populations peaked in presence limits low human presence limits populations of the early 1930s and were again very high in populations of some animal species, e.g., eastern timber the 1940s (Cunningham 1993). White- some animal wolf (Thiel 1985), black bear, bobcat, tailed deer populations peaked in the species. moose, eastern cougar and spruce grouse. 1940s with 40 to 50 deer per square mile These species are known as extensive forest of deer range (Keith McCaffery, Dep. of specialists. These are usually large, wide- Natural Resources, pers. comm.). These ranging, or sensitive animals. The forest very high deer and hare populations caused need not be mature and can be intensively widespread damage to vegetation. managed. However, it must have low Current Deer Management Unit human presence. population goals reflect current forest Other mammal species dropped to habitat conditions. Management Unit goals very low numbers when logging and Euro- in the northern forest average 18 (range ten American settlement drastically altered to 25) deer per square mile of deer range. their habitat, then increased as the forest Snowshoe hare populations are currently began to mature again. These include gray low because of widespread predation, squirrel, porcupine, flying squirrel, and particularly by fisher. The impact of white- beaver. Still other species, such as raccoon, tailed deer and snowshoe hare on the striped skunk, woodchuck, thirteen-lined composition and structure of forests needs ground squirrel, and eastern cottontail, to be viewed on a broad temporal and became much more abundant as young spatial scale (Mladenhoff and Stearns forests, edge, resorts, small towns, and 1993).

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 59 NORTHERN FOREST COMMUNITIES

Many of the human and ecological In the past, forest birds adapted to forces that impacted mammal species also large blocks of mature forest decreased in affected bird species. Habitat changes and numbers as these forests were converted to unregulated commercial hunting extirpated brushland. Examples include the barred the passenger pigeon from Wisconsin; the owl, pine warbler, Blackburnian warbler, species became extinct in 1914. Although black-throated blue warbler, yellow-bellied common in presettlement and early Euro- sapsucker, pileated woodpecker, eastern American settlement times in Wisconsin, wood-pewee, Swainson’s thrush, wood bald eagles, osprey, and Cooper’s hawks thrush, solitary vireo, cerulean warbler, and dropped to low numbers by the mid-1900s scarlet tanager. The northern hardwood because of indiscriminate shooting and component of the northern forest is recov- reproductive failure caused by pesticides. ering (Stearns 1990) but occurs in smaller In the early 1900s, red-shouldered hawks blocks (Mladenhoff et al. 1993). It now declined as mature lowland deciduous averages 70 years of age, is developing an forests declined. Extensive logging, fire, all-aged structure, and again supports and scattered agriculture created favorable populations of mature forest birds habitat for species such as sharp-tailed (Hoffman 1989). grouse, upland Forest practices sandpiper, eastern can negatively affect bluebird, American Many of Wisconsin’s amphibian and some species of forest goldfinch, golden- reptile species are found throughout the birds (Temple 1988, winged warbler, state, often in wetlands located within Howe et al. 1992). American crow, gray other vegetative communities. However, However, properly catbird, northern some species with highly specific habitat modified forest harrier, red-tailed practices, in the requirements are found only in the hawk and American context of the exten- kestrel. These species extensive northern forests . sive northern forest, are now declining as can enhance habitats the forests have for forest birds grown back and are maturing. (Temple et al. 1979, Hoffman and Species that are adapted to young or Mossman 1990, DeGraaf et al. 1992, Probst disturbed forests have increased as this et al. 1992, Thompson et al. 1992, DeGraaf successional stage has increased. These et al. 1993, Thompson et al. 1993, Welsh species include ruffed grouse, woodcock, and Healy 1993). chestnut-sided warbler, mourning warbler, Forest ponds are breeding habitat for blue jay, rufous-sided towhee, brown many species of frogs and salamanders. thrasher, Nashville warbler, indigo bunting, Abundant decaying logs on the forest floor rose-breasted grosbeak, and great horned as well as an uncompacted forest floor litter owl. layer are important habitats for salamanders One bird species, the brown-headed and invertebrates. Many of Wisconsin’s cowbird, has increased dramatically in the amphibian and reptile species are found eastern United States and in southern throughout the state, often in wetlands Wisconsin. In agricultural areas this nest present within other vegetative communi- parasite can cause forest bird species to ties. However, some species with highly decline. In the northern forest, cowbirds specific habitat requirements are found are uncommon but present in local agricul- only in the extensive northern forests (Vogt tural areas and near towns. In the forested 1981). Examples are the mink frog, red- environment, cowbirds are present in first- backed salamander, and spotted sala- year aspen clearcuts, young conifer planta- mander. Other species such as the wood tions, and large grassy openings. The frog, northern red-bellied snake, and wood impact of cowbirds on northern forest bird turtle are most common in the northern populations is unknown. forest but occur elsewhere in Wisconsin as

60 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE well. Because no thorough inventories have Portions of current aspen-birch type will been conducted for Wisconsin’s reptiles and be replaced by various mixtures of white amphibians, we have no basis to compare pine, red maple, and occasionally red current distribution and abundance with oak. A significant proportion will that of the past. succeed to mixed stands of mesic Except for pest species, little research hardwoods, with sugar maple playing has been directed at forest invertebrates. the largest role. Lack of knowledge in this area is a serious concern since invertebrates are a very All forests currently dominated by mesic diverse group and perform important hardwoods will remain so, but species ecosystem functions. composition will vary greatly depending on geographic location, site type, and management practices. Sugar maple will THREATENED AND ENDANGERED SPECIES become more dominant on many mesic Threatened and endangered bird and sites. mammal species that have a significant portion of their range in the northern forest The acreage of red pine plantations is include eastern timber wolf, Canada lynx, likely to dominate local areas, particu- pine marten, bald eagle, osprey, red- larly on forest industry lands. Jack pine shouldered hawk, and wood turtle. Threat- acreage is decreasing. Most is going to ened and endangered plants of the north- red pine plantations. ern forest include moonwort, goblin fern, Smith melic grass, pine-drops, small Because of great disparity between shinleaf, foamflower, calypso orchid, ram’s economic and biological maturity of head lady’s-slipper, small round-leaved most tree species, the increase of old- orchid, Braun’s holly fern, drooping sedge, growth forests, in a biological sense, is auricled twayblade, broad-leaved tway- unlikely. Increased utilization prevents blade, and hawthorn-leaved gooseberry. development of old-growth characteristics in managed mature forests. PROJECTED Clearcuts and plantations will continue Projections of future dynamics of to fragment large, uniform blocks of Wisconsin’s forests are difficult to make mature mesic hardwoods. Temporary without a knowledge of future management edges caused by forest cutting will or utilization objectives of a changing continue to dominate the northern society. Barring major changes in forest landscape. ownership and resource utilization policies, the following trends can be expected: Small, permanent grassy openings will The total forested area will probably continue to decline to less than 1% of remain the same or increase slightly. public and forest industry lands. Wild- life dependent on grassy, open areas will The aspen-birch type will gradually decline (McCaffrey and Creed 1969). decrease as forest succession progresses. The area in aspen has declined 1.8 Balsam fir and tag alder will continue to million acres since 1936 (Spencer et al. dominate the former white cedar forests. 1988). Aspen stands today are perpetu- White cedar and Canada yew reproduc- ated almost entirely by commercial tion will be restricted to scattered, local clearcutting. Current utilization is not areas. keeping up with the rapid maturation rate of this short-lived species.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 61 NORTHERN FOREST COMMUNITIES The major forest The scattered relict stands containing result is a mosaic of many small stands of cover types of the hemlock and yellow birch will continue widely different age classes. Temporary to decline. Reproduction of these species edges are abundant. Large blocks of unbro- northern forest are will be restricted to scattered, local areas ken mature mesic forest remain rare. Fire as managed at an (Eckstein 1980). a natural process is rare and is not currently economic rotation used as a management tool in most areas. age. Old-growth Fire will not play a significant role as an There is pressure by hunters to raise forests and old- ecological agent in the northern forest. white-tailed deer population goals. Some growth plants such as Canada yew, hemlock Road networks will continue to be saplings, and some orchids are quite characteristics in improved and expanded. Currently, 46% sensitive to deer and snowshoe hare managed forests of the northern commercial forest is herbivory and decline with locally high 1 do not develop. within /4 mile of an improved road deer and hare populations. Only selected (Smith 1986). Road networks are improving and economic tree expanding so that they dominate the species, a few The demand for forest products such as landscape in most areas. Housing and forest game pulpwood, sawlogs, white-tailed deer, recreation interests are developing in more ruffed grouse, characteristics such as and more wild land, particularly in Oneida, species, and wild country and solitude will continue Sawyer, Vilas, and Washburn counties. selected to increase. Large blocks of undeveloped country are endangered or declining throughout the northern forest. threatened species State and county agencies currently receive funding use no regional landscape overview and do ACTIONS CAUSING CONCERN and management not utilize a unified regional classification system. Forests are managed on a stand-by- attention. The The major forest cover types of the stand basis with a bottom-up forest recon- result is a mosaic northern forest are managed at an economic rotation age. This perpetuates a naissance system. There is little consider- of many small simpler local and regional age structure of ation of forest patterns and processes using stands of widely forest communities. Old-growth forests and a top-down regional landscape approach. different age old-growth characteristics in managed In many cases economic rather than classes. forests do not develop. More intensively ecological decisions determine management managed forests lack the snag and den-tree direction. National, state, county, and local component as well as the horizontal and public land units plan management strate- vertical structure typical of old-growth gies independently. stands. Only selected economic tree species, a few forest game species, and selected SOCIO-ECONOMIC ISSUES endangered or threatened species receive In the recent past the forest was used funding and management attention. The primarily as a source of wood products. With the exception of a few periods when Some orchids are quite sensitive to deer and there was some concern over the diminish- snowshoe hare ing forest resource, the public was generally herbivory and decline unconcerned about the treatment of forests. with locally high deer and hare populations. Biological resources not conspicuously Showy ladyslipper. related to timber were largely unrecog- Photo by Staber nized. Reese. These attitudes have changed greatly in recent years. Conflicts between traditional uses of forests, recreational demands, and concerns for natural ecosystem preservation are intensifying. While all factions

62 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE agree that each has valid concerns, agreements on the future use of forest resources are becoming more and more difficult to reach. Although the public is better educated about environmental issues than it has been in the past, numerous misconceptions about the nature of forest ecosystems persist. Many see any disturbance, particularly fire and clearcutting, as unnatural and always detrimental. The process of change through natural succession is seldom appreciated. There are numerous attempts at “preserving” community types that are successional. The hands-off approach is often considered as the only solution to many problems, even though indirect refers to all age classes from seedling Sugar maple stand with a history of effects of humans are most often present through old growth. These successional logging. Heavy sapling (e.g., introduced insects and diseases, stages should occur in all stand sizes from layer shades out exotic plant species, air pollution, acid small 40-acre stands to large 2,000-acre ground layer except for stands. maple seedlings. deposition, exclusion Photo by Michael J. of fire, etc.). Public lands Mossman. Development of Although the public is better educated occur across the ecologically sound, entire northern forest about environmental issues than it has cost-effective tech- on all major land- niques encouraging been in the past, numerous forms and soil types. natural processes on misconceptions about the nature of forest The distribution and the forest landscape ecosystems persist. abundance of public will require partner- lands present an ships with the forest opportunity to meet landowners, including the forest industry. multiple objectives on a landscape scale. Public pressure to pay more attention to Different landscape objectives can be met maintaining complete and functional forest on different public land ownerships, ecosystems will surely continue. depending on the degree of cooperation among agencies. For example, large unmanaged tracts could (and do) occur on POTENTIAL FOR COMMUNITY National Forests, smaller unmanaged tracts on state forests, and small natural-area- RESTORATION sized tracts on county forests. A regional There is great potential for maintain- landscape approach can incorporate ing and enhancing biodiversity in the management of some forest ecosystems to northern forest. The basic elements of the feature certain species such as white-tailed conservation of biodiversity in forests deer and ruffed grouse while managing include tree species composition, stand age, other forest ecosystems for plants and stand structure, and stand area. The key is animals that require large blocks of mature to use a landscape management approach forest or old-growth forest. The challenge is that accounts for all the characteristic determining what agency does what, how successional stages with forest stands much, and where. ranging from small to very large (Hunter We suggest that the record of 1990, Crow 1991, Probst and Crow 1991, presettlement vegetation be used as an aid Freemark et al. 1993, Haila et al. 1994) but not an absolute model for determining (Fig. 14). Characteristic successional stage the “desired state” of forest vegetation in a

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 63 NORTHERN FOREST COMMUNITIES

particular area. There are numerous reasons Resources Board for consideration begin- for this. Most importantly, the ning in the 1995-97 biennium.. presettlement vegetation, as reconstructed 1. Facilitate inter-agency cooperation by from survey records, represents communi- creating a Northern Forest Working ties based only on dominant tree species Group. This Working Group would present in a particular time period. Subse- coordinate information exchange among quent studies, based on hundreds of the various agencies and groups manag- stands, clearly show that forest communi- ing the northern forest community. The ties sharing common dominants often Working Group could act as a clearing- exhibit significantly different floristic house for information and could facili- compositions when entire floras are com- tate coordinated landscape planning. pared (Kotar 1987). Similar differences also For example, meetings have begun exist in productivity, rates of succession, between USDA Forest Service, the associated animal species, and perhaps in Department, and the County Forest other ecological conditions not yet studied. Association collaborating on research The forest habitat type classification and information-sharing. system (Kotar et al. 1988) is another tool that can be used for assessing the desired 2. Encourage the integration of the plan- state of vegetation on different sites and ning and management functions within especially for evaluating the potential for each of the land management agencies restoring a chosen condition. in Wisconsin. All featured-species forest management guidelines (including forest game, forest vegetation, and endangered, POSSIBLE ACTIONS threatened, and nongame species) and The following possible actions are all new ecosystem management guide- consistent with ecosystem management, lines should be integrated into one but require more analysis and discussion. handbook. Figure 14 How priorities are set within this list will be based on ecoregion goals, staff workload, 3. Encourage inclusion of ecosystem Framework for fiscal resources, public input and support, management elements in the Managed application of a Forest Act. Develop guidelines for landscape approach and legal authority. We will work with our within ecosystem customers and clients to set priorities and private landowners to enhance management. bring recommendations to the Natural biodiversity. Local diversity could be maintained and improved by developing

A Landscape All species and Need for Compositional, Management natural processes community or structural, and Emphasis are important ecosystem functional management diversity means ➤ which implies ➤ (not just commodity species) by maintaining ➤ to enhance ➤

64 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE guidelines for snags, den trees, old how the various public properties can growth, sensitive habitats, protection of be managed to meet local, regional, hemlock and white cedar, reserve trees, and national objectives. Examples and extended rotations. include county forest lands, Depart- ment managed lands, lands managed 4. Plan and manage public lands using a by the Board of Commissioners of landscape-scale ecosystem approach. Public Lands (School Trust Lands) Use a top-down hierarchical approach to and National Forest Lands. Protect plan management across large landscape the unique biological, scientific, ecosystems (Noss 1983, 1992; aesthetic, and educational opportuni- Mladenhoff and Pastor 1993; Bailey et ties on these lands. al. 1994) (Fig. 15). Continue implementation of a system Implement the Forest Accord by of designated natural areas that using the National Hierarchy of represent the full spectrum of bio- Ecological Units (Bailey et al. 1994) logical communities across the combined with the Habitat Classifica- northern forest. tion System (Kotar et al. 1988) to the greatest extent possible. Ecologically 5. Develop an old-growth policy for state based maps would provide informa- land and encourage the application for tion on spatial patterns and interac- old growth on county land. tions of landform, soils, climate, cover types, and potential natural Develop operational definitions of vegetation (Albert 1992, 1993; Bailey old growth for Department managed et al. 1994). Landscape-scale ecosys- lands. tem mapping must be coordinated between agencies and landowners Defer cutting of existing old growth across the northern forest so termi- on state land until an old-growth nology and techniques are consistent. policy is established.

Determine how the various public Old-growth areas in the northern lands ﬁt into regional and large forest must be large enough to meet landscape ecosystems. Then, based compositional, structural, and on the type of public land, identify functional objectives (Vora 1994).

A range of A range of patch Forest habitat Regional developmental sizes type Landscape stages of (from small areas to in the ➤ communities very large blocks) (successional as on each ➤ well as mature) in ➤

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 65 NORTHERN FOREST COMMUNITIES

Current literature suggests that there animals. This recommendation applies are three factors that contribute to to all upland forest types. Landscape the effective size of an old-growth planning can help determine the best patch: (1) actual size (with a mini- opportunities to reduce edge and mum area constraint), (2) distance increase forest interior conditions. from similar old-growth patches, and (3) degree of habitat contrast of 7. Continue to improve structure and intervening forest (Harris 1984, composition in managed forests. Vankat et al. 1991, Vora 1994). Apply big-tree silviculture methods, a A way to enhance each old-growth system originally designed to achieve patch’s effective area is to surround aesthetic objectives, on state and each with a mature forest zone county forests. Big-tree silviculture is managed with single-tree or group a powerful tool to enhance diversity selection methods (Mladenhoff et al. within and between stands. 1994). The approach of imbedding old-growth in mature forest zones Based on a landscape analysis, serves to enhance composition, determine the need to extend the structure, and function. Efforts economic rotation for some even- should be made to link old-growth aged stands. patches through use of riparian zones, aesthetic zones, or natural Develop guidelines for structural and areas. compositional characteristics in managed stands. These include large- 6. Increase relative stand size to reduce diameter trees, supercanopy trees, edge and increase forest interior condi- large standing snags, mast trees, large tions. Patch (stand) size is smaller in den trees, and large downed trees. today’s forests compared to presettlement forests (Mladenhoff et al. Continue developing guidelines for 1993). In the context of the extensive sensitive habitats such as riparian Figure 15 forest, stands with sizes of 200 to 2,000 zones, rare-plant zones, and sensi- acres tend to develop interior conditions tive-soil zones. Decision framework for favored by a variety of plants and managing on a landscape scale.

Wisconsin’s Five forest Landscape Forest habitat Forest cover northern habitat type ecosystem types types (i.e., forest regions units occupied successional stages) on divided divided on which by ➤ public lands into ➤ into ➤ occur ➤ These cover types provide ➤

66 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Develop prescriptions to maintain Aerial view of small Iron County farms tree species diversity. Maintain fragmenting the hardwoods such as oak and red northern forest. Photo maple in pine stands and intolerant by Michael J. hardwoods such as basswood, yellow Mossman. birch, and white ash in sugar maple stands.

Protect and enhance relict stands of hemlock and white cedar. Enhance- ment of these stands for species regeneration may require active or non-active management scenarios.

8. Analyze road densities and develop policies for roads on Department managed lands; encourage consideration of this issue on state and county lands. Use landscape scale units to analyze logging road distribution, quality, and abundance. Reduce road densities to protect sensitive plant and animal species and sensitive areas.

Young Forest with ➤ to benefit ➤ Stands of many sizes composed mostly of Deer and other forest game and associated intolerant types including forest openings species

to Mature Forest with ➤ to benefit ➤ Stands of many sizes composed mostly of Interior and old growth specialists and tolerant types including old growth associated species

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 67 NORTHERN FOREST COMMUNITIES

Case Study

MARATHON COUNTY FOREST: USING GIS TO MANAGE FOREST INTERIOR HABITAT

Contributed by Mark Heyde, Ron Eckstein, and Becky Isenring.

Marathon County owns and manages a 26,747-acre public forest made up of many tracts, mainly concentrated in three areas of the county. Each tract, large and small, is imbedded in a matrix of agricultural and private forest lands. Although this county is on the southern edge of the range of northern forest, it provides a good example of how ecosystem management principles help us address issues of biodiversity across the northern forest. Populations of a large group of songbirds, neotropical migrants, are in decline world- wide. Although it is not clear which of several factors are most responsible for their decline, the Marathon County Forest wanted to do what they could to contribute to the long-term viability of neotropical migrant bird numbers. Some evidence suggests that nests in small forest blocks are susceptible to high rates of parasitism, predation, and competition from species that tolerate edge habitat. In general, small forested tracts situated in agricultural landscapes provide little habitat suitable for species that are dependant upon forest interior conditions. Marathon County decided to try to address the needs of the neotropical migrants using a Geographical Information System (GIS). They are using the GIS to analyze the county forest, generating an overview of forest stand types, sizes, and ages within the context of the Marathon County landscape. The GIS is queried for the location of possible and potential interior-forest bird habitat, using guidelines from research in the Hoosier National Forest in Indiana that were adapted to reflect conditions on the southern front of Wisconsin’s northern forest. For example, edge was defined in terms of forest stand structure, size of forest openings, location of roads, and the location of nearby agricultural fields. These parameters, applied to GIS map layers, are being used to design a forest management system that reduces edge effects and enhances the area of interior forest habitat. Marathon County is using a hierarchical approach to look at multiple scales of space and time in planning and designing management activities. The manager considers where the Marathon County Forest is located in the state while considering the position of individual county forest parcels and their composition. With this broad array of information at hand, the manager can lay out a variety of possible future conditions for the Marathon County Forest. In the planning, a wide range of options can be considered, including those that benefit interior forest songbirds.

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Hoffman, R.M. 1989. Birds of Wisconsin Mladenhoff, D.J. and J. Pastor. 1993. northern mesic forests. Passenger Sustainable forest ecosystems in the Pigeon 51(1):97-110. northern hardwood and conifer forest Hoffman, R.M. and M.J. Mossman. 1990. region: concepts and management. Birds of northern Wisconsin pine pp.145-180. In: G.H. Aplet, N. forests. Passenger Pigeon 52(4):339- Johnson, J.T. Olson and V.A. Sample, 355. eds. Defining sustainable forestry. Island Press. Washington, D.C. 328pp. Howe, R.W., M.J. Mossman and S.A. Temple. 1992. Effects of natural forest Mladenhoff, D.J., M.A. White, T.R. Crow, management on birds in northern and J. Pastor. 1994. Applying principles Wisconsin. Passenger Pigeon of landscape design and management to 54(4):297-305. integrate old-growth forest enhancement and commodity use. Hunter, M. L., Jr. 1990. Wildlife, forests, Conservation Biology 8(3):752-762. and forestry; principles of managing forests for biological diversity. Prentice- Mladenhoff, D.J., M.A. White, J. Pastor and Hall, Englewood Cliffs, New Jersey. 370 T.R. Crow. 1993. Comparing spatial pp. pattern in unaltered old-growth and disturbed forest landscapes for Knapp, J. G. 1871. The native vegetation of biodiversity design and management. Wisconsin. Trans. Wis. State Hort. Soc. Ecological Applications 3:293-305. 1:119-25. Noss, R.F. 1983. A regional landscape Kotar, J. 1987. The role of phytosociology approach to maintain diversity. in forest site classification in the Great BioScience 33:700-706. Lakes region. pp. 76-80 D. W. Cole and S. P. Gessel, eds. Forest site evaluation Noss, R.F. 1992. Issues of scale in and long-term productivity. Univ. Wash. conservation biology. pp.139-250. In: Press, Seattle. P.L. Fiedler and S.K. Jain, eds. Conservation biology. Chapman and Kotar, J., J. A. Kovach, and C. T. Locey. Hall. New York. 507pp. 1988. Field guide to forest habitat types of northern Wisconsin. Univ. Wis.- Probst, J.R. and T.R. Crow. 1991. Madison Dep. of For. and Wis. Dep. Integrating biological diversity and Nat. Resour. 212 pp. resource management. J. Forestry. 89:12-17. Lorimer, C. G., L. E. Frelich, and E. V. Nordheim. 1988. Estimating gap origin Probst, J.R., D.S. Rakstad and J. Rugg. probabilities for canopy trees. Ecology 1992. Breeding bird communities in 69:778-85. regenerating and mature broadleaf forests in the USA Lake States. Forest Lorimer C. G. and L. E. Frelich. Natural Ecology & Management 49:43-60. disturbance and eastern old growth. J. For. In press Raile, G. K. 1985. Wisconsin forest statistics, 1983. U.S. For. Serv. N. Cent. McCaffery, K. R. and W. A. Creed. 1969. For. Exp. Stn. Res. Bull. MC-94. 113 Significance of forest openings to deer pp. in northern Wisconsin. Wis. Dep. Nat. Resour. Tech. Bull. No. 44. 104 pp. Roth, F. 1898. On forestry conditions of northern Wisconsin. Wis. Geol. and Mladenhoff, D.J. and F. Stearns. 1993. Nat. Hist. Surv. Bull. 1. Madison Eastern hemlock regeneration and deer browsing in the northern Great Lakes Smith, W.B. 1986. Wisconsin’s fourth forest region: a reexamination and model inventory: area. USDA. For. Serv. simulation. Conservation Biology Research Bull. NC-97. 48 pp. 7(4):889-900.

70 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Spencer, J. S., Jr., and H. W. Thorne. 1972. Thompson, F.R., W.D. Dijak, T.G. Kulowiec Wisconsin’s 1968 timber resource—a and D.A. Hamilton. 1992. Effects of perspective. U.S.D.A. For. Serv. N. even-aged management on breeding Cent. For. Exp. Stn. Res. Bull. NC-15. bird densities. J. Wildl. Manage. 80 pp. 56(1):23-30. Spencer, J. S., Jr., W. B. Smith, J. T. Hahn Thompson, J.W. 1990. Lichens in old- and G. K. Raile. 1988. Wisconsin’s growth woods in Wisconsin. The fourth forest inventory, 1983. U.S.D.A. Bulletin. The Botanical Club of For. Serv. N. Cent. For. Exp. Stn. Res. Wisconsin. 22(1):7-10. Bull. NC-108. 158 pp. Vankat, J.L., J. Wu and S.A. Fore. 1991. Stearns, F. 1990. Forest history and Old-growth forests by design: applying management in the northern Midwest. the concepts of landscape ecology. pp.107-122. In: J.M. Sweener, ed. pp.153-169. In: D. Henderson and L.D. Management of dynamic ecosystems. Hedrick, eds. Restoration of old-growth North Central Section. The Wildlife forests in the interior highlands of Society. West Lafayette, IN. 180pp. Arkansas and Oklahoma. Proceedings Stone, R. N. and H. W. Thorne. 1961. of the Conf. Morrilton, AR. Sept 19-20, Wisconsin’s forest resources. U.S.D.A. 1990. For. Serv. Lakes States For. Exp. Stn. 52 Vogt, R. C. 1981. Natural history of pp. amphibians and reptiles of Wisconsin. Swift, E. 1946. A history of Wisconsin deer. Milwaukee Publ. Mus. 205 pp. Wisconsin Conservation Department. Vora, R.S. 1994. Integrating old-growth Pub. 323. Madison. 96pp. forest into managed landscapes: a Temple, S.A. 1988. When is bird habitat northern Great Lakes perspective. not habitat? Passenger Pigeon 50(1):37- Natural Areas Journal 14:113-123. 41. Ward, R.T. 1956. The beech forests of Temple, S.A., M.J. Hoffman and B. Ambuel. Wisconsin-changes in forest 1979. The ecology and management of composition and the nature of the avian communities in mixed hardwood- beech border. Ecology 37:407-419. coniferous forests. pp.132-153. In: R. Warden, J. A. 1881. Forests and forestry in Degraaf and R. Evans, eds. Management Wisconsin. Trans. Wis. State Hort. Soc. of north-central and northeastern 11:143-56. forests for nongame birds. USDA. For. Welsh, C.J. and W.M. Healy. 1993. Effect of Serv. Gen. Tech. Rep. NC-51. even-aged timber management on bird Thiel, R.P. 1985. The relationship between species diversity and composition in road densities and wolf habitat in northern hardwoods of New Wisconsin. American Midland Hampshire. Wildl. Soc. Bull. 21: 143- Naturalist 113:404-407. 154. Thompson, F.R., J.R. Probst and M.G. Raphael. 1993. Silvicultural options for neotropical migratory birds. pp.353- 362. In: D.M. Finch and P.W. Stangel, eds. Status and management of neotropical migratory birds. USDA. For. Serv. Gen. Tech. Report RM-229. 422pp.

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ecological importance of several species of oak (red, white, black, bur, northern pin, and swamp white) and by the presence of several tree species normally not found CHAPTER 5 north of the tension zone (shagbark hickory, hackberry, boxelder, and black walnut). Bitternut hickory and butternut, which occur only occasionally in northern forests, are common although not abundant Southern in the south. Equally important is the general absence of conifers (white spruce, balsam fir, hemlock). Pines, especially jack pine, occur in scattered areas of sandy soils. Forest Curtis (1959) classified southern Wisconsin forests into five community types, based on existing overstory composition: wet, wet-mesic, mesic, dry-mesic, and Communities dry. Only the upland types (mesic, dry- mesic, and dry) will be considered here. by John Kotar Mesic forests are characterized by the Department of Forestry dominance of so-called mesic hardwoods, University of Wisconsin-Madison mainly sugar maple, basswood, and Ameri- and can beech in the extreme eastern part of the Paul Matthiae state. Ironwood, American elm, and white Bureau of Endangered Resources ash are common associates. Dry-mesic Department of Natural Resources forests are dominated by red and white oak, and dry forests are dominated by black, white, and bur oak. Hickories are common associates in both dry-mesic and dry types, while mesic hardwoods are frequently present as less important associates in dry-mesic types. A system of finer divisions into DESCRIPTION “habitat types” based on ground layer and shrub species as well as canopy species has escribing forests in southern been developed by Kotar et al. (1988). The The southern Wisconsin (south of the system has been valuable in developing forest is tension zone) is more difficult forest management plans for specific sites. contrasted with than describing forests in This system takes into account the fact that northern Wisconsin. One the northern all tree species have wide ecological ampli- complication is that this region forest by the tudes and often occur as temporary domi- includes both glaciated and unglaciated nants on sites where they do not maintain ecological landforms, which together include soils themselves in competition with other importance of that range in age from 15,000 to perhaps species. For example, stands dominated by several species of 750,000 years. Another complicating factor red and white oak may occur as a result of oak and by the is the important role of fire during a period fire disturbance on both mesic and dry- lasting perhaps 5,000 years, up to the time presence of mesic sites. However, the associated flora of Euro-American settlement, with the peak on the two sites will differ significantly. several tree of this xerothermic period occurring 3,500 species normally Thus the two communities are less alike years ago. than their canopy composition would not found north of In broadest terms, the southern forest is suggest. Also, with the exclusion of fire, the the tension zone contrasted with the northern forest by the oak community on the mesic site will

72 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE rapidly succeed to mesic dominants, Wisconsin’s landscape is covered by varying whereas the community on the dry-mesic depths of wind-blown silt (loess), originat- site will either remain as an oak-dominated ing in the Mississippi floodplains. Thus the Current differences community or will composition of the soil in the species succeed to mesic parent material and composition of species over a much age of soils in glaciated longer period. Because the function of an ecosystem is and driftless regions communities in Because the dependent on both its biotic and abiotic differ less than would similar function of an makeup, it is important to distinguish be expected. environments are ecosystem is depen- between communities with different overall Current floristic presumed to be dent on both its species composition. In other words, two distribution in the due to differences biotic and abiotic state suggests that communities dominated by the same trees in disturbance makeup, it is enough time has species may be functionally very different. important to elapsed since the histories and distinguish between retreat of the continen- chance events. communities with tal ice sheet for most different overall species composition. In plants and animals to reach suitable habi- other words, two communities dominated tats. Current differences in species compo- by the same trees species may be function- sition of communities in similar environ- ally very different. If biological diversity is ments are presumed to be due to differ- to be a factor in the development of man- ences in disturbance histories and chance agement strategies, our understanding of events. relationships between physical site and community composition must be enhanced. COMPOSITION OF PRESETTLEMENT FORESTS As is true for the northern forest, the exact nature of the floristic and structural STATUS composition and the geographic variation of the southern forest before Euro-Ameri- PAST can settlement has never been described and will probably never be known with POST-GLACIAL ENVIRONMENT certainty. However, descriptions and occurrences of prominent forest types, at Only the eastern half of the state least in terms of tree species composition, south of the tension zone was glaciated were recorded by numerous early observers during the most recent stage of the Pleis- (e.g., Knapp 1871, Chamberlin 1877, tocene. The relationship between various Warden 1881). These observers recognized glacial landforms and basic soil types, as southern forests as distinct from northern outlined in the section on the northern types even though many tree species forest, also applies here. Most of the occurred in both regions. The predomi- southwestern portion of the state, known as nance of oaks and general absence of the Driftless Region, has escaped glaciation conifers were key distinguishing features at least over the last 750,000 years. How- noted by all observers. Another feature of ever, contrary to common belief, the region southern forests often singled out by early was not entirely surrounded by ice at any travelers was the relative openness or park- time; thus there was always an open route like appearance due to the lack of small for migration of flora and fauna. In fact, the trees and shrubs. For example, one could significance of the Driftless Region has easily ride on horseback through the more to do with its function as a source of woods, a condition much less common in flora and fauna for post-glacial reinvasion northern forests. of glacial regions than it does with unique- The best source of information on the ness of its soil parent material. Most of composition of vegetation in Wisconsin

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during the earliest period of Euro-American Oak openings: bur oak—white oak— settlement comes from the records of the black oak. This savanna community rectangular survey of public lands (General type often occurred as a transition Land Office Surveys). The nature of these between oak forest and prairie. This type records and methods used to interpret could not always be distinguished from them for purposes of constructing maps of Type 3 on the basis of surveyors’ presettlement vegetation have already been records. Often it was not possible to summarized in the section on the northern determine whether the trees occurred in forest community. A simplified map of a close enough spacing to represent a presettlement vegetation constructed from true forest or whether they occurred as survey records by Finley (1976) is shown openings or savanna. This forest type is in Figure 10. discussed in detail in the “Oak Savanna Seven of the 11 forest types recog- communities” section of this report. nized by Finley occur in northern Wiscon- sin and have already been described. The four southern types are as follows: FACTORS CONTROLLING THE DYNAMICS OF Sugar maple-basswood with red oak, PRESETTLEMENT FORESTS white oak, or black oak as major associ- Explaining the composition, distribu- ates. This type of forest occurred in tion, and dynamics of southern Wisconsin’s three major forests has been a blocks, one challenge to plant centered in There is ample evidence that the ecologists and Richland and foresters for genera- vegetation mosaic at the time of Euro- Vernon counties, tions. Although we another in Wash- American settlement was largely a result do not yet have all ington and Dodge of fire regimes that existed for 5,000-6,000 the answers, a counties, and a years prior to that time. consensus is emerg- third in Pierce ing on many issues. County. Numerous Over the last century, small segments the region south of the tension zone has occurred in other counties, particularly been regarded by some as part of the more Grant, Green, Lafayette, and Sauk extensive eastern oak-hickory forest or even counties. oak savanna (Kuchler 1964) and by others as maple-basswood forest (Daubenmire American beech—sugar maple— 1936, Braun 1950). The presence in the basswood with red, white, or black oak region of both mesic maple-basswood as major associates. This type was forests (greatly resembling the clearly mesic similar to Type 1 above, except that northern hardwoods forests) as well as beech was often dominant or shared drier oak forests and even savannas and dominance with sugar maple. This type prairies caused much misunderstanding occurred in a narrow north-south belt and confusion. However, ecological evi- along Lake Michigan. It also coincided dence accumulated to date clearly suggests with the geographic range of beech in that without regular, moderate to severe fire southern Wisconsin. disturbance, southern Wisconsin’s climate can support mesic forests on most loamy White-oak—black-oak—bur-oak. This soils. Only on sands or shallow loams on loosely defined type occurred in a southern and western exposures can oak seemingly random pattern throughout forests be expected to persist. Without fire, the region south of the tension zone. perpetuation of prairies and savannas, including oak openings, is virtually impossible.

74 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE There is ample evidence that the sure that the floristic differences between vegetation mosaic at the time of Euro- two physiographic types or soil types are American settlement was largely a result of due to site constraints or to fire history. We fire regimes that existed for 5,000-6,000 are currently conducting extensive floristic years prior to that time. Because of differen- sampling of forest stands stratified by site tial sensitivity of tree species to fire damage, factors and presettlement vegetation types. the communities in existence prior to Euro- This information should help us to better American settlement were clearly related to understand the dynamics of southern forest the frequency and intensity of fires. All types. mesic hardwoods and particularly sugar The concept of “old growth,” as maple are easily killed by fire at all stages of understood in the context of northern growth. Oaks, on the other hand, have forests, is applicable in the south only to many adaptations to fire environment. the mesic community types, which are the Saplings and seedlings of all oak species only types capable of maintaining them- native to Wisconsin resprout readily when selves without disturbance. However, old- tops are killed. Bur oak has the greatest growth dry-mesic and dry forests, while capacity for resprouting, followed by black, very rare today, were maintained by natu- white, and red oak. Mature trees also rally occurring disturbances such as fire. possess varying degrees of resistance to fire These natural ecological dynamics are damage, in the same species order. essential to the maintenance of these and Thus, mesic other climax and old- forests could persist Euro-American settlers converted growth communities. in southern Wiscon- There are probably no extensive acreages of southern forest sin only on those true old-growth oak landscapes relatively to agriculture. forests left in south- free of fire distur- ern Wisconsin, with bance. Surveyors’ records clearly showed the possible rare exception of those oak that such forests occurred where rivers or forests growing on the mid-slope area of lakes formed firebreaks against fires driven north and east slopes of very steep south- by the prevailing southwesterly winds. western Wisconsin ridges. Some of these Landscapes subject to moderate fire fre- forested ridges of the Driftless Region quency supported oak forests, while those exceed 450 feet in height, and only those more frequently burned supported oak portions of the side slopes that could be openings or other savanna types. Each of reached with cable were logged. This these community types, once developed, commonly left old-growth strips 200-300 contributed toward its own perpetuation. feet wide at mid-slope, extending the Thus, open grasslands burned most readily length of the ridge. Today, those strips are while mesic forests were far less likely to imbedded in second-growth forest growing burn due to their more humid interior above and below. condition, lower wind speed, and lack of flammable vegetation. THE LOGGING AND EURO-AMERICAN SETTLEMENT ERA CLIMAX AND OLD GROWTH There was a significant difference in In the section on northern forests, we the impact of Euro-American settlement on discussed how site conditions (e.g., soils, northern and southern Wisconsin forests. topography) limit the development of While in the north the impact was mainly climatic climax and how the floristic on forest composition; in the south, Euro- composition of a community can be used to American settlement meant elimination and characterize and classify communities and conversion to agriculture of extensive forest sites. In southern Wisconsin this process is acreage. Forests not cleared for farming complicated by fire history. We cannot be were almost universally high-graded for

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lumber, fuelwood, railroad ties, and other revert at least temporarily to shrub commu- products and were subsequently or simul- nities. taneously grazed by cattle or sheep. Be- cause wild fires were also suppressed with Euro-American settlement, former oak ANIMALS savannas not used for farming rapidly The forests of southern Wisconsin transformed into oak forests of generally prior to Euro-American settlement sup- low economic value. ported a rich fauna that included large herbivores and carnivores such as bison, PRESENT elk, white-tailed deer, cougar, bobcat, and black bear, and a great variety of smaller VEGETATION mammals as well as wet-forest furbearers— mink, otter, beaver, and muskrat (Jackson Perhaps the most conspicuous charac- 1961)—and a rich avifauna. Remaining Because oaks are teristic of the present southern forest as a habitat patches, most of them less than 125 intolerant of shade, whole is its fragmentation. Percentage of acres, appear to still support most of the the heavy cutting forested area for various southern counties species found at the time of Euro-American that went on for ranges from almost zero in some eastern settlement. Many of the generalists and counties to 30% or 35% in the western several decades adaptive species have increased their “coulee” region. The remaining forests exist after Euro- populations (e.g., deer, raccoon, skunk, red mostly in small blocks or patches. Some fox, robin, blue jay, and cowbird). The wild American notable larger blocks are found in the turkey has been successfully reintroduced settlement Baraboo range, the northern unit of the over the past 15-20 years. stimulated oak Kettle-Moraine State Forest and the Today, except for the deer and coyote, reproduction, even Kickapoo River valley all of the large region. The general in mesic forests herbivores and condition of south- Perhaps the most conspicuous originally carnivores are absent ern Wisconsin forests characteristic of the present southern from southern dominated by is perhaps of greater forest as a whole is its fragmentation. Wisconsin, and a maple and concern to foresters number of them are basswood. than it is to ecolo- gone from the state. Subsequent gists. Red and white These species losses selective cutting of oak are of considerable economic value, but and other concerns in faunal composition their supply is decreasing. The initial these forests again and survival in the southern Wisconsin impact of Euro-American settlement created the forests are a result of forest fragmentation actually resulted in an increase of oaks in and ecological simplification brought on by environment more present stands. Because oaks are intolerant the rapid spread of agriculture and urban- favorable to of shade, the heavy cutting that went on for ization along with unregulated subsistence tolerant several decades after Euro-American and commercial hunting. hardwoods. settlement stimulated oak reproduction Birds provide the best insight into the even in mesic forests originally dominated status of southern forest animals, for birds by maple and basswood. Subsequent have been far more intensively researched selective cutting of these forests again and are subject to more regular surveys created the environment more favorable to than any other animal group. Though it tolerant hardwoods. Thousands of acres of remains largely intact today, this faunal previous oak savannas not utilized for group is faced with mounting problems. farming rapidly grew into dense oak forests The passenger pigeon, a colonial forest bird through sprouting of the fire suppressed that inhabited the southern Wisconsin root stalks called “grubs.” However, these forests, is extinct. While only two other forests are not regenerating. If mesic birds, the carolina parakeet (extinct) and hardwood seed source is lacking, many of the swallow-tailed kite (extirpated), have these forests will gradually break down and been lost from the southern forest land-

76 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE scape, many species have been negatively Many observers have noticed a Birds provide the impacted by habitat loss, reduced size of significant increase in forest edge bird best insight into habitat area, and changes in the composi- species (Bond 1957; Howe and Jones 1977; the status of tion and structure of forests and woodlots. Ambuel and Temple 1982; Robbins 1991; These changes have affected bird distribu- David Sample, Wis. Dep. Nat. Resour., southern forest tion and abundance to the point where pers. comm.). One species that has ben- animals, because many species are listed as endangered, efited from increased edge is the brown- birds have been threatened, or of special concern, and headed cowbird. Brittingham and Temple more intensively others show signifi- (1983) have shown researched and cant population that the cowbird, a are subject to more declines. The forests of southern Wisconsin prior to brood parasite on For example, Euro-American settlement supported a forest songbird regular surveys Bond (1957) noted rich fauna that included large herbivores species, has reduced than any other that interior forest and carnivores . . . . Today, except for the reproductive success animal group. species preferred deer and coyote, all are absent from for a number of forest larger and more songbirds and may be southern Wisconsin, and a number of mesic forests, while responsible for their generalists and them are gone from the state. recent declines. An disturbance-prefer- additional concern is ring species showed the role that edge affinity for smaller, pioneering forests. In a birds play in predation of interior species. study of southern flood-plain forest, As crows, blue jays, and grackles increase Mossman (1988) found that at least 20 in number, so too will their predation on species appeared to require stands at least forest nesting birds increase. Brood parasit- 40 acres in size, and some required much ism and predation, along with other larger tracts. There are at least 12 songbirds elements of habitat loss and modification, that depend on forests in excess of 40 acres have combined to create “population in size, with three requiring a minimum of sinks,” poor-quality habitats in which The extensive 161 acres and five more requiring either populations produce deficit numbers that floodplain forest along 200-acre or 240-acre woodlots to have at require subsidization from other popula- the St. Croix River in Polk County harbors least a 50% chance of supporting a breed- tions (Whitcomb et al. 1981). many species of ing population (Temple 1988). The average Mammals are much more poorly interior-nesting birds, size of a southern Wisconsin woodlot is understood than are birds in relation to the while the small southern forests. Lack of comprehensive patches of forest in the currently 47 acres. Consequently, many of distance lack these these area-sensitive, interior-dependent inventories and population surveys means bird species. Photo by songbird species are decreasing in fre- that most current knowledge is based on Eric Epstein. quency and undergoing population declines (Ambuel and Temple 1982, Wis. Dep. Nat. Resour. 1991). Ecological simplification has also impacted southern forest avifauna. Reduc- tion in area is often accompanied by grazing, logging, and cutting and gathering of firewood, activities which have altered both forest composition and structure. For example, over-grazing eliminates understory grasses, herbs, and shrubs, depriving insect and foliage-gleaning foragers of a source of food. Habitat for cavity-nesting and insect-foraging birds is removed through logging and wood-gathering.

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observations and relatively few studies of species (Jansen and Anderson 1981, local species populations. However, mam- Mossman and Hine 1984, Mossman and mals in general don’t seem to show the Huff 1990, Huff 1992). This survey was same correlations with habitat area and initiated because of the concern that The average size quality decline as birds do (e.g., species amphibian populations were declining for of a southern decline, or in mammals, abnormal some species in Wisconsin and globally Wisconsin woodlot rhythms) (Frank Iwen, UW Zoological (Modern Medicine 1973, Les 1979, Hine et Museum, pers. comm.). In general, forest al. 1981, Vogt 1981) is currently 47 mammals are secure, with some species’ Several southern forest amphibians acres. populations apparently increasing, such as are susceptible to changes in habitat Consequently, voles, mice, and shrews, where understory structure. These primarily include the many area- composition and structure are well pro- species dependent on ephemeral or vernal sensitive, interior- tected and maintained. There also appears ponds for breeding, such as the chorus dependent to be an increase in forest mammalian frogs, eastern gray and Copes gray tree songbird species predators that are capitalizing on the frogs, wood frogs, and blue spotted and increased small mammal populations eastern tiger salamanders (see “Actions are decreasing in (Frank Iwen, pers. comm.). Causing Concern” section). Snakes associ- frequency and The white-tailed deer has, over the ated with the southern forest that are of undergoing last 25-30 years, expanded its range concern include the black rat snake, timber population southward and is present in great abun- rattlesnake, and massasauga rattlesnake. declines. dance in southern forest and woodlots. In The impacts of natural succession, forestry some areas their numbers are so great that practices, and other land-use and manage- browse impacts are ment activities on readily observed by these species are not the elimination of Little is known about the historic or current well understood. The some plant species abundance of southern forest amphibians timber rattler and (e.g., certain orchids and reptiles . black rat snake are and Canada yew) communal denning and reduced repro- snakes whose local duction of cedar, oak, and maple, among populations are susceptible to losses of others. Deer negatively affect cover for critical hibernating sites. These and other ground nesting birds. Numbers of deer communal denning snakes are also more continue to increase, and this species’ range vulnerable to destruction or collection since is expanding into all suitable habitat. they are clustered in quantities especially A few mammalian species have not during spring emergence from den sites adapted well to current conditions. Loss of (Robert Hay, Wis. Dep. Nat. Resour., pers. forest structure and spraying for insect comm.). Both the massasauga and timber control in agricultural areas has posed rattlers have seen demonstrable declines in problems for the southern forests’ solitary populations throughout Wisconsin and the bats. Fox squirrels also appear to be rest of their ranges. Both have been im- declining in southern forest edge, as these pacted by habitat loss and bounties which areas convert to closed forest. have virtually eliminated them from many Little is known about the historic or areas. They are still killed because of their current abundance of southern forest unfounded reputation as being very dan- amphibians and reptiles (herptiles). Re- gerous. The massasauga is the most seri- gional distributions studies for herptiles are ously endangered species of reptile in the ongoing by a group of amateur and profes- state, now restricted to only a few lowland sional herpetologists as part of the Wiscon- hardwood forests, forest edges, and adja- sin Herpetological Atlas Project (Casper cent upland ﬁelds (Vogt 1981). 1986). The DNR has been conducting an Little is known about the inverte- annual frog and toad survey since 1981 to brates of the southern forests of Wisconsin. determine the population trends of these Diversity in forest structure plays an

78 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE important role in meeting the needs of Fire, perhaps the most important lepidopterans as well as other insects and ecological tool in establishing and invertebrates. Recent surveys have focused maintaining oak forests, will not be on lepidoptera, which may in the future employed sufﬁciently as a prescribed serve as indicators of change because of management practice. their frequent association with host plants and species-speciﬁc food sources and their relative sensitivity to habitat perturbations. ACTIONS CAUSING CONCERN

PROJECTED Ecological consequences of altered vegetation dynamics, as described above, Given the rate and means by which There is no are difficult to assess. There is no question southern forest fragments in some areas, question that particularly southeastern Wisconsin, are that natural community diversity of the natural community being harvested and developed as rural southern forest landscape has been reduced homesites, the following trends can be since the suppression of presettlement fire diversity of the expected: regimes, and lack of fire in the oak wood- southern forest lands and forests is a concern. In addition, landscape has Fragmentation and reduction in size of native vegetation is extremely vulnerable to been reduced woodlots will continue. replacement by exotics. However, there is since the no clear evidence that any forest plant Highest quality woodlands will continue species have been lost. The southern forest suppression of to be lost. fauna, both vertebrate and invertebrate, has presettlement fire apparently been more regimes, and lack At the current rate severely impacted. It of fire in the oak of harvest, oak Little is known about the invertebrates of appears that forest woodlands and may cease to be a the southern forests of Wisconsin. fragmentation is of forests is a commercially primary concern in viable product in terms of faunal concern. In the future. diversity. Additionally, structural and addition, native compositional changes from intensive land- vegetation is Emphasis on hardwood saw logs will in use practices, exotic and edge species extremely the near future shift from oak to other encroachment, and grazing have adversely vulnerable to southern forest hardwoods such as sugar impacted southern forest fauna. The replacement by maple, black cherry, hackberry, walnut, southern forest avifauna is particularly exotics. However, and white ash, further reducing both vulnerable to fragmentation and simplifica- long-term veneer and saw-log supply tion. there is no clear and overall species composition and Forest amphibians also are a primary evidence that any stand structure. concern because of their vulnerability to forest plant habitat changes and pesticide use in species have been Forest composition will vary greatly, adjoining agricultural lands. Intensive lost. The southern with both commercially and ecologically forest management and woodlot scavenging forest fauna, both less desirable species (such as black can significantly open or disturb large areas locust, box elder, and persistent dense of forest, which leads to siltation and vertebrate and shrubs) replacing oak and maple forest premature drying of vernal ponds, reducing invertebrate, has communities in some areas. or prohibiting amphibian metamorphosis apparently been (Robert Hay, Wis. Dep. Nat. Resour., pers. more severely Poor management practices will reduce comm.). Also of concern in these disturbed impacted. productivity, decrease long-term eco- areas is the loss of structural habitat com- nomic value, and diminish sustainability posed of large, dead woody debris, heavy of the southern forest community loam, and thick surface litter, which are complex. habitat characteristics essential for amphibians (Gary Casper, UW-Milwaukee, pers.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 79 SOUTHERN FOREST COMMUNITIES

many areas forest regeneration may be frequently reduced by deer browsing.

SOCIO-ECONOMIC ISSUES

As in all forested regions, conflicts between traditional uses of forests, recreational demands, and concerns for preserving natural communities is intensifying. Numerous misconceptions about the nature of forest ecosystems exist among forest owners as well as the general public. The process of change through natural succession is seldom appreciated. Forest owners too often agree to sell only the highest quality trees, usually oaks, and thus slowly convert their woodlots to tolerant hardwoods. On the other hand, the general public often sees any disturbance, particularly clearcutting and fire, as unnatural and always detrimental. In order to maintain a desired diversity of forest communities, extensive education of forest owners and the public will be required. Because most of the forest land in southern Wisconsin is in private ownership it may seem that public opinion does not matter. However, the public everywhere is becoming progressively more proactive, and its influence on the legislature and the courts is increasing. We should expect management decisions to be increasingly questioned.

This stand of southern comm.). Roads near breeding areas are also hardwoods is a threat along with the “clean-yard” prac- POTENTIAL FOR COMMUNITY converting from oak to more tolerant species tices of homeowners building in woodlots. RESTORATION such as ironwood and Gypsy moth control efforts pose a red maple. Photo by serious threat to native lepidopterans. If the desired state is considered to be John Kotar. Aerial application of the biological control some representation of all presettlement agent Bacillus thuringensis (B.t.) can kill forest communities, considerable difficul- In order to native species larva. Because susceptibility ties will be encountered with its implemen- maintain and to B.t. is dependent on the time of emer- tation. Restoration of oak savannas would restore a diversity gence, and time of emergence is variable be the most difficult, both from an ecologi- of forest over several weeks, native lepidopterans cal as well as an economic standpoint. with synchronous emergence patterns are Intensive management through the use of communities, equally vulnerable to mortality. Agricultural fire would be necessary, and without extensive pesticide use is also of concern for the economic incentives it is not likely to be education of forest invertebrate community. applied to private lands. Restoration and owners and the Of increasing concern for the south- maintenance of mixed oak forests is cer- public will be ern forest is the artificially maintained high tainly possible from the ecological point of required. deer density. There is now evidence that in view, but greater economic incentives and

80 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE technical support will be needed to enable landowners to apply proper management techniques. Without such incentives and support, high-grading and degradation of oak forests are likely to continue. Mesic forest restoration and maintenance would be relatively easy. However, because of direct competition with farming, most forest communities will probably remain confined to terrain unsuitable for cultivation. Today, large tracts of floodplain are limited to the Lower Wisconsin and Missis- sippi river valleys; most of the southeastern floodplain forests have been destroyed or reduced to small patches. Large areas of upland forest are restricted to parts of Crawford and Vernon counties, the Baraboo Hills, the northern unit of the Kettle Moraine State Forest, and parts of Manitowoc County. The potential for restoring additional large tracts of each forest type is relatively good in at least some areas of both western and eastern Wisconsin. There are also a number of swamp and bog forests still intact, though often degraded, in the south-central and southeastern counties. Many of these forests have been reduced in size by drainage, agricultural encroachment, and grazing; however, many could be restored over time by reversing the drainage processes. The best way to enhance biodiversity based on ecoregion Mature southern across the southern The best way to enhance biodiversity mesic forest of sugar forest landscape is to goals, staff workload, across the southern forest landscape is to maple and basswood. increase the size of fiscal resources, Lost Lake State individual woodlots increase the size of individual woodlots public input and Natural Area in the support, and legal eastern end of the and reduce their and reduce their fragmentation. Baraboo Hills. DNR fragmentation. authority. We will photo Achieving this goal work with our will be difficult, but potential does exist in customers and some parts of the southern forest. clients to set priorities and bring recommendations to the Natural Resources Board for consideration beginning in the 1995-97 biennium. POSSIBLE ACTIONS 1. Community-specific actions: The following possible actions are consistent with ecosystem management, Mesic Forest (including oak-dominated but require more analysis and discussion. forests with presence of shade-tolerant How priorities are set within this list will be mesic hardwoods such as sugar maple,

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 81 SOUTHERN FOREST COMMUNITIES

extensive, even before Euro-American settlement, it is recommended that they be maintained wherever they still occur.

Mixed Oak and Oak-Hickory Forest. Where no tolerant hardwoods are present, these forests are not threatened by succession; however, neither do they regenerate without periodic disturbance. It is recommended that these forests be maintained by appropriate silvicultural methods, including prescribed burning. While no single method has been shown to work in all situations, a number of techniques have been used successfully across a range of site types.

Oak Openings (Savanna). Although this community type is treated in detail separately in this report, it is included here because it is dynamically related to oak forests. Oak openings and savannas in general are among the rarest community types in Wisconsin. Over most of their former range, they have been eliminated by farming or have naturally converted to closed-canopy oak woodlots. Restoring these community types would clearly enhance local as well as regional biodiversity. Although restoration methods are still in developmental stages, it is almost certain that prescribed burning rather than mechanical manipulation of vegetation will be Mixed oak and hickory American beech, Because the conifers are rare in southern required. Because of stands do not basswood, white regenerate without Wisconsin’s forest communities, restoring predominantly private ash, or bitternut periodic disturbance. the presence of white pine and its ownership, large-scale This stand of young hickory). Mesic restoration of these oak is growing back forests are associated communities to their previous after clearcutting. communities will be relatively stable. southern range would enhance Photo by John Kotar. difﬁcult without In the absence of biodiversity. providing additional periodic major economic incentives. disturbances, the dominance of shade-tolerant hardwoods Mixed Pine-Oak Communities. With the increases. Mixed composition can be exception of the “central sands” region maintained with proper silvicultural and a narrow belt along the shores of techniques (e.g., shelterwood, group Lake Michigan, pines have not generally selection, clearcutting). For optimal been considered as a natural component biodiversity, some mesic forests should of southern forest. However, a number be maintained in a mixed state. Because of scattered oak-white pine communities southern mesic forests were never (e.g., Devil’s Lake State Park) and several

82 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE white pine relicts suggest that this type 3. Whenever possible, reduce fragmenta- could be maintained on many land- tion of woodlots by enlarging current scapes. White pine and red oak have blocks and providing wooded corridors similar ecological requirements and the through reforestation. two species can be managed together. Because the conifer component is rare in 4. Work toward the development of southern Wisconsin’s forest communi- economic incentives for private land- ties, restoring the presence of white pine owners to enable them to participate in and its associated communities to their resource management programs that previous southern range would enhance protect biodiversity. Only through such biodiversity. programs will it be possible to implement the specific recommendations 2. Old-growth restoration and maintenance listed in this report on a region-wide areas in southern mesic forests should basis. first be addressed on Department lands where the largest southern mesic forest 5. In order to coordinate management tracts remain. Areas of remnant old practices consistent with state-wide growth should be objectives, some type maintained and of regional “informa- enhanced by tion center” will have allowing sur- Old-growth restoration and maintenance to be created. For rounding forest to areas in southern mesic forests should example, a land attain old-growth first be addressed on Department lands manager on a given condition through where the largest southern mesic forest property may be natural processes. tracts remain. taking all the correct On appropriate measures to optimize Department lands, local biodiversity, but designated old- without some source growth areas can be enhanced by of information on wider, regional needs surrounding each with a mature forest he/she may nevertheless be acting management zone based on selection inappropriately. In order to allow for harvest practices (see the “Northern more natural type conversions (through Forest” section Possible Actions). Private succession), there will be a need for woodland owners should be encouraged planning regional rotations of cover to apply selection management prac- types. Forest nurseries are producing a tices, which would allow trees to reach a much greater diversity of planting stock, much older age before harvest and including pioneer species such as aspen, would build old-growth structural because management through natural characteristics into their forests. Where succession must go hand in hand with woodlands occur in close proximity, the establishment of compensating encourage blocking through the refores- pioneer stages. tation of intervening open lands, thus enhancing mature old-growth forest 6. Bring together the large amount of characteristics of the existing patches. existing technical information on Encourage participation in private forest silviculture, forest ecology, and wildlife assistance programs such as the Man- ecology by establishing a natural com- aged Forest Law and the U.S. Forest munity information system. The system Service Stewardship Incentive Program. should have the following characteristic:

A basic inventory of wildlife species.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 83 SOUTHERN FOREST COMMUNITIES

A basic inventory of wildlife habitats Procedures for applying species and plant communities. habitat relationships to integrate resource planning and management. A basic model of the relationships of wildlife species to these habitats Guidelines for managing special (Verner et al. 1986). habitats or stand conditions.

A computerized storage and retrieval A monitoring strategy. system.

Case Study

THE BARABOO HILLS: PARTNERS PROTECTING AND MANAGING IN AN ECOSYSTEM CONTEXT

Contributed by Eric Epstein and Becky Isenring.

The ancient quartzites of the Baraboo Hills rise hundreds of feet above much of the surrounding landscape. Thin soils, steep slopes, low fertility, and public interest discouraged the intensive development now characterizing the vast majority of southern Wisconsin. Today the Hills are mantled with the most extensive upland deciduous forests (totalling about 55,000 acres) remaining in the southern part of the state. In a landscape dominated by agriculture, where most remnant natural vegetation occurs in small, isolated, and often highly disturbed stands, the Hills are an oasis for one of the most diverse arrays of natural communities, plants, and animals in the upper midwest. Naturalists, conservationists, and scientists from many disciplines have been drawn to the Baraboo Hills for well over a century. In 1911, the creation of Devil’s Lake State Park marked the first effort to protect a portion of them in perpetuity. The state purchased and acquired other significant properties in the years that followed, including Parfrey’s Glen (Wisconsin’s first State Scientific Area), Natural Bridge State Park, Lost Lake State Natural Area, McGilvra Woods State Natural Devil’s Lake is Area, and Pewit’s Nest State Natural Area. Significant conservation ownerships are also held imbedded in the by the University of Wisconsin Foundation (Potter Preserve), the University of Wisconsin- Baraboo Hills and the Baraboo (Van Zelst Barrens), the Village of Rock Springs (Weidman Park), and Wisconsin most extensive forest in southern Wisconsin. Society for Ornithology (Honey Creek). The band of continuous forest of the south The Nature Conservancy (TNC), a private conservation organization with a large range of the Hills membership and history of involvement in the Baraboo Hills protection efforts dating back narrows at this point some thirty years, is a leading partner. Many of their active projects are focused on the most from a wider band to the west. Photo by ecologically important sites in the Hills, including Baxter’s Hollow, Hemlock Draw, Pine Michael J. Mossman. Hollow, and Misty Valley.

84 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Until recently, most of the conservation work in the Hills had been devoted to individual projects. To take advantage of the unique opportunity for protection and management at a large scale, it was clear that a unified, expanded vision encompassing the scope and attributes of the entire ecosystem was necessary. In 1991, TNC took a major step to support this vision by initiating a two-year biological inventory in parts of Sauk and Columbia counties, targeting an area of 144,000 acres. The area inventoried was defined principally by the underlying Baraboo quartzite. Field staff included ecologists, botanists, and zoologists. Information was collected on all types of natural communities occurring in the Baraboo Hills, and on many plant and animal species. DNR personnel from the Bureaus of Research, Endangered Resources, Parks and Recreation, and Forestry provided assistance through training, consultation, and development of sampling design. Many individuals in these programs also contributed personal records to the Baraboo Hills Inventory. The Natural Heritage Inventory provided existing computerized records for the area surveyed, a database to store and maintain records, a format to record field data, and a methodology for ranking natural communities and rare species populations. To meet the existing and anticipated needs of forest managers in the Hills, vegetation data were collected by TNC’s inventory teams under the guidance of the UW-Madison’s Forestry Department. These data are being analyzed to identify habitat types, as part of a statewide Forest Habitat Classification system. Other key partners in this endeavor included the UW-Madison herbarium staff (speci- men identification), U.S. Forest Service (Forest Stewardship Fund), UW-Stevens Point (Biotic Index analysis of stream samples), numerous volunteers, and the cooperation of numerous landowners who willingly gave inventory staff access to their properties. To enhance the value and utility of the data collected through inventory, TNC worked with the UW Land Information and Computer Graphics Facility and the UW Institute for Environmental Studies to develop a GIS for the Baraboo Hills. Information incorporated into this system includes Natural Heritage Inventory data, “presettlement” vegetation, current land ownership, natural community covers, and geology. A computer model simulating the effects of land-use changes on neotropical migrant birds has been adapted using the GIS. Today, an exceptional opportunity exists in the Baraboo Hills to protect and manage an existing functional, diverse, forested ecosystem. The inventory has documented the ecological context for the Baraboo Hills. The new technologies have provided tools to aid in the synthesis and analysis of the information available. Now, many different public and private interests will work to support local and county leadership as plans for the future come into place. These plans will need to incorporate the inventory and other ecological, socio-economic, and political information and provide strategies to address threats, resolve conflicts, and ensure long-term success. Strong commitment to the eventual success of this unprecedented project is needed from all partners, public and private, large and small.

LITERATURE CITED

Ambuel, B. and S.A. Temple 1982. Ambuel, B. and S.A. Temple 1983. Area- Songbird populations in southern dependent changes in the bird Wisconsin forests: 1954 and 1979. J. of communities and vegetation of Field Ornithol. 53(2):149-158. southern Wisconsin forests. J. of Ecology. 64(5):1057-1068.

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Bond, R.R. 1957. Ecological distribution of Harper, K.T. 1963. Structure and dynamics breeding birds in the upland forests of of the maple-basswood forests in southern Wisconsin Ecol. Mono. 27(4). southern Wisconsin. Univ. Wis.- Braun, E.L. 1950. Deciduous forests of Madison. PhD. Thesis. North America. Blackstone Co., Heinselman, M.L. 1973. Fire in the virgin Philadelphia. forests of the Boundary Waters Canoe Bray, J.R. 1960. The composition of savanna Area, Minnesota. Quaternary Res., vegetation in Wisconsin. Ecology 3:329-82. 41:721-32. Hine, R.L., B.L. Les and B.F. Hellmich. Brittingham, M.C. and S.A. Temple 1983. 1981. Leopard frog populations and Have cowbirds caused forest songbirds mortality in Wisconsin. WDNR Tech. to decline? Bio. Science 33(1):33-35. Bull. No. 122. Cawley, E.T. 1960. A phytosociological Howe, R.W. and G. Jones 1977. Avian study of the effects of grazing on utilization of small woodlots in Dane southern Wisconsin woodlots. Univ. County, Wisconsin. Passenger Pigeon Wis.-Madison. PhD. Thesis. 39(4):313-319. Chamberlin, T.C. 1877. Native vegetation Huff, J. 1992. Wisconsin frog and toad of eastern Wisconsin. Geol. of Wis. 2: survey. pp. 114-17 in WDNR Bureau of 176-87. Research Surv. Rept. Christensen, N.L. and R.K. Peet. 1984. Jackson, H.H.T. 1961. Mammals of Convergence during secondary forest Wisconsin. Univ. Wis. Press, Madison. succession. J. Ecology., 72:25-36. 504 pp. Christensen, N.L. 1987. Succession and Jansen, D.K. and R.K. Anderson. 1981. natural disturbance: paradigms, Frog inventory for Wisconsin. UW- problems, and preservation of natural Stevens Point. unpubl. rep. systems. Agee, J.K. and D.R. Johnson, Kline, V.M. and G. Cottam. 1979. eds. Ecosystem management for parks Vegetation response to climate and ﬁre and wilderness—workshop. Univ. of in the Driftless Area of Wisconsin. Wash. Ins. For. Resour., Seattle. Ecology 60:861-868. Contrib. No. 65. Knapp, J.G. 1871. The native vegetation of Curtis, J.T. 1959. The vegetation of Wisconsin. Trans. Wis. State Hort. Soc. Wisconsin. Univ. Wis. Press, Madison. 1:119-25. 657 pp. Kotar, J. 1987. The role of phytosociology Daubenmire, R.F. 1936. The “big woods” of in forest site classiﬁcation in the Great Minnesota. Ecol. Monogr. 6:233-68. Lakes region. pp. 76-80. In D.W. Cole Davis, M.B., K.D. Woods, S.L. Webb, and and S.P. Gessel, eds. Forest site R.P. Futyma. 1986. Dispersal versus evaluation and long-term productivity. climate: expansion of Fagus and Tsuga Univ. Wash. Press, Seattle. into the upper Great Lakes region. Kotar, J., J.A. Kovach, and C.T. Locey. 1988. Vegetation 67:93-103. Field guide to forest habitat types of Dorney, J. 1981. The impact of Native northern Wisconsin. Univ. Wis.- Americans on presettlement vegetation Madison Dep. For. and Wis. Dep. Nat. in southeastern Wisconsin. Trans. Wis. Resour. 212 pp. Acad. Sci., Arts and Lett. 69:26-36. Kuchler, A.W. 1964. Potential natural Finley, R.W. 1976. Original vegetation cover vegetation of the conterminous United of Wisconsin (map). U.S. For. Serv., N. States [map] Am. Geogr. Soc. Cent. Exp., St. Paul.

86 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Leitner, L.A., C.P. Dunn, G.R. Mossman, M.J. and R.M. Hoffman 1989. Guntersperger, F. Stearns and D.M. Birds of southern Wisconsin upland Sharpe. 1991. Effects of site, landscape forests. Passenger Pigeon 51(4):343- features, and fire regimes of vegetation 358 patterns in presettlement southern Neuenschwander, H.E. 1956. The Wisconsin. Ecology 5:203-217. vegetation of Dodge County 1833- Les, B.L. 1979. The vanishing wild. Wis. 1837. Trans. Wis. Acad. Sci., Arts and Dep. Nat. Resour. 36 pp. Lett. 46:233-54. Lorimer, C.G., L.E. Frelich, and E.V. Robbins Jr., S.D. 1991. Wisconsin birdlife Nordheim. 1988. Estimating gap origin population and distribution past and probabilities for canopy trees. Ecology present. Univ. Wis. Press, Madison. 702 69:778-85. pp. Matthiae, P.E. and F. Stearns 1981. Steinbrenner, E.G. 1951. Effects of grazing Mammals in forest islands in on floristic composition and soil southwestern Wisconsin. In Burgess, properties of farm woodlands in R.L. and D.M. Sharpe eds. Forest island southern Wisconsin. J. For. 49:906-10. dynamics in man-dominated Stroessner, W.J. and J.R. Habeck. 1966. The landscapes. Springer-Verlag, New York. presettlement vegetation of Iowa 310 pp. County Wisconsin. Trans. Wis. Acad. McIntosh, R.P. 1950. Pine stands in Sci., Arts and Lett. 55:167-80. southwest Wisconsin. Trans. Wis. Acad. Temple, S.A. 1988. When is a bird’s habitat Sci., Arts and Lett. 40:243-57. not habitat? Passenger Pigeon 50(1):37- Modern Medicine. 1973. Where have all 41. the frogs gone? Modern Medicine, 12 Temple, S.A. 1990. Sources and sinks or Nov. regional bird populations. Passenger Mossman, M.J. 1988. Birds of southern Pigeon 52(1):35-37 Wisconsin floodplain forests. Passenger Warden, J.A. 1881. Forests and forestry in Pigeon 50(4):321-337 Wisconsin. Trans. Wis. State Hort. Soc. Mossman, M.J. and J. Huff. 1990. Results of 11:143-56. Wisconsin’s frog and toad survey 1984- 1989. unpubl. rep. Mossman, M.J. and R. Hine. 1984. Wisconsin frog and toad survey: establishing a long-term monitoring program. Wis. Dep. Nat. Resour., ER report #9.

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most subject to controversy.” Today there is still no widely accepted, clear-cut definition of what is meant by savanna. Fortunately for us in the Midwest, the term savanna has a relatively narrow CHAPTER 6 definition. Here it is generally used to describe an ecosystem that was historically part of a larger complex bordered by the prairies of the west and the deciduous forests of the east. This complex was a Oak mosaic of plant community types that represented a continuum from prairie to forest. Savannas were the communities in the middle of this continuum. The mosaic Savanna was maintained by frequent fires and possibly by large ungulates such as bison and elk. Oaks were the dominant trees, hence the term oak savanna. Communities Because savannas grade into both prairie and forest, there are no clear divid- by Richard Henderson ing lines between savanna and these two Bureau of Research communities. In classifying the plant Department of Natural Resources communities of Wisconsin, Curtis (1959) was forced to set limits for what he called savanna. He ultimately defined it as having no less than one tree per acre and no more than a 50% tree canopy. However, Curtis made it clear that these limits were arbitrary and chosen purely for convenience. Curtis also subdivided Wisconsin savannas DESCRIPTION into four categories based on plant composition: oak barrens, pine barrens, oak opening, he term savanna has never been and cedar glade. He defined oak barrens as well defined. It has its origin in savannas with black/Hill’s oak on infertile, the early Spanish colonization of droughty sand or sandstone-derived soils. the Caribbean in the 16th Pine barrens were defined as savannas with The term century, where it was applied to jack/red pine on similar soil types as oak savanna is used treeless grassy plains (Johnson barrens. Oak openings were defined as in the Midwest to and Tothill 1985). By the end of the 19th savannas on rich, mesic soils with mostly describe an century, this Spanish term was widely used bur or white oak. Cedar glades were defined ecosystem by plant geographers to describe tropical as savannas on dry limestone bluffs, with grasslands. Also by this time, woody plants red cedar more prevalent than oaks. bordered by the had became an accepted and, in some Another savanna community type, wet and prairies of the west cases, even mandatory part of the defini- wet-mesic soil savannas, was not listed by and the deciduous tion. By the mid-20th century ecologists Curtis, because not enough intact examples forest of the east— were still struggling with the definition of could be found at the time of his study. Bur a mosaic savanna, especially in North America and swamp white oak were probably the maintained by (Penfound 1962). Cole (1960) summed up dominant trees of this community historically. The following discussion mostly frequent fires and the situation this way: “Perhaps of all types of vegetation the savanna is the most covers the community types Curtis called possibly by large difficult to define, the least understood, and oak opening, but it applies to other savanna ungulates. the one whose distribution and origin is the types as well. The sandy soil oak and pine

88 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE barrens are covered in a separate chapter in this report. Just what the understory and ground layer vegetation of oak savannas was like is largely unknown. Bray (1960) described the oak savannas as having less grass and more forbs and woody shrubs than prairie, but more grass and fewer forbs than forest. Historically, the savanna community was probably a slowly shifting mosaic of plant species associations that had varying degrees of shade and sun tolerance. Conse- quently, the flora of oak savanna was probably a blend of the following species: True “sun-loving” prairie species that can tolerate or survive only light shad- ing. for Bray, Curtis, and others to study were An oak opening is a limited in number and size and had prob- savanna on rich, mesic soils with mostly Prairie-associated species that do well, ably already been altered to some degree by bur or white oaks. or perhaps slightly better, in light shade absence of fire and a history of domestic Here is a white oak than in full sun. livestock grazing. Recent information and with prairie-like understory in a observations resulting from savanna subdivision in Dane True savanna restoration attempts County. This tree has species that do best over the past typical open-grown Historically, the savanna community was decade suggest that architecture, is more in, or are restricted than four feet in to, a blend of shade probably a slowly shifting mosaic of plant the original oak diameter, and and sun. species associations that had varying savanna vegetation probably got its start may have been even around the Revolution- degrees of shade and sun tolerance. ary War. Photo by Forest-associated more diverse and Richard Henderson. species that do well specialized than the with fire and moderate amounts of Bray and Curtis sunlight. studies indicate (Packard 1988a, 1988b; Bronny 1989; Clewell 1989; Pruka 1994; True forest species that can persist, but W. Pauly, Dane Co. Parks, unpubl. data; R. do not necessarily thrive, with occa- Henderson, Wis. Dep. Nat. Resour., sional fire and moderate sunlight. unpubl. data). The more wooded part of the histori- Although oak savannas were probably cal prairie-forest complex (i.e., savanna or relatively dynamic communities compared woodlands with 50%-100% tree canopy) is with prairies or forests, major vegetation known to us only through the early ac- changes within these savannas still took counts of explorers and settlers. This decades if not centuries to occur. community was already so distorted by lack Detailed descriptions of Wisconsin of fire and other disturbances by the mid- oak savanna vegetation can be found in 1900s that it was not even classified and works by Bray (1958, 1960) and Curtis studied as a separate community by Curtis (1959). These studies provide the best and his students. What remained of this available data on savanna vegetation; community at the time of the Curtis studies however, they should not be considered the (i.e., grown-in savannas) was lumped with final word on historical savanna. By the the dry or dry-mesic southern hardwood time these studies were done, the savanna forest communities based on the residual as a complete ecosystem had already been oak trees, often independent of the actual gone for 100 years. The remnants available soil moisture regimes of the sites. Recent

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 89 OAK SAVANNA COMMUNITIES

research is now starting to shed some light savanna. At the time of Euro-American on this plant community. Pruka (1994) settlement, Wisconsin had an estimated 5.5 studied the sorting out of groundlayer plant million acres of oak savanna (not including species along the natural sunlight gradients the 4.1 million acres of oak and pine found in savanna and woodland. barrens) (Curtis 1959) and an additional This more heavily wooded portion of 1.4 million acres of oak forest, much of the prairie-forest complex (up to and which may have been open oak woodland including 100% closed canopy) might best (see Fig. 10). have been described as an open oak woodland. Although much work needs to PRESENT be done in describing and understanding this community, it should most likely be In the early to mid-19th century, the viewed as separate from oak forest. Based oak savanna as an ecosystem was thor- on historical accounts, it had a “park-like” oughly fragmented and nearly totally structure, with the destroyed throughout its range. Most of its dense shrub and acreage suffered one of the following fates: understory tree Oak savanna now shares equal billing layers associated (1) clearing and with tallgrass prairie as the most with oak forests of plowing, (2) overgraz- today kept sparse threatened plant community in the ing, or (3) invasion by and low in stature by Midwest and among the most threatened dense shrub and tree ﬁre. The ground in the world. Intact examples of oak growth due to lack of ﬁre, lack of grazing, layer was probably savanna vegetation are now so rare that or both. Oak savanna dominated by forest less than 500 acres are listed in the species of low- to now shares equal Natural Heritage Inventory as having a mid-shade tolerance billing with tallgrass (e.g., summer- and plant assemblage similar to the original prairie as the most fall-blooming oak savanna. This is less than 0.01% of threatened plant grasses, sedges, the original 5.5 million acres. community in the legumes, and com- Midwest and among posites) that are the most threatened today doing best in forest gaps and edges, in the world. Intact and savanna species of mid- to high-shade examples of oak savanna vegetation are tolerance. now so rare that less than 500 acres are listed in the Natural Heritage Inventory as having a plant assemblage similar to the original oak savanna. This is less than STATUS 0.01% of the original 5.5 million acres. PAST Many plant species that were probably savanna specialists are now uncommon and Oak savanna has probably been in are found only in the fringes and openings North America for 20-25 million years of oak woods, brushy areas, and lightly (Barry and Spicer 1987), shifting about and grazed pastures. Some examples are yellow expanding and contracting with climatic pimpernel, pale Indian plantain, woodland changes. For the past several thousand thistle, downy wild rye, elm-leaved golden- years it has existed in a more or less stable rod, New Jersey tea, sessile-leaved eupato- and continuous band covering millions of rium, and horse gentian. Two likely sa- acres in what is now Minnesota, Wisconsin, vanna specialists (purple milkweed and Iowa, Illinois, Michigan, Indiana, Ohio, wild hyacinth) are listed as endangered in Missouri, Arkansas, Oklahoma, and Texas. the state and three others (kitten tails, Historically, what is now Wisconsin was cream gentian, and Virginia lespedeza) are probably a leader in total acres of oak listed as threatened.

90 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Fortunately, most of the savanna species, especially the mammals, birds, reptiles, and amphibians, have readily adapted to the changed landscape, or they have managed to hang on and survive to this point in suboptimal habitat (e.g., the fringes of other less devastated communities such as oak forests). The success of the vertebrate animals has been due to the fact that major elements of the savanna structure are still well represented today in various “edge” habitats, including wooded pastures, lawns, and woodlots. The fact that the plant species may be different in those habitats has not affected savanna vertebrate species for the most part. species; and two others, the loggerhead This property in Many of the mammal species that Waukesha County were closely associated with our historical shrike and barn owl, are on the state’s shows what is thought oak savannas are still doing well today endangered species list (D. Sample and M. to be the typical tree (e.g., long-tailed weasel, cottontail rabbit, Mossman, Wis. Dep. Nat. Resour., pers. structure of oak comm.). Although loss of habitat has not openings. Since Euro- woodchuck, fox squirrel, red fox, and American settlement, white-tailed deer). However, others have been the cause of decline in all these oak openings have been either extirpated from the former species, it certainly is affecting many of almost disappeared them. The abandonment and loss of from the landscape savanna regions (e.g., timber wolf, bison, because of clearing, and elk) or reduced to very low numbers savanna/woodlot pastures in the past few plowing, overgrazing, (e.g., bobcat and black bear). The loss of decades may be playing a role in some of or suppression of fire these recent declines in savanna bird followed by invasion by these species, however, was due more to dense shrub and tree incompatibility with high human densities species. growth. As Curtis than to loss or degradation of the oak Most of the amphibian and reptile (1959) observed, savanna plant communities. Some mam- species that were closely associated with “Beyond question, an our historical oak savannas are still doing at oak savanna with an mals associated with the most open savan- intact groundlayer is nas (and the prairies) have not fared as well least moderately well today (e.g., Cope’s the rarest plant with the changes. For example, the least gray treefrog, five-lined skink, eastern community in hognose snake, smooth green snake, Wisconsin today.” shrew and the Franklin’s ground squirrel Photo by Eric Epstein. are of special concern in the state. western fox snake, eastern milk snake, and Most savanna bird species are still Dekay’s snake). However, two reptiles doing very well today (e.g., American associated with savanna habitat are suffer- robin, indigo bunting, blue jay, American ing from habitat loss. These are the western goldfinch, and brown thrasher). Only one slender glass lizard and the eastern massas- oak savanna bird, the passenger pigeon, has auga rattlesnake; both are now on the state become extinct, and another, the turkey, list of endangered species. Oak savanna was extirpated but restored; both of these sites may be important nesting sites for were lost to unregulated hunting rather turtle species such as the threatened than loss of habitat. However, a number of Blanding’s turtle in some areas, as agricul- savanna bird species have not thrived or ture continues to dominate open spaces have begun to decline in recent years (e.g., traditionally used for turtle nesting. black-billed cuckoo, northern flicker, red- Unlike the vertebrate communities, headed woodpecker, warbling vireo, vesper our knowledge of oak savanna invertebrates sparrow, bobwhite quail, and field spar- is very limited. We don’t know what species row). One species, the orchard oriole, is on were characteristic or restricted to the the state’s list of special concern; one, Bell’s community, let alone their current status. It vireo, is on the state’s list of threatened is likely that many species were lost or are now very rare.

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PROJECTED Invasion by aggressive exotics (i.e., honeysuckle, buckthorn, and reed In the absence of active management, canary grass). the future of oak savanna looks very bleak in Wisconsin and throughout its entire Increasing human population pressures, range. The increasing abandonment of often expressed as rural home and lightly to moderately grazed wooded suburban development. pastures and the accelerating succession of oak woodlots toward heavy-shade-producing trees and shrubs will lead to the decline and possible loss of much of what remains SOCIO-ECONOMIC ISSUES of the savanna flora and fauna, including eventual decline of the oaks themselves. Oak savanna was probably the optimum habitat for many game species (e.g., bobwhite quail, turkey, squirrels, deer, and rabbits). Thus, management for ACTIONS CAUSING CONCERN oak savanna is compatible with traditional Threats to the future survival of oak wildlife management and hunter interests. savanna can be summarized in five catego- The popularity of savanna songbirds, such ries. as bluebirds, should also lend public support to oak Loss of recovery savanna restoration. opportunities due Threats to the future survival of oak Light to moderate to savanna include the lack of knowledge cattle grazing can be about the community, the resistance to the compatible with ✓ accelerating use of prescribed fire, the lack of maintaining the plant forest succes- understanding of the importance of fire in structure needed by sion to dense- many savanna maintaining oak savanna, and increasing shade-produc- species. There is ing species, human population pressures, often support among expressed as rural home and private conservation ✓ lack of recruit- suburban development. groups for oak ment and savanna protection eventual die- and recovery; it is a out of long- high priority for The Nature Conservancy. lived plants in suboptimal habitat, However, the public in general lacks knowledge about savannas. ✓ increasing or decreasing grazing pressure, due to changes in pasturing practices. POTENTIAL FOR COMMUNITY RESTORATION General neglect and lack of knowledge about the community by the public, The recovery potential of oak savanna professional resource managers, and in Wisconsin is substantial (Holtz 1985; scientists. Bronny 1989; R. Henderson, Wis. Dep. Nat. Resour., unpubl. data). Degraded sites Resistance to the use of prescribed fire, in the dry and wet ends of the spectrum especially in wooded areas, and lack of can be recovered with relative ease. Mesic understanding by the public and profes- savannas with deep, rich soils will take sionals as to the importance of fire in more time and work, but recovery is still maintaining the state’s biodiversity. feasible. The pieces can still be found and put back together with a reasonable amount of effort (Packard 1988b). How-

92 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE ever, biological and socio-economic opportunities are gradually and steadily disap- pearing. Currently there are hundreds if not thousands of acres of overgrown but retrievable oak savanna on Department- managed lands. In addition there are probably thousands of acres of private land, both overgrazed and overgrown, with retrievable oak savanna. Much of this land, especially low productivity sites, could be restored within a decade or two simply by tree thinning, brushing, and burning. Well- drained, rich soil sites will require more work and time to restore. Some plant reintroduction may be necessary, but much can be accomplished with fire alone. Light education program is greatly needed for Fire is an essential component of savanna grazing may also have potential as a sa- developing support for its recovery and ecosystems. To vanna management tool and as a means of maintenance. The Department’s Bureau simulate wild fire, maintaining the open habitat required by of Parks and Recreation and the Bureau managers use of Information and Education should prescribed burning as many savanna vertebrates. Grazing, how- an important tool in ever, should not be considered the best play a major role in this effort. restoration of oak management tool for most savanna plants, openings and other 2. Develop a policy on prescribed burning fire-dependent although some may do well under light communities. Photo grazing. that recognizes the dependence of some from Department State ecosystems, including oak savanna, on Natural Area Files. fire and examines the resources and staff POSSIBLE ACTIONS support necessary to effectively and safely use fire to manage these fire- The following possible actions are dependent communities. In addition, air consistent with ecosystem management, quality standards and policies within the but require more analysis and discussion. Division of Environmental Quality will How priorities are set within this list will be need to be clarified. based on ecoregion goals, staff workload, fiscal resources, public input and support, 3. Pursue, as a high priority, protection and and legal authority. We will work with our maintenance of all high-quality rem- customers and clients to set priorities and nants (i.e., with high savanna species bring recommendations to the Natural richness and community integrity) and Resources Board for consideration begin- mildly degraded sites with high recovery ning in the 1995-97 biennium. potential. Small, high-quality sites should not be 1. Develop an Small, high-quality ignored, for they are education and probably the last sites should not be awareness pro- The recovery potential of oak savanna in refuge for many of ignored, for they gram to enhance Wisconsin is substantial. the savanna plants, are probably the public and profes- insects, and soil last refuge for sional appreciation microflora and of what oak many of the microfauna. Sites as small as a few acres savanna is, its past prevalence, its rapid savanna plants, may be contributing substantially to the decline and current rarity, and its genetic variation and survival of many insects, and soil management needs. Because of the species. This is a critical prerequisite to microflora and current rarity and long-time absence of the success of Action 5, below. microfauna. oak savanna on the landscape, an

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4. Provide buffer lands to these small, Southern District...... 45% high-quality sites. Buffer lands are Western District ...... 32% needed if remnant oak savannas are to Southeast District ...... 14% hang on to the species they have re- Lake Michigan District ...... 7% tained through 150 years of continual North Central District ...... 2% decline. Buffer lands provide remnants with protection against the negative To reach these recovery/restoration impacts of external influences and acreage goals, some acquisition and stochastic events and provide space into protection of private land will be which the community can expand and needed, but only for a limited number rebuild. Buffer lands should be restored of high-quality sites. Much can be done with communities that are compatible for oak savanna in Wisconsin without with the remnants. new land acquisition. Many opportunities exist for recovery on land already 5. Pursue recovery and restoration efforts managed by the Department, especially on as large, varied, and intact tracts as within state parks and wildlife areas. For are available. There should be several example, the Kettle Moraine State sites 1,000-5,000 acres or more in size. Forest-Southern Unit region is an area Habitat fragmentation issues should be with recovery potential on a large scale, considered in selecting candidate sites. and the Department’s Southern District Large tracts are needed because of the Headquarters grounds are a small but dynamic nature of oak savanna vegeta- highly visible site with exceptional tion, due to the educational potential. shifting mosaic of There are also oppor- sun and shade tunities to encourage Many opportunities exist for recovery on over time. The management for larger and more land already managed by the Department, savanna, or at least varied the restora- especially within state parks and components of it, on tion area, the wildlife areas. private lands through greater the tax incentives, likelihood that the educational programs, savanna community and its associated and the offering of technical advice, species will be able to maintain them- assistance, and partnerships. The selves in the long run. Habitat Restoration Areas component of the Wisconsin Stewardship Program 6. Just what total recovery/restoration may also provide some opportunities for acreage goal in the state would ensure regaining oak savanna. the long-term survival of the oak savanna community is unknown. Two to 7. Conduct research on oak savanna and three percent (110,000-165,000 acres) related oak woodland ecosystems of the original acreage may be a reason- regarding plant community association able target. This goal, of course, would and classification, effects of management include both public and private lands. on maintenance and recovery, and status Whatever the final acreage goal, it of rare species and remnants. should include representation of a variety of soil and topographic types as 8. Become an active partner in the Midwest well as geographic locations. Based on Savanna Ecosystem Recovery Plan to be the historical range of the community, proposed by the U.S. Environmental distribution of the acreage goal within Protection Agency. The plan will include Department Districts should be approxi- recommendations on research, inven- mately as follows: tory, management, and protection of Midwest savannas. This plan was first

94 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE discussed at the Midwest Oak Savanna 9. Encourage the establishment of sufﬁ- Conference held in Chicago (February cient sources of seeds and plant material 18-20, 1993), organized by the Illinois using local genotypes of oak savanna Chapter of The Nature Conservancy, the species. U.S. Environmental Protection Agency Region 5, and the College of Natural Resources, UW-Stevens Point.

Case Study

KETTLE MORAINE OAK OPENING: NATURAL COMMUNITY PROTECTION AND RESTORATION THROUGH MASTER PLANNING

Contributed by Mark Martin, Randy Hoffman, and Signe Holtz.

The Natural Resources Board approved the master plan for the Kettle Moraine State Forest in 1991 after a long planning process that included a Department task force, a vegetation management committee, a citizen’s advisory committee, various resource management specialists, citizens, and other organized groups. The state forest, as its name indicates, lies in the kettle moraine area of southeastern Wisconsin. Along the moraines in the Southern Unit are oak openings and oak woodland, and in the kettles and lowlands lie vast wetlands of prairie, fen, and sedge meadow. Dry prairies cover the southern- and western-facing hillsides. The Southern Unit also contains many populations of rare species (listed as endangered or threatened or of special concern), including 11 bird species, 18 plant species, seven insect species, and two mammal species. As the planning process progressed, it became apparent that this property could contribute greatly to the protection of Wisconsin’s natural heritage because it harbored degraded oak openings, one of the rarest natural communities in the state. As the largest block of public land in the southeast with more than 29,000 acres in the project boundary, it would also be one of the only opportunities in southeastern Wisconsin to restore an oak opening at the scale that it had occurred in the past. There were several sites with great restoration potential because of the existing tree structure and because surrounding public land ownership gave the Department the ability to manage effectively using prescribed burning. Out of this discussion came the proposal to create the Kettle Moraine Oak Opening, which would include the existing Blue Springs Oak Opening and three parts of the Messinger Dry Prairie and Savanna Preserve. The proposal became part of the master plan and since then the Department has been preparing the site for larger prescribed burns. First, crews have been removing buckthorn and honeysuckle, both non-native species, by cutting and using spot-herbicides. Second, they have burned small prairie patches to stimulate existing prairie plants to produce more seeds. This seed production, combined with the removal of the non-native shrub layer, should allow prairie to expand more easily across the site. Soon, the Department will burn much larger parts of the oak opening: 100-700 acres at a time, and at fairly short intervals (two or three years). As Randy Hoffman of the Department’s Bureau of Endangered Resources explains, “This is a 100-year work-in- progress.” As time goes by, the Department will examine the results, monitor restoration research, and change management as needed.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 95 OAK SAVANNA COMMUNITIES

LITERATURE CITED

Barry, A. T. and R. A. Spicer. 1987. The Holtz, S. L. 1985. Cutting and burning a evolution and palaeobiology of land degraded oak barrens: management plants. Croom Helm, London. 309 pp. techniques that simulate natural Bray, J. R. 1958. The distribution of savanna disturbance. M.S. Thesis, Univ. Wis.- species in relation to light intensity. Madison. Can. J. Bot. 36:671-81. Packard, S. 1988a. Rediscovering the Bray, J. R. 1960. The composition of tallgrass savanna of Illinois. Article savanna vegetation in Wisconsin. 01.14 in A. Davis and G. Stanford, eds. Ecology 41(4):721-32. The prairie: roots of our culture; foundation of our economy. Proc. 10th Bronny, C. 1989. One-two punch: grazing N. Am. Prairie Conf., Denton, Texas. history and the recovery potential of oak savanna. Restor. Manage. Notes Packard, S. 1988b. Just a few oddball 7(2):73-76. species: restoration and the rediscovery of the tallgrass savanna. Restor. Manage. Clewell, A. F. 1989. Observations Notes 6(1):13-20. corroborate savanna community concept. Restor. Manage. Notes 7(2):82. Penfound, W. T. 1962. The savanna concept in Oklahoma. Ecology 43(4):774-75. Cole, M. M. 1960. Cerrado, caatinga and pentanal: the distribution and origin of Pruka, B.W. 1994. Distribution of the savanna vegetation of Brazil. understory plant species along light and Geograph. J. 126:168-79. soil depth gradients in an upland oak savanna remnant in southern Curtis, J. T. 1959. The vegetation of Wisconsin. M.S. Thesis, Univ. Wis.- Wisconsin. Univ. Wis. Press, Madison. Madison. 271 pp. 657 pp. Johnson, R. W. and J. C. Tothill. 1985. Deﬁnition and broad geographic outline of savanna lands. pp. 1-13 in J. C. Tothill and J. J. Mott, eds. Ecology and management of the world’s savannas. Australian Acad. Sci. 384 pp.

96 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 97 OAK & PINE BARRENS COMMUNITIES

“Barrens” are plant communities that occur on sandy soils and are dominated by grasses, low shrubs, small trees, and scattered large trees. Curtis (1959) described these communities as pine barrens CHAPTER 7 in northern and central Wisconsin and as oak barrens in southern and west-central Wisconsin. Because of their dynamic nature and the variability in structural types and species composition, they are difficult to describe and classify. For example, Eric Oak and Epstein (Wis. Dep. Nat. Resour., pers. comm.) studied the original land survey notes for the area that is now southern Fort Pine Barrens McCoy and found that the surveyor had characterized the vegetation variously as “oak openings,” “oak brush,” “pine-oak woodland,” “pine brush,” “oak forest,” and Communities “level prairie.” Many northern pine barrens are referred to as “brush prairie.” This range by Ronald Eckstein of names derives from the many forms that North Central District barrens have. Prior to Euro-American settlement, the vegetative structure of large Department of Natural Resources barrens landscapes was quite variable and and dynamic. Inclusions of variously sized and Bruce Moss aged forest stands such as mature red pine, Northwest District mature oak (bur, red, Hill’s, or black), Department of Natural Resources aspen groves, and numerous wetlands were typical of most presettlement pine and oak barrens (Murphy 1931). One consistent element of all barrens, though, is the dependence of barrens on DESCRIPTION fire and the major role that fire plays in their dynamics. Fires have burned on he oak and pine barrens commu- Wisconsin barrens for thousands of years. nities of Wisconsin are two of the Prior to Euro-American settlement, fires four types of savanna described were caused by lightning or were set by by Curtis Native Americans. (1959). Native Americans Oak “Barrens” are plant communities that used fire to maintain opening is discussed occur on sandy soils and are dominated game habitat, drive in the previous by grasses, low shrubs, small trees, and game, and enhance chapter. Cedar glade, fruit and berry crops. scattered large trees. One consistent a very specialized Historically, behavior savanna, is not element of all barrens, is their of fire was greatly discussed in this dependence on fire and the major role it influenced by report. In this chapter plays in their dynamics. Fires have burned topography and soil the other two types of on Wisconsin barrens for thousands factors. Natural savanna, the closely of years. wildfires usually related oak and pine produce a complex barrens, are covered mosaic of burned along with bracken grasslands. and unburned

98 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE patches depending on fire intensity, topography, soil moisture, and local weather (Niemi and Probst, 1990). True savanna (widely scattered large trees over a prairie- like understory) was likely maintained by frequent fires of relatively low intensity. Brush prairie may have been subject to a more erratic fire regime with occasional catastrophic events that reduced the oaks to the grub stage. Because of this long association with fire, the plants and animals that live on the barrens are adapted to periodic fire. Vogl (1970) states: “The question of whether fire is necessary to maintain northern Wiscon- sin pine barrens is perhaps not an appro- barrens is the extraordinary development This is a typical pine priate question, for all factors including soil barrens in Florence type, soil fertility, topography, climate, of shrubs. This is . . . far higher than for County. The dominant drought, and fire are inseparably linked any other community in Wisconsin. Two tree is jack pine with and operate together or in chain reactions of the shrub species, redroot . . . and an understory huckleberry . . . , reach their maximum dominated by sweet and cannot be considered individually. Fire fern, hazelnut, and is one of the essential ingredients of pine Wisconsin levels in this community; but sedges. Also found barrens, but the the blueberry . . . is there are species in of even greater the heath family and critical factor in native grasses found determining the Pine barrens . . . are true savannas, in importance . . . . on poor soils, such as presence of barrens that the dominant plants are grasses, Another shrub which Kalm’s brome grass is highly character- and poverty oat grass. among northern forbs and shrubs, with scattered Photo by Eric Epstein. pine-hardwoods stands of trees. istic of the barrens is forests is not so much the sweet fern . . . . fire, but the presence The 134 [plant] of sandy plains; sites with low fertility that species found in the barrens are distrib- lend themselves to droughts and fires of the uted in 48 families, of which these five proper intensities and frequencies to contain over one-half of the total: produce a vegetational structure and Compositae-23.9 per cent, Gramineae- composition called barrens.” Much still 10.4 per cent, Rosaceae-8.2 per cent, needs to be learned about the relationships Liliaceae-6.7 per cent, and Ericaeae-6.0 between fire and barren structure and per cent . . . . [T]here is no doubt that the composition (Mossman et al. 1991). immediate cause of a pine barren is fire. In this case, soil and topography are PINE BARRENS major contributing factors, since it is essential that the fires be repeated at such Curtis (1959) describes pine barrens short intervals as to prevent the active as follows: reseeding of jack pine from its serotinal These barrens are true savannas, in cones. that the dominant plants are grasses, Vogl (1964a, 1970) studied northern forbs and shrubs, with a scattered stand Wisconsin pine barrens and found “the of trees. The most usual tree is jack pine, locations of northern Wisconsin pine although red pine may be the main barrens correlated with the distribution of species in unusual cases. Hill’s oak is sandy soils, great forest fires, present fire usually present as a grub or as a scatter- hazard areas, sites subject to local drought, ing of larger trees . . . . The outstanding the last strongholds of prairie grasses, and feature of the groundlayer in the pine areas of past farming failures and forest

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 99 OAK & PINE BARRENS COMMUNITIES

planting difficulties.” Vogl found that the Dunbar Barrens occur in Marinette County. pine barrens possess some characteristic Although we have no 19th-century descrip- plant species even though plant communi- tion of this site, in the 1960s LeRoy ties vary in different barrens. Prairie- Lintereur (Wis. Dep. Nat. Resour., retired, influenced pine barrens in far northwestern pers. comm.) found various species of and northeastern Wisconsin averaged 26 reindeer moss present, with sedges more more plant species than pine barrens in abundant than grasses. Sand willow and north-central Wisconsin. Prairie plants sand cherry were common. Barnes (1974) were present in the far northwest and quotes the description of the northwestern northeast, but absent in the north-central Wisconsin pine and oak barrens published pine barrens. Shively and Temple (1994) by E. T. Sweet in 1875: “The trees are either describe pine barrens as an open grassland scrub-pine or black-jack oak, averaging in with scattered trees and shrubs, i.e., a pine diameter about three or four inches and in savanna. They describe a pine-shrub- height not over fifteen feet. In some places, grassland ecosystem as a varying mosaic of as in the sand hills of the barrens, the trees vegetation structural types that occur on are at considerable distances from each sandy glacial outwash plains, developing other, and in other places the little scrub and deteriorating in response to periodic pines, not over two inches in diameter, are disturbance. so close together as to constitute a nearly Prior to Euro-American settlement impenetrable thicket. On the sides of the many pine barrens were diverse. Some barrens, and in low places, quite large resembled a pine savanna with mature red groves of norway pine are frequently pine occurring in densities of two to eight found.” trees per acre and an average diameter at breast height (dbh) of 13 inches. Early OAK BARRENS logging eliminated the mature trees. Severe, repeated fires, along with more cutting and Curtis (1959) describes oak barrens as land-clearing, removed the seedlings and follows: remaining red pine seed sources. Several of the early writers Several specific Wisconsin barrens mentioned that the bur oaks and white sites were described historically. Fassett oaks of the heavy soil openings were (1944) described barrens near the Brule replaced by black oaks on the sandy River in Douglas County in 1854 as a areas, but few detailed descriptions of the region of frequent fires, covered with small type exist. Most of the comments refer to jack pine and occasional large scattered red the jack pine barrens found on similar pine. Oak trees and oak brush often sites in the north. Actually the two types accompanied and sometimes replaced the are closely related and intermediate jack pines. Matthiae and Stearns (1978) mixtures of both oak and pine are and Vora (1993) recount historical records widespread in central and northwestern describing the Moquah Barrens in Bayfield counties. For purposes of this discussion, County in 1858 as a diverse landscape with oak barrens are considered to be those openings of various sizes, areas with savannas which have black oak or Hill’s scattered trees, some open forest and some oak as their most prominent tree and in closed-canopy forests about 60 years old. which jack pine is absent. As such, they Vogl (1964b) recounts historical descrip- are located entirely in the prairie-forest tions of Crex Meadows in Burnett County province south of the tension zone. They in 1853 as a jack pine—scrub oak—prairie are prominent on the outwash-filled savanna. The surveyors’ records refer to a valleys of the Wisconsin River from jack pine savanna consisting of large, open- Portage to Arena and the Sugar River in grown jack pines scattered across a level to Green County, and on the sandy uplands rolling landscape with some scattered red of Marquette and Waushara counties . . . pine and scattered areas of oak bushes. The . The origin of the scrub oak savannas is

100 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE the same as that of the oak openings— degradation of prior forests by fire. Maintenance is also by fire, but with the major difference that the tree component is likely to be completely destroyed at rather frequent intervals. Finley (1976) states: “Some minor portions of the oak region were in neither oak forest nor oak openings. These were the so-called oak barrens where thin stands of scrubby dwarf oak grew on sandy soils. The sparse growth appears to have been due more to the sandy earth material rather than to climatic influences. The uniqueness of the oak barrens resulted from the open spacing of the trees, the small size of the Holtz (1985) described a black oak This barrens has large, trees, and the otherwise barren character of barrens in Sauk County as a dynamic open-grown oaks with the surface. This type of vegetation oc- a sand-prairie community of trees, shrubs, and under- curred in small fragmented areas in Eau understory including story plants that is maintained through such species as lupine, Claire County, periodic fire. After little bluestem grass, eastern Dunn and June grass. If not decades without fire, County, and western subjected to fire, oak many understory barrens over time Chippewa County.” Oak barrens are considered to be those plant species persist become more like Barnes (1974) savannas which have black oak or Hill’s southern dry forest. as dwarfed, found the oak-pine oak as their most prominent tree and in Notice in the fore- nonreproducing ground the oak barrens of Eau Claire which jack pine is absent. As such, they culms and rhizomes seedlings and saplings County very hetero- are located entirely in the prairie-forest or as old seeds. If in the understory which geneous, with the over time may form a remnant barrens oak and pine gener- province south of the tension zone. more closed canopy. plants are on site, Photo by Cathy Bleser. ally forming a mosaic former barrens can be of separate stands of restored by a combination of cutting to various sizes. The Eau Claire County open the canopy and prescribed burning. barrens were probably open areas that contained few trees interspersed with rather dense stands of oak and pine. BRACKEN GRASSLANDS Habeck (1959) describes a general Curtis (1959), Vogl (1964b), and Levy picture of the vegetation in Juneau and (1970) identified bracken grasslands, Jackson counties prior to the turn of the sometimes called frost pockets, as a distinct century provided by Filibert Roth in 1898: vegetation type. Bracken grasslands are “Roth stated that much of the central large forest openings dominated by various Wisconsin sand plains was covered with grasses and bracken fern. Probably some of scrub oak and jack pine openings, with the original pine barrens of northern some portions covered with dense groves of Wisconsin included bracken grasslands. jack pine and a few islands of mature red Bracken grasslands occur on a variety of pine and white pine. Mesic upland hard- soils, from fine sands to loams. Bracken wood forests were apparently not present grasslands on loamy soils are thought to or not common enough to draw Roth’s originate after clearcutting and intense attention. Roth further stated that there wildfire. However, some bracken grasslands were extensive bare wastes which he on sandy soils may be natural communities believed were the result of logging and of the same nature and origin as the burning.” southern Wisconsin prairies (Curtis 1959).

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 101 OAK & PINE BARRENS COMMUNITIES

form is found on sandy soils and is charac- Bayfield Douglas terized by blueberries and sweet fern. Ashland Iron Exotic plants form a high content in both

Vilas forms. Washburn Sawyer Vogl (1964b) and McCaffery and Price Forest Florence Oneida Creed (1969) concluded that several factors Burnett Polk Marinette operate in combination to maintain Barron Rusk Lincoln Langlade bracken grasslands. Tree reproduction is Taylor Oconto Chippewa limited by frost, animal browsing, plant St. Croix Dunn Marathon Menominee allelopathy, dense sod, and a dense bracken Clark Shawano fern canopy cover. Pierce Eau Claire Door Pepin Wood Portage Waupaca Buffalo Outagamie Brown Jackson Kewaunee

Trempealeau STATUS Juneau Adams Waushara Winnebago Manitowoc Figure 16 Monroe

La Crosse Calumet Marquette Green Location of jack Lake Sheboygan PAST pine, scrub (Hill’s) Vernon Fond Du Lac oak forests and Sauk Columbia Dodge Pine barrens originally covered 2.3 Richland Washing- barrens, adapted ton Crawford million acres, or 7% of Wisconsin’s from Finley (1976). Ozaukee Dane presettlement landscape (Figs. 10 and 16). Iowa Jefferson Waukesha Note: There were Oak barrens covered 1.8 million acres, or also barrens known Grant Milwaukee Green Rock 5% of the presettlement landscape. on sandy river Lafayette Walworth Racine terraces along the Kenosha Mossman et al. (1991) state: “Prior to Mississippi River, settlement, barrens habitats were wide- Lower Wisconsin River, Levy (1970) recognized two forms of spread in Wisconsin, always associated Chippewa River, and Black River. bracken grassland. One form is found on with coarse-textured sandy or gravelly soils. loamy soils and is characterized by exotic The most extensive barrens were in large plants such as quack grass, Kentucky areas of sandy glacial outwash, or in the bluegrass, and Canada thistle. The other sandy beds of extinct glacial lakes, but they also occurred on river terraces, old dune Pine Barrens are also systems, gravelly moraine, and sandspits. found along the lower Wisconsin River and Geographically, areas of extensive barrens other rivers in the were concentrated in northeastern, north- Driftless Area. This central, northwestern and central Wiscon- barrens in Richland County along the sin. They were also common on the exten- Wisconsin River has sive outwash terraces along the Lower large jack pine and Wisconsin, Lower Chippewa and Missis- scrub oak with an sippi Rivers. In general, trees occurred in understory of sedges, prickly pear, blueberry, low density, usually as scattered individuals and some typical or in small groves, punctuating an open prairie species. Photo grassy landscape that was often dotted with by Signe Holtz. deciduous brush. Where outwash was pitted, the topography was more pronounced and varied and lakes and wetlands were sometimes frequent. In such areas, the pattern of vegetation was likely to be a mosaic of open prairie-like areas, brush, savanna, and occasional stands of deciduous, coniferous or mixed forest. The interplay of topographic and edaphic factors strongly inﬂuenced the behavior and effects of the primary disturbance factor affecting the barrens—ﬁre—and is

102 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE responsible for much of the structural, and compositional variability demonstrated by this community.” Because of the dynamic nature of barrens and their inherent variability, there is a general lack of knowledge of the exact structure of barrens. Some aspects of barrens that were described by early European explorers appear to have disappeared from today’s landscape. For example, some pine barrens were described as having large mature trees, either as widely scattered individuals or dense clusters of mature trees. Pine savannas with For example, a minimum of 10,000 acres This is the most intact scattered large trees are extremely rare. red pine savanna in of pine barrens has been recommended for Wisconsin. Located on long-term survival of an isolated sharp- the Lake Superior PRESENT tailed grouse population with limited shoreline of one of the hunting (Temple 1992). Apostle Islands, this Eric Epstein (Wis. Dep. Nat. Resour., site has an understory pers. comm.) summarized Natural Heritage of common juniper, native grasses and Inventory data and identified approxi- VEGETATION sedges, blueberry, mately 10,000 acres of pine and oak false heather, and barrens remaining at 65 sites (Table 5). Pine Barrens. Most of northern sand cherry. The groundlayer includes These figures do not include all of the pine Wisconsin’s pine barrens have succeeded to northern dry forest. Recently, Kotar et al. species that are and oak barrens in Wisconsin. The most characteristic of dune significant omissions (1988) published a and lakeshore are portions of the natural classification communities and system for northern many species not large managed Because of the dynamic nature of barrens found in Wisconsin barrens on county, and their inherent variability, there is a Wisconsin. This except near the Lake state, and federal system utilizes Superior shore. Photo general lack of knowledge of the exact by Signe Holtz. lands in northwest- interpretation of structure of barrens. Some aspects of natural vegetation ern Wisconsin and barrens that were described by early on the Necedah along soil moisture European explorers appear to have National Wildlife and nutrient gradi- Refuge in central disappeared from today’s landscape. ents with emphasis Table 5 Wisconsin. Some of on understory Remaining acreage of the managed barrens species. The follow- intact Wisconsin pine ing habitat types from the Kotar system can and oak barrens, 1992, are reclaimed forest or abandoned farmland as listed in DNR’s with reduced floristic compositions. The be used to describe the present status of Wisconsin Natural 1,432 acres of southern oak barrens at 20 former barrens in northern Wisconsin. Areas Inventory. sites is a fairly accurate estimate. The Natural Heritage Inventory lists Acreage pine barrens as G3 (very rare and local throughout its range or found locally) and oak barrens as G2 (imperiled globally Classification Pine Barrens Oak Barrens Total because of rarity) (see Table 1). Most remaining pine and oak barrens exist as Undisturbed 3,952 420 4,372 small, isolated fragments on about a dozen state or federal managed areas. Most of Moderately disturbed 3,421 280 3,701 these fragments are too small and isolated Disturbed 1,205 732 1,937 to ensure long-term viability of all their Total 8,578 1,432 10,010 characteristic native plants and animals. (45 sites) (20 sites) (65 sites)

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 103 OAK & PINE BARRENS COMMUNITIES

The pin oak/wintergreen-New Jersey maple, and balsam fir-white spruce. This tea forest habitat type occurs in Burnett, habitat type has more moisture, is more In central and Washburn, and Douglas counties. The mesic, and quickly succeeds to closed dominant landform is pitted outwash; the canopy forest. southern dominant soil is dry, nutrient-poor sand. Oak Barrens. In central and southern Wisconsin counties The following species are diagnostic: New Wisconsin counties the former barrens the former barrens Jersey tea, sweet fern, wintergreen, bush communities now support extensive pine communities now honeysuckle, cow-wheat, trailing arbutus, plantations, irrigation agriculture, or a support extensive bearberry, and sessile bellwort. Within this natural growth of dry oak forest. In rela- pine plantations, general habitat type the common forest tively undisturbed forests, prairie grasses irrigation types are jack pine, scrub oak forests and and forbs reappear if the forest cover is barrens, jack pine-pin oak, pin oak, aspen, clearcut and the logging slash burned agriculture, or a and red pine. (Holtz 1985). natural growth of The red oak-red maple/trailing Bracken Grasslands. In northern dry oak forest. arbutus forest habitat type which occurred Wisconsin the land area in bracken grass- in the former barrens in Marinette, lands has significantly declined. Fire Menominee, Oconto, Florence, Lincoln, control, tree planting, and aspen sprouting Oneida, and Vilas counties has been following clearcutting of adjacent forest replaced by forest cover of jack pine, red resulted in conversion of most bracken pine, aspen, and red oak-red maple. grasslands to balsam fir, white pine, and Understory vegetation includes bracken aspen. About 1% to 2% of the northern fern, grasses, sedges, blueberry, winter- public forest lands exist in forest openings green, and trailing arbutus. Low shrubs are or bracken grassland. A white-tailed deer more common than tall shrubs. Dry, habitat maintenance program undertaken nutrient-poor soils predominate. by the Department and the U.S.D.A. Forest Most remaining pine A white oak-pin oak/lead plant forest Service has maintained these small, scat- and oak barrens exist habitat type occurs in extreme northwest- tered bracken grasslands. as small, isolated ern Polk County and southwestern Burnett fragments—too small County. This habitat type appears to and isolated to ensure ANIMALS long-term viability of all represent a prairie-forest transition. Com- component plant and mon forest cover types include jack pine, Barrens are inhabited by animals that animal species. It is scrub oak forests, and barrens. estimated that the require open, brushy habitats. Large, open population of one Some barrens communities were barrens are critical habitat for sharp-tailed species, sharp-tailed located on the red maple-red oak/low sweet grouse (Hamerstrom and Hamerstrom grouse, shown here in blueberry habitat type, which occurred in a picture taken in 1942 1952, Gregg 1987); barrens large enough to in Wood County, would the former barrens of north-central and sustain a viable population of sharptails require a minimum of northeastern Wisconsin. The current will also sustain populations of other plants 10,000 acres of pine common forest cover types occurring on and animals common to large, open, barrens for long-term survival with limited this habitat type include aspen-white birch, brushy habitats. The particular structure of hunting. Photo by aspen-red oak, aspen-pines, jack pine, red each barrens will dictate the particular Dorothy Cassoday. pine, white pine, red oak, red oak-red complement of species present and their relative abundance. Jackson (1961) and Hamilton and Whitaker (1979) report that the following mammals find preferred habitat in barrens: thirteen-lined ground squirrel, plains pocket gopher, prairie deer mouse, coyote, badger, white-tailed deer, and striped skunk. Elk may have been another important species on barrens in presettlement times. Pierre Radisson described elk as fairly

104 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE abundant in parts of northwestern Wiscon- [3.1 ft]) woody vegetation: chipping, sin in 1658-60 (Seno 1985), but Schorger clay-colored, field, vesper, grasshopper (1954) gives only one literature reference and song sparrows, upland sandpiper, for elk in northwestern Wisconsin in the brown thrasher, bobolink, western 1800s (in the Superior area). Apparently meadowlark, Brewer’s blackbird, brown- elk were more abundant in the oak savan- headed cowbird and American goldfinch. nas, forest edge, and open woodlands of Common nighthawk is another species southern and central Wisconsin (Schorger common to open barrens and dry sand 1954). prairie. The relatively high abundance of Faanes (1981), Hoffman and Brewer’s blackbirds in open barrens is Mossman (1990), and Mossman et al. partly a consequence of its common (1991) described the birds of barrens and association with recently burned sites and young pine forest. Mossman et al. (1991) dead, fallen wood. In some cases, song describe the barrens bird community: sparrows also seem attracted to recently burned areas where remain charred The barrens is a tenuous commu- stems of shrubs and oak grubs. Nearly nity pulled in opposing directions by fire/ all of the species of Wisconsin’s dry frost and succession. The barrens prairies are well represented in open or avifauna responds to the variety and other types of barrens. Because these pattern of structures and dominant plant barrens are generally larger, and in forms in this dynamic community, and many cases more manageable than can be seen as a variable combination of southern Wisconsin’s isolated, dry prairie elements from related communities such relics, they serve an important role in as dry prairie (Sample and Hoffman maintaining this natural association of 1989), xeric pine and hardwood forest breeding-bird species, especially for those (Hoffman and Mossman 1990) . . . . Yet such as upland sandpiper that require barrens also represent a real natural large tracts. community with unique characteristics, and has undoubtedly been a major Vogt (1981) found Cope’s gray component of the upper Midwest landscape treefrog, American toad, five-lined skink, for centuries; thus it is not surprising that hognose snake, green snake and bullsnake several bird species appear to be especially common in Wisconsin pine and oak adapted to it. barrens. James Hoefler (Wis. Dep. Nat. Resour., pers. comm.) reports prairie skink Altogether, the most common and common in northwestern barrens. Eric regular species of Wisconsin pine and oak Epstein (Wis. Dep. Nat. Resour., pers. barrens are blue jay, common yel- comm.) reports the six-lined racerunner lowthroat, rufous-sided towhee, brown- and slender glass lizard present in west- headed cowbird, and the chipping, clay- central Wisconsin pine and oak barrens. colored, field, and vesper sparrows. Barrens habitats may be important nesting Other characteristic species that are sites for aquatic turtle species that lay their found here equally or more commonly eggs in upland, often sandy soils (Robert than perhaps in any other native Wis- Hay, Wis. Dep. Nat. Resour., pers. comm.). consin community include sharp-tailed In general, little is known about the grouse, upland sandpiper, northern invertebrates that occupy the barrens flicker, eastern kingbird, eastern bluebird, community or the ecological function they brown thrasher, Tennessee warbler, lark fulfill. An exception is the butterfly and sparrow, Brewer’s blackbird and Ameri- moth fauna, which has been extensively can goldfinch. studied by Ferge (1990). Based on his Open barrens are characterized by publication and his personal records, we dry sand prairie birds, most of which were able to compile a list of butterflies and tolerate or prefer some low (<1 m tall moths of barrens habitats (Table 6). Endan-

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 105 OAK & PINE BARRENS COMMUNITIES

Prairie fameflower ern and central Wisconsin, feed only on (Talinum rugospermum) occurs lupine. Midwestern populations of the in sand prairies within wide-ranging northern blue butterfly are barrens complexes in restricted by the distribution of its sole central and west larval food plant, the dwarf bilberry. These central Wisconsin. This individual is growing on species are known from very limited sand that has eroded locations in central and northeastern from a sandy rock Wisconsin (Ebner 1970). The phlox moth outcrop. Photo by Thomas A. Meyer. is found in Eau Claire County. The sand violet is found in west-central Wisconsin pine barrens. The rough white lettuce, phlox moth, and slender glass lizard have recent records at Fort McCoy (Eric Epstein, Wis. Dep. Nat. Resour., pers. comm.). Several singing males of the federally endangered Kirtland’s warbler have been located in west-central and northwestern Wisconsin pine barrens. Threatened Species. The frosted elfin butterfly is restricted to pine and oak barrens that contain large populations of gered, threatened, and other rare species lupine and false wild indigo (larval food are covered in more detail in the following plants). The wooly milkweed has historical paragraphs. records from sandy barrens near Necedah National Wildlife Refuge and other central RARE SPECIES Wisconsin oak barrens and sand prairies. The brittle prickly pear is found in oak Table 6 Endangered Species. Larvae of the barrens, dry cliffs, and sand prairies mostly Butterflies and moths Karner blue butterfly, a federally endan- in central and west-central Wisconsin. of barrens habitats.* gered species found primarily in northwest-

Species Range** Flight Status** Larval Host

Moths Saturniidae—Giant Silkmoths

Hemileuca nevadnesis Burnett & Sept local prairie willow, (Nevada buck moth) Douglas Co. poplar Sphingedae—Sphinx Moths Hemaris gracilis (graceful clearwing) N,C May-Jun rare blueberry Notodontidae—Prominents Hyparpax aurora (pink prominent) C Jun local oak Arctiidae—Tiger Moths Grammia celia CMay-Jun local unknown Pygarctia spraguei (Sprague’s pygarctia) C Jun local Euphorbia Noctuidae—Owlet or Noctuid Moths Schinia indiana phlox ﬂower moth Eau Claire Co. Jun Endangered Phlox pilosa Heliothis borealis N,C May local unknown

106 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Species Range** Flight Status** Larval Host

Butterflies Hesperiidae—Skippers Erynnis brizo (sleepy dusky wing) State May-Jun M oak Erynnis juvenalis (Juvenal’s dusky wing) State May-Jun 4 oak Erynnis persius (Persius dusky wing) C May 3 lupine Hesperia comma laurentina (Laurentian skipper) N Jul 3 grasses Hesperia leonardus leonardus (Leonard’s skipper) N,C Aug M grasses Hesperia Metea (cobweb skipper) N,C May M grasses Hesperia sassacus (Indian skipper) N,C Jun M grasses Atrytoropsis hianna (dusted skipper) W,C May-Jun M Andropogon Amblyscirtes vialis (roadside skipper) N,W May-Jul M grasses Pieridae—Whites and Sulphurs Euchloe olympia (Olympian marble) State May M rock cress Colias interior (pink edged sulpher) N,C Jun M blueberry Lycaenidae—Harvesters, Coppers, Hairstreaks and Blues Lycaena phlaeas americana (American copper) State May-Aug 4 rumex Harkenclenus titus (coral hairstreak) State Jul 4 cherry Satrium edwardsii (Edward’s hairstreak) W,E,C Jul M oak Incisalia augustinus (brown elfin) N,C May M blueberry Incisalia polia (hoary elfin) N,C May M blueberry Incisalia irus (frosted elfin) C May 3 lupine Incisalia henrici (Henry’s elfin) N,C May 3 blueberry? Incisalia niphon clarki (pine elfin) N,C May M jack pine Everes amyntula (western tailed blue) N May 3 vetch? Glaucopsyche lygdamus couperi (silvery blue) State May M lathyrus Lycaeidesidas nabokovi (northern blue) N Jul Endangered dwarf bilberry Lycaeides melissa samuelis (karner blue) C May-Aug Endangered lupine Nymphalidae—Brush-Footed Butterflies Charidryas gorgone carlota (Gorgone checkerspot) W,E,C May-Sep M Helianthus Phyciodes batesii (tawny crescent) N,C Jun M aster Satyridae—Satyrs and Wood Nymphs Oeneis chryxus strigulosa (chryxus arctic) N May 3 grasses

* From Ferge 1990, with additional information from 8 April 1991 Status: 3 = Resident, but rare and/or local in occurrence correspondence from Leslie Ferge to the author. 4 = Common or widespread ** Codes M = Not really rare, but not widespread or numerous enough to N = Northern Highland south to Tension Zone regard as common. C = Central Sands and Burnett County W = Driftless Area and Western Wisconsin E= Eastern Ridges and Lowlands

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 107 OAK & PINE BARRENS COMMUNITIES

Species of Special Concern. Ternate grape fern, common hairgrass, and possibly SOCIO-ECONOMIC ISSUES Hookers’ orchid may occur on or near Landowners and land managers often barrens in pockets of northern dry forest. Without active see barrens management as reducing their Prairie fameflower occurs in sand prairies restoration and ability to grow commercial trees. Some within barrens complexes in central and management the people find the barrens an exceptionally west-central counties. In addition, there are barrens community aesthetic landscape where native plants and some Great Plains plant species that reach animals have adapted to the very poor site will probably their eastern range limits in the Polk, nutrient quality and open character. disappear from all Douglas, and Burnett county barrens Wisconsin’s former barrens landscapes but a few large (Robert Read, Wis. Dep. Nat. Resour., pers. could be used by citizens for a variety of comm.). public land areas products and purposes. These include and a handful of wood fiber, food, game animals (including ROJECTED small, isolated P sharp-tailed grouse), and native plant and managed areas. Without active restoration and animal populations. A combination would management the barrens community will best meet the overall needs of Wisconsin’s probably disappear from all but a few large citizens. public land areas and a handful of small, isolated managed areas. POTENTIAL FOR COMMUNITY RESTORATION ACTIONS CAUSING CONCERN Mossman et al. (1991) state: Since Euro-American settlement, the “Wisconsin’s oak and pine barrens evolved pine and oak barrens communities have in a dynamic landscape governed largely by been reduced to small scattered parcels the forces of succession, fire and frost. with a simplified vegetative structure and a Variety of this landscape was imparted by reduced composition of plants and animals. the variable influences of climate, topogra- Control of wildfire, forest succession, pine phy, soils, moisture regimes and fire plantation development, and agricultural barriers. Despite the neglect and abuse that development have all most barrens have worked to bring the undergone since barrens communities Despite the neglect and abuse that most settlement, this is one to their current rarity. barrens have undergone since settlement, of our most resilient Some citizens this is one of our most resilient natural natural communities, question the neces- communities, and it will respond to careful and it will respond to sity and value of careful management management by controlled [prescribed] timber cutting and by controlled [pre- prescribed burning, burns and cutting. scribed] burns and management tech- cutting. Moreover, its niques used to economic land value restore barrens. Air quality standards is generally low, and comparatively little applicable to prescribed burning need to be has been permanently converted to other clarified. Some land managers and citizens uses. Perhaps for no other native commu- consider the barrens landscape as, indeed, nity are the opportunities for large-scale barren or worthless. The great habitat restoration so great.” values and aesthetic appeal of barrens Department wildlife areas in north- remain largely unrecognized. western Wisconsin, the Necedah National Wildlife Refuge, and the Chequamegon National Forest’s Moquah Barrens form the nucleus for large landscape-scale barrens

108 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE management. Barrens can be restored Shively and Temple (1994) describe a through cutting and prescribed ﬁre (Holtz pine-shrub grassland restoration plan with 1985, Vora 1993, U.S. Fish and Wildlife goals of increasing area and decreasing Service 1994) or through a combination of fragmentation of barrens, restoring the full cutting, limited herbicide use, and pre- natural range of variation in structure and scribed ﬁre (Paul Kooiker and Mike composition of patch types in barrens Zeckmeister, Wis. Dep. Nat. Resour., pers. landscapes, and obtaining statewide comm.). support and involvement of state residents.

POSSIBLE ACTIONS

The following possible actions are Douglas County Wildlife Area. consistent with ecosystem management, Expansion could include about one but require more analysis and discussion. section (640 acres) of additional land. How priorities are set within this list will be based on ecoregion goals, staff workload, Encourage the Chequamegon Na- ﬁscal resources, public input and support, tional Forest to continue to enlarge and legal authority. We will work with our Moquah Barrens. Support creation of customers and clients to set priorities and a large barrens landscape around the bring recommendations to the Natural Moquah Barrens Wildlife Area Resources Board for consideration begin- including old-growth jack pine, ning in the 1995-97 biennium. scattered red pine, frost pockets, seepage lakes, and small wetlands. 1. Restore several large pine and oak barrens communities. These actions may Continue restoration and manage- include the following: ment of a large oak barrens at Necedah National Wildlife Refuge. Continue development of Fish Lake Wildlife Area in Burnett County to Work with local ofﬁcials in Adams establish large barrens and savanna. and Juneau counties and Monroe and Jackson counties to explore the Continue management of Crex opportunity for restoration of large Meadows and Kohler Peet areas. barrens. Because of its geographic Explore opportunities to work with location, this area supports a far Burnett County to manage county greater diversity of barrens plants and forest land between the two areas for animals than other areas of the state, a connecting corridor of ecologically including more threatened and functional barrens habitat. endangered species. Continue restoration and expansion 2. Restore smaller barrens to conserve of barrens habitat within the ap- plant and animal species diversity. For proved Namekagon Barrens Wildlife example, Area. The goal will be restoration and maintenance of an extensive barrens Continue the development of suitable landscape that includes various barrens habitat on Amsterdam barrens types. Sloughs Wildlife Area. Work with Douglas County to Manage the Dunbar Wildlife Area to expand and continue development of restore barrens.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 109 OAK & PINE BARRENS COMMUNITIES

Manage the Spread Eagle Wildlife 4. Maximize the connectivity of pine and Area to restore a barrens landscape oak barrens restorations using tech- that includes wetlands, Pine River niques that are found to be effective by shoreline, and seepage lakes. current conservation research. These may include the establishment of Continue restoration and manage- corridors of open land between barrens ment of oak barrens at the Sandhill habitats. These corridors need to be Wildlife Area. carefully planned to avoid unintended effects such as species traps and the Work with the Nicolet National introduction of exotic species. Utility Forest to explore opportunities for rights-of-way, railroads, or short rotation restoration of barrens in the Lake- timber harvest could be used as corri- wood District. dors.

Work with federal Department of 5. Develop an education and awareness Defense officials to restore an oak program to increase public and profes- barrens at Fort McCoy, Monroe sional knowledge and appreciation of County. what barrens are, their past prevalence, their current rarity, and their manage- Protect and manage scattered small ment needs. Because of the barrens barrens to enhance populations of rarity, an education program is needed locally rare plant and animal species. to develop support for restoration and 1 Included here are the /4-mile-wide management of barrens. fire breaks maintained on state and county forests in northwestern 6. Develop a policy on prescribed burning Wisconsin and tracts along the Lower that recognizes the dependence of some Wisconsin River. ecosystems, including barrens, on fire and that examines the resources and Maintain and restore the scattered staff support necessary to effectively and bracken grasslands on public lands in safely use fire to manage these fire- northern Wisconsin. dependent communities. In addition, air quality standards and policies within the 3. Restore and manage pine and oak Department’s Division of Environmental barrens with a variety of structures Quality will need to be clarified. including brush prairie, pine savanna, older jack pine stands, and mature red 7. Gather more information on the struc- pine pockets. The barrens landscape ture and composition of barrens that should also contain grassland, frost existed at various times in the past, pockets, wetlands, and lakes. This drawing on a variety of sources, to landscape will best meet the needs of a describe the full range of variability of wide variety of barrens plants and these communities. animals.

110 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Case Study

HABITAT CONSERVATION PLANNING: SPECIES PROTECTION THROUGH ECOSYSTEM MANAGEMENT

Contributed by Cathy Bleser, Darrell Zastrow, and Signe Holtz.

Since the federal government listed the Karner blue butterfly as federally endangered in 1992, the Department has been working on its protection and recovery. At present, a Rangewide Federal Recovery Plan is being developed, and in Wisconsin, where the Karner blue butterfly is most abundant and widespread, a Habitat Conservation Plan (HCP) is underway. A HCP is required for any incidental taking of the insect to be permitted by the U.S. Fish and Wildlife Service. This HCP approach calls for working with an extensive group of public agencies, private businesses, and nonprofit conservation organizations to develop a plan for conservation of Karner blue habitat. Although the butterfly lives in several different natural communities that contain its larval food plant, lupine, in Wisconsin it is found most often on barrens. In addition, barrens are home to many other state and federally listed species such as phlox moth, massasauga rattlesnake, prairie fameflower, and frosted elfin butterfly. Although the Karner blue butterfly is the catalyst for the planning process, the Depart- ment will expand the focus to the barrens ecosystem on Department lands and encourage ecosystem considerations across the extensive range of the butterfly in Wisconsin. It has been found on public and private land across central and northwestern Wisconsin. In order to maintain this extensive range, it was apparent that there would be many players The Karner blue with an interest in this plan, including U.S. Fish and Wildlife Service; U.S. Department of butterfly, a federally endangered species, Defense; U.S. Environmental Protection Agency; county forests; Sand County Foundation; occurs most often in Georgia-Pacific Corporation; Consolidated Papers, Inc.; Wisconsin Department of Agricul- Wisconsin on barrens. ture, Trade and Consumer Protection; utility companies; railroad commissions; Menominee Here it rests on black- eyed Susan (Rud- Indian Nation; Winnebago Indian Nation; and Wisconsin Woodland Owners Association. beckia hirta) in Eau The proposed plan would emphasize processes that maintain the shifting barrens areas Claire County. Photo by Eric Epstein. across the landscape such as fire, cutting, and mowing, rather than relying solely on permanent protection of fixed parcels of land. “We want a plan that integrates conservation and economic land use in a manner that is both ecologically and economically sound. And we want to involve landowners and other individuals and groups that have interests in both using and protecting this habitat,” explains Cathy Bleser of the Department’s Bureau of Endangered Resources. The plan would integrate Karner blue butterfly conservation with existing land uses and would identify ways for landowners to carry out activities on their lands that will avoid or minimize harm to the butterfly and even provide additional areas of barrens for the butterfly to colonize. The plan would take a landscape-scale approach to conservation and would focus restoration efforts on those areas where the best opportunities exist. The planning process was proposed to the U.S. Fish and Wildlife Service in April 1994, and the Department has initiated discussions with many groups. A goal of July 1997 was agreed upon as the target date for a final plan.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 111 OAK & PINE BARRENS COMMUNITIES

LITERATURE CITED

Barnes, W. J. 1974. A history of the Hoffman, R. M. and M. J. Mossman. 1990. vegetation of Eau Claire County, Birds of northern Wisconsin pine Wisconsin. Trans. Wis. Acad. Sci. Arts forests. Passenger Pigeon 52(4):339-55. and Lett. 62:357-75. Holtz, S.L. 1985. Cutting and burning a Curtis, J. T. 1959. The vegetation of degraded oak barrens: management Wisconsin. Univ. Wis. Press, Madison. techniques that simulate natural 657 pp. disturbance. M.S. Thesis. Univ. of Dunn, C. and F. Stearns. 1980. Vegetation Wisconsin. Madison. 111 pp. of the Moquah Barrens Research Jackson, H. T. 1961. Mammals of Natural Area. Unpubl. rep. to N. Cent. Wisconsin. Univ. Wis. Press. 504 pp. For. Exp. Stn. Coop. Aid Study. Kotar, J., J. A. Kovach, and C. T. Locey. Ebner, J. A. 1970. Butterflies of Wisconsin. 1988. Field guide to forest habitat types Milwaukee Public Mus. Pop. Sci. of northern Wisconsin. Univ. Wis.- Handb. No. 12. 205 pp. Madison Dep. For. and Wis. Dep. Nat. Faanes, C. A. 1981. Birds of the St. Croix Resour. River Valley: Minnesota and Wisconsin. Levy, G. F. 1970. Phytosociology of U.S. Dep. Inter. N. Am. Fauna No. 73. northern Wisconsin upland openings. 196 pp. Am. Midl. Nat. 83:213-37. Fassett, N. C. 1944. Vegetation of the Brule Matthiae, P. E. and F. W. Stearns. 1978. basin, past and present. Trans. Wis. Ecological analysis of Moquah Barrens Acad. Sci., Arts and Lett. 36:33-56. Wildlife Management Area. Univ. Wis- Ferge, L. A. 1990. Checklist of Wisconsin Milwaukee. [unpubl. rep.] butterflies. Wis. Entomol. Soc. Misc. McCaffery, K. R. and W. A. Creed. 1969. Publ. No. 1. 4 pp. Significance of forest openings to deer Finley, R. W. 1976. Interpretation of map of in northern Wisconsin. Wis. Dep. Nat. the original vegetation of Wisconsin. Resour. Tech. Bull. No 44. 104 pp. U.S. For. Serv. N. Cent. For. Exp. Stn., Mossman, M. J., E. Epstein, and R. M. St. Paul. Hoffman. 1991. Birds of Wisconsin Gregg, L. 1987. Recommendations for a pine and oak barrens. Passenger Pigeon program of sharp-tail habitat 53(2):137-63. preservation in Wisconsin. Wis. Dep. Murphy, R. E. 1931. The geography of the Nat. Resour. Res. Rep. 141. 24 pp. northwestern pine barrens of Habeck, J. R. 1959. A phytosociological Wisconsin. Trans. Wis. Acad. Sci., Arts study of the upland forest communities and Lett. 26:69-120. in the central sand plain area. Trans. Niemi, G.J. and J.R. Probst. 1990. Wildlife Wis. Acad. Sci., Arts and Lett. 48:31- and fire in the Upper Midwest. in J.M. 48. Sweeney (ed). Management of Dynamic Hamerstrom, F. and F. Hamerstrom. 1952. Ecosystems. Proc. of a Symposium. 51st Sharp-tails into the shadows? Wis. Midwest Fish and Wildl. Conf., Conserv. Dep. Wis. Wildl. No. 1. 35 pp. Springfield, Illinois. 5 Dec 1989. 180pp. Hamilton, W. J. and J. O. Whitaker. 1979. Mammals of the Eastern United States. Sample, D. W. and R. M. Hoffman. 1989. Cornell Univ. Press. 346 pp. Birds of dry-mesic and dry prairies in Wisconsin. Passenger Pigeon 51(2): 195-208.

112 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Schorger, A. W. 1954. The elk in early Wisconsin. Trans. Wis. Acad. Sci., Arts ADDITIONAL READING and Lett. 43:5-23. Anderson, R. D. and L. E. Brown. 1983. Seno, W. J., ed. 1985. Up county: voices Comparative effects of ﬁre on trees in a from the Midwestern wilderness. Midwestern savannah and adjacent Round River, Madison. 239 pp. forest. Bull. Torrey Bot. Club 1:87-90. Shively, M.M. and S.A. Temple. 1994. An Blewett, T. 1976. Prairie and savanna ecosystem recovery plan for Wisconsin restoration in the Necedah National pine-shrub-grassland ecosystems (pine Wildlife Refuge. pp. 154-57 in Proc. barrens). Univ. of Wisconsin. Madison. Fifth Midw. Prairie Conf. Aug. 22-24, 48 pp. 1976. Iowa State Univ., Ames. Temple, S.A. 1992. Population viability Bray, J. R. 1960. The composition of analysis of a sharp-tailed grouse savanna vegetation in Wisconsin. metapopulation in Wisconsin. in Ecology 41:721-32. Wildlife 2001:populations. D.R. Brewer, L. 1989. A survey of the oak McCullough and R.H. Barrett. (eds). barrens and savannas of the Oak Elsevier Applied Science, New York. Openings Preserve Metropark. Nat. 1163 pp. Conservancy, Columbus, Ohio. [unpubl U.S. Fish and Wildlife Service. 1994. rep.] Barrens restoration and management Collins, S. L. and R. E. Good. 1987. plan. Necedah National Wildlife Canopy-ground layer relationship of Refuge. Unpublished Manuscript. oak-pine forests in the New Jersey pine 136pp. barrens. Am. Midl. Nat. 117(2):280-88. Vogl, R. J. 1964a. The effects of ﬁre on the Forman, R. T. (ed). 1979. Pine barrens: vegetational composition of bracken- ecosystem and landscape. Academic grasslands. Trans. Wis. Acad. Sci., Arts Press. New York. 601 pp. and Lett. 53:67-82. Gratson, M. W., J. E. Toepfer, and R. K. Vogl, R. J. 1964b. Vegetational history of Anderson. 1990. Habitat use and Crex Meadows, a prairie savanna in selection by male sharp-tailed grouse, northwestern Wisconsin. Am. Midl. Tympanuchus phasianellus campestris. Nat. 72:157-75. Can. Field-Nat. 104(4):561-66. Vogl, R. J. 1970. Fire and the northern Grimm, E. C. 1984. Fire and other factors Wisconsin pine barrens pp. 175-209 in controlling the big woods vegetation of Proc. Annu. Tall Timbers Fire Ecology Minnesota in the mid-nineteenth Conf. No. 10. century. Ecol. Monogr. 54(3):291-311. Vogt, R. C. 1981. Natural history of Hamerstrom, F. N. 1963. Sharptail brood amphibians and reptiles in Wisconsin. habitat in Wisconsin’s pine barrens. J. Milwaukee Public Mus. 205 pp. Wildl. Manage. 27(4):743-802. Vora, R.S. 1993. Moquah barrens, pine Hamerstrom, F. N. and F. Hamerstrom. barrens restoration experiment initiated 1951. Mobility of the sharp-tailed in Chequamegon National Forest. grouse in relation to its ecology and Restoration and Manage. Notes distribution. Am. Midl. Nat. 46(1):174- 11(1):39-44. 226. Payne, N.F. and F.C. Bryant. 1993. Techniques for wildlife habitat management of uplands. McGraw-Hill. New York. 500 pp.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 113 OAK & PINE BARRENS COMMUNITIES

Sando, R. W. 1967. The effects of repeated Vanderschaegen, P. V. 1977. Status of short- spring burning on the oak stands of the tailed grouse in Wisconsin, 1975. Wis. Cedar Creek Natural History Area. Dep. Nat. Resour. Res. Rep. 95. 9 pp. Univ. Minnesota. M.S. Thesis. Vogl, R. J. 1965. Effects of spring burning Schneberger, E. and A. D. Hasler. 1954. The on yields of brush prairie savanna. J. Brule River, Douglas County. Wis. Range Manage. 18(5):202-05. Conserv. Dep. 255 pp. Schwegman, J. E. and R. C. Anderson. 1984. Effect of eleven years of ﬁre exclusion on the vegetation of a southern Illinois barren remnant. pp. 146-48 in The Prairie: Prairie Conf.

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of these communities are (1) lack of trees and tall shrubs and (2) dominance by graminoid (grass and sedge) species. Wisconsin’s grasslands are at the periphery of North America’s extensive CHAPTER 8 mid-continental grassland biome which lies south and west of the state. Historically, Wisconsin grasslands were maintained primarily by frequent fires, as was most of the North American grassland biome. Grassland Treelessness is generally the optimum state for maximum development and health of these grassland systems. Although grasses and sedges dominate vegetative Communities biomass in this community type, forbs (non-graminoid wildflowers) dominate the by Richard Henderson species composition. The most represented Bureau of Research families of forbs are the composite (aster), legume, milkweed, carrot, and rose fami- Department of Natural Resources lies. Over 400 species of native vascular and plants are characteristic of Wisconsin David Sample grasslands, and most of these are restricted Bureau of Research to grassland or grassland/savanna commu- Department of Natural Resources nities. Detailed descriptions of Wisconsin’s grassland plant communities can be found With contributions from: Eric Epstein, Randy Hoffman, June in the classic text by Curtis (1959). Wis- Dobberpuhl, Thomas Meyer, and Mark Martin, Bureau of consin grasslands also have a diverse and Endangered Resources, and Mike Mossman, Bureau of specialized fauna, especially among the Research invertebrates, herptiles, and birds. Prairie (French for “meadow”) was the only word early French explorers had to describe the extensive, treeless, and grass- covered landscapes of central North America. Prairie subsequently became the DESCRIPTION term used to describe the grassland type most prevalent in Wisconsin prior to Euro- n this document, the term grassland American settlement. Prairie in Wisconsin refers collectively to several native was located mostly in the southern and Wisconsin plant western parts of the communities. state. It occurred These include Prairie (French for “meadow”) was the across a wide range prairie, brush only word early French explorers had to of topographies, soil prairie (i.e., prairie describe the extensive, treeless, and types, and soil with oak grubs and grass-covered landscapes of central North moisture regimes. shrubs less than six America. Prairie subsequently became the This variety of feet tall), sand bar- edaphic conditions term used to describe the grassland type rens, bracken- resulted in a great grassland, fen, and most prevalent in Wisconsin prior to Euro- diversity of prairie sedge meadow American settlement. flora. (southern and Fen is a highly northern) as defined restricted type of wet by Curtis (1959). Common characteristics prairie that supports an unusually special-

116 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE ized flora. It forms on wet to moist and often peaty, calcareous soils that have developed over a diffuse groundwater discharge area that is often under artesian pressure. Sedge meadow is at the extreme wet end of the prairie continuum and was the second most common grassland type in the state. It is distinguished from wet prairie by having (1) more sedge than grass vegetation, (2) more organic than mineral soil, and (3) seasonally standing water. It also supports a less diverse flora than wet prairie. Bracken-grassland was the northern version of prairie and was found mostly north of the tension zone, which is a band millions of years. Originating in the rain Northern sedge meadow in Douglas 10-30 miles wide, running from the shadow that developed with the uplifting of County. Structurally, northwest to southeast corners of the state, the Rocky Mountains, they have been sedge meadow is a separating the two expanding and grassland but contracting with hydrologically, it is a major floristic wetland. Photo by Eric provinces of Wiscon- North America’s mid-continental major climatic Epstein. sin (Curtis 1959). It grasslands have been in existence for changes ever since. They made their most was not abundant millions of years . . . . Original land survey historically. Although recent incursion into records of the 1830s indicate there were similar to prairie in what is now Wiscon- structure, bracken- 3.1 million acres of treeless grassland in sin approximately grassland is floristi- Wisconsin, or 9% of the total land cover five to six thousand cally very different (Curtis 1959) . . . . Tallgrass prairie and years ago and re- (Curtis 1959:314- related oak savanna are now the most mained relatively stable here until 21), with bracken decimated and threatened plant fern being a domi- Euro-American communities in the Midwest and nant species. This settlement in the limited vegetation in the world. mid-1800s. Original type is covered in the land survey records “Oak and Pine of the 1830s indicate Barrens” section. there were 3.1 million acres of treeless Sand barrens is also a limited grassland grassland in Wisconsin, or 9% of the total type. It is similar to dry sand prairie, but land cover (Curtis 1959). A little over two- has far sparser vegetation, and it generally thirds of this open land (2.1 million acres) includes exposed sand or sandblows. Most was prairie, and approximately one-third sand barrens today are artifacts of post- (1 million acres) was sedge meadow (see Euro-American-settlement activity, prima- Fig. 10). rily failed attempts at agriculture. PRESENT Over the past 150 years, the mid- STATUS continental grassland biome has been greatly reduced and degraded throughout PAST its range. Most grassland acreage has North America’s mid-continental suffered one of the following fates: (1) grasslands have been in existence for conversion to crop production, (2) over-

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 117 GRASSLANDS COMMUNITIES

grazing, or (3) invasion by shrubs and trees Midwest and among the most decimated in due to lack of fire, lack of grazing, or both. the world. With productive soils and ample precipita- According to the State Natural Heri- tion, the eastern portion of the grassland tage Inventory, Wisconsin has only 0.5% Table 7 biome (including Wisconsin), known as (13,000 acres) of its original grassland Rare and declining tallgrass prairie, was thoroughly fragmented ecosystem remaining in a relatively intact Wisconsin grassland and almost totally converted to agricultural condition, and much of this remnant plants. Compiled from use. Tallgrass prairie and related oak acreage has been degraded to some degree Wisconsin Department of Natural Resources savanna are now the most decimated and by livestock grazing or woody invasion. (1992). threatened plant communities in the Over 80% (11,000 acres) of this remaining acreage is sedge meadow, and the rest (2,000 acres) is native prairie. However, the Status Scientific Name Common Name inventory is not nearly as complete for sedge meadow as it is for prairie; there are many acres of secondary and small tract Extinct (none) sedge meadows not included in the acreage Extirpated Asclepias meadii* Mead’s milkweed total. Platanthera blephariglottis white-fringed orchid These remnants represent only 1.1% Endangered Agalinis skinneriana** pale false foxglove and 0.1% of the original sedge meadow and Anemone caroliniana Carolina anemone prairie acreage, respectively. Most of the Anemone multifida Hudson Bay anemone surviving prairie is either dry or wet; the Astragalus crassicarpus prairie plum intermediate type (mesic prairie), once the Astragalus neglectus Cooper’s milk vetch most common type in the state, is now Fimbristylis puberula chestnut sedge virtually gone. Only about 100 acres * Lespedeza leptostachya prairie bush clover (0.01%) of an original million acres of Liatris punctata dotted blazing star mesic prairie are known to exist, and these Parnassia parviflora small-flowered grass-of-parnassus are in small (often linear), scattered parcels Phlox glaberrima smooth phlox of a few acres at best. Platanthera leucophaea* prairie white-fringed orchid Wisconsin’s grassland plants and Polygala incarnata pink milkwort animals responded to the changes that Prenanthes crepidinea great white lettuce came with Euro-American settlement in Prenanthes aspera rough white lettuce various ways. Some species adapted well Ruellia humilis wild petunia and maintain healthy populations today, Scirpus cespitosus tussock bulrush while some are persisting only in low Scutellaria parvula small skullcap numbers. Others are restricted to prairie Threatened Agastache nepetoides yellow giant hyssop and sedge meadow remnants, and a few Agalinis gattingeri round-stemmed false foxglove have been extirpated. Asclepias lanuginosa wooly milkweed An estimated 15%-20% of the state’s Asclepias sullivantia prairie milkweed original grassland flora is now considered Cacalia tuberosa prairie Indian plantain rare in the state. Seventeen species are ** Cirsium hillii prairie thistle currently on Wisconsin’s endangered ** Cypripedium candidum white lady-slipper species list; 17 species are on the threat- Echinacea pallida pale purple coneflower ened species list; and 29 species are of Eleocharis rostellata beaked spike-rush Hypericum sphaerocarpum round-fruited St. John’s wort special concern in the state (Table 7). This Lesquerella ludoviciana bladderpod pervasive rarity among grassland plants is Opuntia fragilis brittle prickly-pear due to the extensive loss of the original Parnassia palustris marsh grass-of-parnassus grassland sod and the conservative nature Parthenium integrifolium wild quinine of many grassland plants, which are rarely Polytaenia nuttallii prairie parsley found outside of native vegetation rem- Platanthera flava tubercled orchid nants. Some, such as prairie gentian and Tofieldia glutinosa false asphodel hoary puccoon, are so conservative that Continued on next page

118 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE they are rarely if ever successful in restoration attempts. Status Scientific Name Common Name The current rarity of many of these species is not limited just to Wisconsin but Special Agoseris cuspidata prairie dandelion Concern Aristida dichotoma poverty grass is also characteristic throughout their Artemisia frigida prairie sagewort range. Three Wisconsin species, Mead’s Callirhoe triangulata poppy mallow milkweed, prairie bush-clover, and prairie Carex richardsonii Richardson’s sedge white fringed orchid, are on the federal list Carex suberecta prairie straw sedge of threatened species, and six others, Carex torreyi Torrey’s sedge prairie thistle, glade mallow, tubercled Cassia marilandica Maryland senna orchid, prairie fame-flower, pale false Dasistoma macrophylla mullein foxglove foxglove, and eared false foxglove, are Eleocharis robbinsii Robbin’s spike-rush being considered for federal listing. Eleocharis wolfii wolf spike-rush Most of Wisconsin’s grassland verte- Gentianopsis procera small fringed gentian brates adapted to the changes in the land. Houstonia caerulea bluets Noted exceptions are the extirpated mega- Liatris spicata marsh blazing star Napaea dioica glade mallow fauna (i.e., bison, elk, and wolves), and Oenothera serrulatus toothed evening primrose smaller, specialized animals such as the Orobanche ludoviciana Louisiana broomrape ornate box turtle and the long-billed Orobanche uniflora one-flowered broomrape curlew. Species that did adapt made use of Panicum wilcoxianum Wilcox’s panic grass croplands, pastures, old fields, roadsides, Penstemon hirsutus hairy beardtongue and other highly altered, surrogate “grass- Penstemon pallidus pale beardtongue lands.” However, in the past few decades Petalostemum villosum villous prairie clover even these areas have declined in acreage Physalis grandiflora white ground cherry and quality due to changing agricultural Polygala cruciata cross milkwort practices and land use (e.g., increased use Psoralea argophylla silvery scurfy pea of pesticides, extensive conversion of small Psoralea esculenta pomme-de-prairie grain and pasture acreage to row crops, and Solidago ohioensis Ohio goldenrod Talinum rugospermum** prairie fame-flower changes in the nature and timing of agricul- Tomanthera auriculata** eared false foxglove tural disturbances, including the early and frequent mowing of alfalfa) and invasion by * Federally threatened. ** Concern at federal level. woody growth into fence lines and open fields. Indiana little short-tailed shrew, white- Table 7 (cont’d) Some prairie mammals adapted to the tailed jack rabbit, Franklin’s ground squir- Rare and declining initial loss of prairie vegetation and more rel, and prairie vole. Wisconsin grassland recent land-use changes and are thus still Grassland bird populations were plants. Compiled from Wisconsin Department doing well in Wisconsin today. These substantially altered by Euro-American of Natural Resources include the prairie settlement. But (1992). mole, thirteen-lined because grassland ground squirrel, Wisconsin’s grassland plants and animals birds are not strictly harvest mouse, and responded to the changes that came with dependent upon prairie deer mouse. Euro-American settlement in various native vegetation, Other species have ways. Some species adapted well and they are one group not adapted well to maintain healthy populations today, while that generally did not the changes, and decline solely because some are persisting only in low numbers. have been either of the loss of native extirpated as men- Others are restricted to prairie and sedge vegetation. They are, tioned above or are meadow remnants, and a few have been however, sensitive to now of special extirpated. An estimated 15%-20% of the both the structure of concern in the state. state’s original grassland flora is now vegetation (e.g., The special concern considered rare in the state. degree of treelessness, species include vegetation height and

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density, and amount of residual ground (area-sensitive species) such as sharp-tailed cover) and size of habitats, as well as to the grouse, greater prairie-chicken, and short- nature and timing of eared owl, are not agricultural distur- thriving today, in bances mentioned Little is known about the status of part because of the above. Radical invertebrates in our native grasslands. In fragmentation and changes in these fact, there are probably dozens of reduction of large habitat features have grassland insects in Wisconsin still habitat tracts. been occurring over unknown to science . . . . Many species Today, grass- the past 150 years; may have already been extirpated or land bird species often these changes vary in their status. become extinct without our having known have had direct or Of those that were indirect ramifications of their existence. historically present Table 8 for bird populations. in Wisconsin, a few For example, species are still doing very Rare and declining with large minimum area requirements well, often because they (1) are generalists Wisconsin grassland birds. that can use a variety of habitat types (e.g., red-winged blackbird, mourning dove, and song sparrow) or (2) have adapted to Status Common Name intensive row-crop agriculture (e.g., killdeer and horned lark). However, the status of most grassland birds is far less secure Extinct (none) than that of these few species. A variety of Extirpated whooping crane* species adapted well to the low intensity long-billed curlew agriculture that occurred before the late swallow-tailed kite 1950s, and they thrived until then. How- Threatened greater prairie chicken ever, in the past 30 years many of them Special Concern northern harrier (e.g., bobolink, eastern meadowlark, field sharp-tailed grouse sparrow, and grasshopper sparrow) have upland sandpiper begun to decline, due in part to the short-eared owl changes in land use and agricultural dickcissel practices mentioned above. Other species, Henslow’s sparrow such as greater prairie-chicken and sharp- grasshopper sparrow** tailed grouse, did well after Euro-American Le Conte’s sparrow settlement, but they did so partly by sharp-tailed sparrow expanding their ranges into the vast logged lark sparrow and burned-over lands of northern Wiscon- bobolink** western meadowlark** sin. As the northern habitat grew back to yellow rail forest, these species eventually declined as Wilson’s phalarope well. sedge wren As a result of a combination of factors northern pintail including habitat changes over the past 150 Declining savannah sparrow** years on breeding grounds, wintering eastern meadowlark** grounds, or both, and habitat-related vesper sparrow** problems with nest productivity, 16 of field sparrow** Wisconsin’s grassland bird species are now blue-winged teal** of special concern in the state (Table 8); one (greater prairie-chicken) is on the * Federally endangered. state’s list of threatened species. In addition, ** Declining in recent years based on federal breeding bird surveys conducted in Wisconsin. three other grassland birds, whooping crane, long-billed curlew, and swallow- tailed kite, have been extirpated from the

120 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE state due in part to their inability to adapt brates and vegetation (e.g., host plant to land-use changes and unregulated specificity at the species, genus, and family hunting. All three extirpated species now levels), many grassland invertebrates would have reduced ranges, but none is extinct. be considered rare and endangered at both One, the whooping crane, is federally state and federal levels if distribution and endangered. population data were available. For ex- Only about one-half of Wisconsin’s ample, some information is available about prairie-associated reptiles and amphibians Lepidoptera (moths and butterflies); are still at good population levels today. consequently, 19 grassland Lepidoptera are These include eastern tiger salamander, six- now on the state’s list of special concern; lined racerunner, blue racer, eastern plains two species, swamp metalmark and regal garter snake, and Butler’s garter snake. Like fritillary, are on the Wisconsin threatened many other vertebrates, their success has species list (regal fritillary is also being been due to their ability to adapt to surro- considered for federal listing); and three gate “grasslands.” The rest of the prairie species, Powesheik skipper, phlox moth, reptiles have not adapted as well and are and silphium borer moth, are on the apparently suffering from habitat loss and Wisconsin endangered species list. fragmentation. Of this group, three (ornate box turtle, western slender glass lizard, and PROJECTED massasauga rattlesnake) are on the state list of endangered species, one (Blanding’s What remains of our native grassland turtle) is on the state list of threatened systems has neither long-term nor short- species, and two term security. In the (prairie ringneck absence of additional snake and bull If recognition, protection, management, recognition and management, grass- snake) are on a list of and restoration are actively pursued and land species and special concern in fostered at levels greater than they have the state. remnant vegetation been in the past, most of the biotic Little is known will continue to be about the status of diversity of our original grassland lost due to area invertebrates in our ecosystems can be retained within the reduction, fragmenta- native grasslands. In state over time. But time is running out tion, isolation, and degradation (ecosys- fact, there are prob- fast. With each passing year, options for ably dozens of tem simplification). retention or recovery are lost at an However, if recogni- grassland insects in accelerating rate, and the costs and Wisconsin still tion, protection, efforts needed to retain grassland unknown to science. management, and For example, a biodiversity increase. restoration are cursory search for actively pursued and leafhoppers at 14 fostered at levels Wisconsin prairie remnants in 1993 and greater than they 1994 revealed five leafhopper species new have been in the past, most of the biotic to science and 24 species never before diversity of our original grassland ecosys- recorded from the state (K.G.A. Hamilton, tems can be retained within the state over Agriculture Canada, Ottawa, pers. comm.). time. But time is running out fast. With Many species may have already been each passing year, options for retention or extirpated or become extinct without our recovery are lost at an accelerating rate, and having known of their existence. In light of the costs and efforts needed to retain this ignorance and the fact that there are grassland biodiversity increase. often close relationships between inverte-

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✓ tree planting for “wildlife,” “aesthet- ACTIONS CAUSING CONCERN ics,” and timber/ﬁber production. (Planting trees into prairie remnants Threats to the future survival of our is a common practice, so as to make native grassland ﬂora, fauna, and vegetation economic use of sites people perceive remnants can be summarized in six catego- as being “worthless.”) ries: Continued loss of native remnants (both ✓ public opposition to tree removal high-quality sites and those moderately needed to restore or maintain grass- degraded by grazing) due to: lands.

[top] Spring Green dry ✓ accelerating invasion by woody ✓ rural home building; this is often prairie seen in 1975 growth on both wet and dry sites focused on nonagricultural lands with the beginnings of red cedar invasion. (e.g., red cedar is now invading dry and, thus, prairie remnants. Photo by Bill Tans. bluff prairie so fast that in 20 years ✓ [bottom] The same most unmanaged bluff prairie re- conversion of traditional prairie view of Spring Green maining in the Midwest will be pastures (unplowed but grazed dry prairie in 1991 with completely overgrown, and wetland prairie) to crops. red cedar nearly shrubs and trees are increasingly covering the hillside. ✓ Photo by Richard taking over extensive areas of sedge drainage and conversion of sedge Henderson. meadow). meadow and wet prairie to muck farming.

Continued loss of surrogate post- settlement “grasslands” used by grassland animals (especially birds), due to intensive agriculture and urban development.

General lack of attention to native grassland communities by the public, resource managers, and scientists.

Resistance to the use of prescribed fire and lack of understanding by the public and professionals of fire effects (i.e., the consequences of both too much and too little fire).

Invasion by aggressive exotics (e.g., honeysuckle, common buckthorn, reed canary grass, leafy spurge, parsnip, purple loosestrife, etc.).

Habitat fragmentation, which results in patch isolation and the creation of edge effects. This is especially harmful to vertebrate animals.

122 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE SOCIO-ECONOMIC ISSUES POTENTIAL FOR COMMUNITY RESTORATION Strong public support will play a vital role in retaining and regaining grassland Recovering and maintaining native biodiversity in Wisconsin. Management of grassland biodiversity in Wisconsin is very the grassland ecosystem, or at least ele- feasible for many but not all components ments of it such as open treeless habitat for (e.g., birds, plants, and invertebrates) of the grassland birds, is compatible with many system. It is unlikely that we will ever again traditional wildlife management and hunter be able to accommodate mega-fauna such interests (i.e., species such as ring-necked as bison, elk, and pheasant, upland wolves in a naturally nesting ducks, sharp- functioning grassland tailed grouse, and Strong public support will play a vital role ecosystem in Wiscon- prairie chicken). in retaining and regaining grassland sin. Livestock grazing and biodiversity in Wisconsin. Retention of crop production are grassland biodiversity also potentially will require more compatible with maintaining the open than just the preservation of existing high- habitat structure needed by many grassland quality remnants of native vegetation. Most animal species. Agricultural and soil remnants are less than ten acres in size and conservation programs, such as the Conser- very few exceed 50 acres. They are just too vation Reserve Program (CRP), and water small for many if not most vertebrate quality protection can also be very compat- animal species. Small sites, however, are Restoration will not ible with grassland habitat interests. capable of supporting viable populations of be easy, and it will There is already some public aware- most plant species, most soil microflora take much time, ness and interest in both prairie flora (as and microfauna, and many other inverte- maybe even exemplified by the popularity of prairie brate species for decades if not centuries to centuries, before restoration and landscaping) and grassland come, especially if the sites contain soil the prairie birds that are well known and popular with moisture gradients and are provided with bird watchers, such as sandhill crane, buffer land. community is prairie chicken, meadowlark, bobolink, Remnants degraded by grazing or significantly Henslow’s sparrow, and upland sandpiper. woody growth invasion can also play a restored. However, These combined interests should translate significant role. Degraded areas are much restoration is into public support for grassland habitat more common and often larger than high- feasible. Many preservation and restoration. There is quality remnants. Their value is in the elements of the already much support among private residual species they still harbor and the conservation groups for prairie and sedge great potential they have for recovery. Their system can still be meadow protection and recovery; for condition is often such that recovery can be found in forgotten example, mesic prairie is a top preservation accomplished solely by brush removal, corners of the priority for The Nature Conservancy; there restrained grazing, or fire. The greatest landscape, and is a new and growing regional conservation opportunities for recovery of degraded sites they can be group called The Prairie Enthusiasts; and are at the dry and wet ends of the soil brought back The Madison Audubon Society is support- moisture spectrum, where several thousand together with ing a large prairie restoration on land they acres of degraded dry prairie and sedge hold. meadow still exist. reasonable effort. Recovery of the mesic prairie system is a different situation. Because mesic prairie remnants of any quality are very rare, retaining or regaining components of this system will require extensive buffering of the few remaining remnants and much

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POSSIBLE ACTIONS

The following possible actions are consistent with ecosystem management, but require more analysis and discussion. How priorities are set within this list will be based on ecoregion goals, staff workload, fiscal resources, public input and support, and legal authority. We will work with our customers and clients to set priorities and bring recommendations to the Natural Resources Board for consideration beginning in the 1995-97 biennium. Efforts at recovery and maintenance of grassland biodiversity in Wisconsin should focus on three general areas of concern: (1) Mesic prairie remnant restoration from scratch (i.e., on sites bird, herptile, and small mammal commu- between hay and corn highly altered by tillage or other intense fields. Most mesic nities that require large habitat areas but disturbance). Such restoration will not be prairie has been not necessarily native vegetation; (2) native converted to agricul- easy, and it will take much time, maybe community remnants (vegetation, soil, and ture. The soil on which even centuries, before the prairie commu- mesic prairies occurred invertebrates); (3) endangered or threat- nity is significantly restored. However, is deep and fertile. ened animal species that have requirements Ipswich Prairie, Grant restoration is feasible. Many elements of the for both native vegetation and large areas County. Photo by Eric system can still be found in forgotten Epstein. (e.g., ornate box turtle). Species in the corners of the landscape and they can be latter category will need specific recovery brought back together (i.e., translocation of plans, which are not addressed here. individuals or reintroduction by seed) with Possible actions that address the first two reasonable effort. As reduced and frag- concerns are as follows: mented as the prairie ecosystem is in Wisconsin, local genetic variations of 1. Establish treeless grassland habitat at species, particularly plants and inverte- several landscape scales to meet area brates, still survive in low numbers on the requirements for species ranging from landscape. This genetic diversity can still be the prairie chicken to the grasshopper accessed for restoration. Although restora- sparrow. Examples of both lowland and tions should be viewed as long-term, they upland habitats should be sought, and can, in as little as a decade, result in most projects should be in former native reasonable facsimiles of prairie that support grassland areas. Rationale for the latter far more biotic diversity than alternative requirement is that historical grassland grass covers such as brome or switchgrass. areas will have the soil, topography, Given the fragmented nature and remnant vegetation, lack of large trees, small size of native remnants and even and climatic conditions most conducive potential restorations, the main hope for to restoring and maintaining open grassland vertebrates lies with surrogate grassland habitat. However, some “grassland” habitat that does not necessarily regions of existing cleared forest or have native vegetation. The opportunities drained marsh may prove suitable as for establishing this habitat are extensive on well. both private and public lands, especially DNR-managed lands. In many cases The total acreage of permanent grass/ establishment would only require removal forb cover needed for maintaining viable and control of woody growth. In others it populations of grassland birds in the would require the establishment of perma- state is unknown. At least 3%-4% nent grass/forb cover. (90,000-125,000 acres) of the original

124 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE acreage may be required. However, even status and management is presently more important than total acreage is the being written. placement and configuration of these acres. The following three-part strategy 2. Manage, enhance, and restore native is recommended: vegetation remnants as refugia for flora, invertebrates, and ecological processes. Large Landscapes. Establish 4-5 For the most part these efforts will be at landscape regions of at least 10,000 scales far smaller than those used for the acres each that are as treeless and “grassland” habitat discussed above. open in character as possible. Within However, some acreage overlap of the each region there should be a core two programs is likely to occur and area of permanent grass cover that is should be encouraged. Recommended at least 2,000 acres in size. Within strategies for maintaining and recovering the rest of the region there should be the two major grassland types of the at least 35% permanent cover, 75% state—sedge meadow and prairie—are of which should be in units of 40 as follows: acres or more. The remainder (52% of total) can stay in crop production. Sedge Meadow. The total acreage Small grain and hay crops should be needed to ensure the long-term encouraged. survival of the sedge meadow plant and invertebrate communities in the Small Landscapes. Establish 10-12 state is unknown. One to two percent landscape regions of 1,000-5,000 (10,000-20,000 acres) of the original acres each, with permanent cover one million acres may be a reasonable cores of at least 250 acres and a 15% target. permanent cover over the rest of the region. Follow the same guidelines Highest priority should be given to used in the large landscape regions. the protection, maintenance, and recovery of the largest and most Scattered Tracts. Establish numer- intact examples. However, small, ous scattered grass/forb fields that are high-quality sites (as small as ten at least 20 acres in size when there acres) should not be overlooked; are no edge effects from trees or other such areas may represent the last obstructions. If edged by trees, the refuge for many sedge meadow plant, minimum acreage should be 40 insect, and soil microflora and acres. Total acreage goals should be microfauna species. Special priority approximately 50,000 acres. When should also be given to sedge mead- possible these fields should be placed ows that are part of larger grasslands in close proximity to other perma- or wetland complexes. nent grass/forb cover. Once the high-quality sites are secure In the development of these “grass- (including adequate buffer lands), land” habitat areas, a variety of grass/ degraded areas with high recovery forb cover types should be used. potential should be considered for Incorporation of native remnants and completing the acreage goals. Resto- restoration of native vegetation ration of sedge meadow from scratch should be encouraged, but not made is not a desirable recovery strategy at an absolute requirement. For more this time, because of the adequate detailed habitat recommendations, amount of remnant acreage that still see Sample and Mossman (1990). In exists and the great difficulties addition, a Bureau of Research associated with sedge meadow Technical Bulletin on grassland bird restoration.

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These repositories must not be lost. Their genetic holdings will be needed in any future prairie restorations. Buffer lands will be crucial to the long-term (100 years or more) survival of all prairie remnants— especially the smallest ones—and their dependent species. Buffer lands are needed to protect remnants against the negative impacts of external influences and stochastic events, and to provide living space into which the prairie community can spread and rebuild marginal populations. Ideally, buffer lands should also provide those portions of Curtis Prairie, UW Prairie. As with sedge meadow, the the soil-moisture spectrum not found Arboretum, an total acreage needed for long-term example of dry mesic in the remnant. In most cases this survival of the prairie plant and prairie. Restorations will be mesic soil. It would also be such as this will be invertebrate communities in the state ideal to buffer remnants by including needed to meet the is unknown. Again, 1%-2% (21,000- possible acreage goals them in larger grassland bird habitat 42,000 acres) of the original 2.1 for native grasslands areas. restoration. Photo by million acres may be a reasonable Richard Henderson. target. In the case of prairie, however, 3. If we are serious about long-term this goal is at least ten times greater retention of grassland biodiversity in than the total known acreage of all Wisconsin, eventually we will need three high- to moderate-quality prairie or four large-scale restorations (greater remnants combined, and it probably than 1,000 acres) that encompass exceeds the combined acreage of all clusters of existing remnants. Such remnants, including degraded ones acreage is needed for natural landscape (i.e., remnants not included in the processes to occur. Having such areas inventory). Therefore, some restora- will also eventually reduce management tion from scratch will be needed to costs per acre; for the effort required to Buffer lands will be meet the acreage goal, preferably on maintain a remnant community is buffer lands surrounding remnants. inversely proportional to the size of the crucial to the long- remnant. term (100 years or Highest priority should be put on the more) survival of protection and maintenance of all 4. The current DNR/DOT Native Plant high-quality remnants of an acre or all prairie Seed Program, which will be supplying more in size, followed by degraded local genotypes, must be encouraged remnants, remnants of five acres or more. High- and expanded if restoration on a large especially the quality sites as small as 1-2 acres scale is to become feasible. smallest ones, and should not be ignored, especially 5. Whatever the final acreage goal for their dependent when they contain mesic prairie, for either surrogate “grassland” habitat or species. they are probably the last refugia for many prairie plant, insect, and soil native remnant vegetation, it should micro-organism species. In addition, include representation of a variety of soil because of the near total loss of and topographic types, as well as prairie, these small remnants now geographic locations. Based on historical collectively function as the repository occurrence, the total acreage goal should for the genetic diversity of most be shared among DNR Districts in the prairie plants and invertebrates. following approximate proportions:

126 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Southern District...... 45% Western District ...... 32% Northwest District...... 12% North Central District ...... 5% Southeast District ...... 5% Lake Michigan District ...... 1% To reach these goals, some acquisition of private land will be needed, especially for remnant sites. However, much of Wisconsin’s native grassland biodiversity can be maintained and regained without new land acquisition. Many opportunities exist for maintenance and recovery on land already managed by the DNR, especially in the programs of the Bureau of Parks and Recreation and the Bureau Department’s Division of Environmental of Wildlife Management. Much could A policy for prescribed Quality will need to be clarified. burning that also be accomplished outside of DNR recoginzes the lands through cooperation and partner- 8. Qualitative inventories of selected dependance of some ships with other agencies (e.g., roadside invertebrate taxa in remnants of prairie ecosystems, including grasslands, on fire programs) and private conservation and sedge meadow and other grassland need to be developed. groups. There are also many opportuni- types (including non-native surrogates) Photo by Richard ties for encouraging surrogate habitat are needed for the purpose of determin- Henderson. and remnant management on private ing what specialized, remnant-restricted lands through tax incentives (e.g., the species still exist, their distribution, and Minnesota Prairie Bank Program), their status. This information, which is educational programs, agricultural currently lacking for the most part, programs (i.e., agricultural policy, farm would be of great assistance in setting bills, continuation of CRP and annual protection and management priorities. set-aside), technical advice and assis- 9. Much additional research is needed on tance, and the Habitat Restoration Areas the effects of grassland management component of the Wisconsin Steward- techniques, such as burning, mowing, ship Program. and grazing, on grassland vegetation and 6. Because of the current rarity and long- fauna. Obtaining this information will time absence of prairie on the landscape, be crucial to our long-term ability to a program of education/awareness is manage grasslands for the entire array of greatly needed for developing support native grassland biodiversity. for prairie recovery and maintenance. 10.Planning and coordination among all The Department’s Bureau of Parks and land management interests, especially Recreation and the Bureau of Informa- within the DNR, is crucial, so that tion and Education should play major programs do not inadvertently cancel roles in this. each others’ efforts. For example, tree 7. Develop a policy on prescribed burning planting programs should strive to avoid that recognizes the dependence of some destruction of prairie remnants or ecosystems, including grasslands, on fire fragmentation of grassland habitat and, and examines the resources and staff conversely, grassland habitat projects support necessary to effectively and should avoid areas that are more appro- safely use fire to manage these fire- priate for reforestation. Integrated dependent communities. In addition, air management is the key to overcoming quality standards and policies within the these types of management and policy conflicts.

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Case Study

THE GLACIAL HABITAT RESTORATION AREA: AN APPROACH FOR RESTORATION OF GRASSLAND COMMUNITIES

Contributed by Becky Isenring and Richard Henderson.

Native grasslands are among the most threatened natural community types in the state. Thus the Department faces a tremendous challenge in addressing the needs for restoration of these communities and preserving the native species in them. One approach that takes a step towards this end is the Glacial Habitat Restoration Area (HRA). The primary objective of the Glacial HRA is to provide nesting habitat on a landscape scale for upland nesting waterfowl, native grassland non-game birds, and pheasants. This is to be achieved by restoring 10% of the land within a selected region of the state to a suitable condition. Restoration will be accomplished through a program of acquiring land rights through fee title and perpetual easements and then restoring the land to grassland and wetland habitat. Innovations of the Glacial HRA project are its size and scope. The area covers 530,000 acres within the glaciated, former prairie/savanna area of southeast Wisconsin. Restoration goals are 38,000 acres of upland grassland habitat and 11,000 acres of wetland habitat distributed in small, scattered units (10-250 acres) within high priority sub-units of the entire HRA project area. The idea is to take an area of the state of manageable size and, applying research-based species management guidelines, reintroduce complexity into a simplified landscape. The priority sub-units were identified using Geographic Information System (GIS) analysis of three landscape habitat models developed for prairie waterfowl, native upland grassland birds, and pheasants. Any area that met the minimum criteria for at least two of the three habitat models became high priority for the programs restoration activities. The Glacial HRA focuses on distributing most of the restoration into small scattered units (smaller than 100 acres). The largest unit goal is 250 acres, of which 12 are being sought within the total HRA. Restoration activities include planting native prairie species mix of 6-20 species. The Glacial HRA program will do an excellent job of providing the scattered tract component of habitat for upland and wetland grassland birds. It will also meet some needs for native prairie restoration and protection of native sedge meadows. In the process of implementing the Glacial HRA, Department managers are realizing that it can be used as a springboard into a program that does even more for grassland biodiversity needs. Some ideas being considered in statewide discussions include (1) making some management units larger (250-2,000 acres) because the program could go farther to meet the habitat needs of grassland birds, mammals, and reptiles if the restoration units were larger; (2) using seed from local sources and planting it in mixes of 80 plus species to increase benefits to include a broader range of wildlife species; (3) making native prairie remnants acquisition priorities so that the Glacial HRA would help to meet the total protection and restoration needs of native prairie vegetation, its associated soil communities, and prairie-restricted macro-invertebrates; and finally (4) having Department managers discuss how they can take what they have learned from the Glacial HRA and apply it to other areas of the state that have native prairie and sedge meadow restoration opportunities.

128 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE LITERATURE CITED

Curtis, J. T. 1959. The vegetation of Wisconsin Department of Natural Wisconsin. Univ. Wis. Press, Madison. Resources. 1992. Wisconsin Natural pp. 657. Heritage Inventory working list, Bureau Sample, D. and M. Mossman. 1990. Habitat of Endangered Resources. management guidelines for grassland birds on public and private land in southern Wisconsin. Unpubl. memo to public and private land managers on ﬁle at Wis. Dep. Nat. Resour. Res. Cent., Monona, Wis.

ADDITIONAL READING

Collins, S. and L. Wallace (eds.). 1990. Fire Risser, P., E. Birney, H. Blocker, S. May, W. in North American tallgrass prairies. Parton, and J. Wiens. 1981. The true University of Oklahoma Press, Norman. prairie ecosystem. US/IBP Synthesis 271 pp. Series No. 16. Hutchinson Ross, Hoffman, R.M. and D. Sample. 1988. Birds Stroudsburg, PA. 557pp. of wet-mesic and wet prairies in Sample, D.W. and R.M. Hoffman. 1989. Wisconsin. Passenger Pigeon Birds of dry-mesic and dry prairies in 50(2):143-152. Wisconsin. Passenger Pigeon Reichman, O. 1987. Konza Prairie: a 51(2):195-208. tallgrass natural history. University Wali, M. (ed.). 1975. Prairie: a multiple Press of Kansas, Lawrence. 226pp. view. University of North Dakota Press, Grand Forks. 433pp.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 129 WETLAND COMMUNITIES

area where water is at, near, or above the land surface long enough to be capable of supporting aquatic or hydrophytic vegetation and which has soils indicative of wet conditions.” People often think of wetlands as CHAPTER 9 cattail marshes utilized by waterfowl and muskrats. However, many other types of wetlands occur in Wisconsin and are given names such as wet meadow, swamp, bog, fen, sedge meadow, shrub-carr, alder Wetland thicket, conifer swamp, and bottomland or lowland hardwood forest. Curtis (1959) described wetland communities for Wis- consin and discussed their general charac- Communities teristics and relative diversity of plant species (Table 9). Wetlands vary in their by Steven W. Miller plant and animal composition and in their Division of Resource Management diversity. Northern bogs, for example, are Department of Natural Resources generally acidic and support fewer plant and animal species in fewer numbers than the alkaline marshes of southern Wiscon- sin. Wetland communities Detailed wetland classification sys- vary widely in their tems have been developed. The earliest, plant and animal most widely used major classification composition. For system for wetlands in the United States example, northern bogs, such as this was developed by Shaw and Fredine muskeg and bog along DESCRIPTION (1956). However, this system was overly the Black River in simplistic and was replaced with the U.S. Douglas County, ll wetlands have a common are generally acidic Fish and Wildlife Service’s Classification of and support characteristic—soils or a Wetlands and Deep-Water Habitats of the species adapted to substrate that is periodically United States (Cowardin et. al. 1979). very different saturated with or covered by conditions than the Wisconsin’s classification system (Wis. Dep. alkaline marshes of water. The statutory defini- Nat. Resour. 1992c) is based on this southern Wiscon- tion of a wetland used in system, but incorporates some modifica- sin. Photo by Eric Section 23.32 (1), Wisconsin Statutes is “an tions to make it easier to use and under- Epstein. stand. (See Payne 1992 for comparison of wetlands classification systems.) Wetlands are part of the water cycle of all ecosystems, and their location in the landscape allows them to function as a buffer between upland areas and surface waters (Weller 1981). Wetlands perform a number of natural functions that benefit natural ecosystems and society. Water quality is often dependent upon wetlands because they serve to trap sediment, remove nutrients, protect shorelines, and slow the effects of flood water. They also serve as both discharge and recharge areas for groundwater and provide habitat for

130 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Table 9. Wetland Communities of Wisconsin (from Curtis 1959)

Community Description Approximate Original Area

Southern Lowland Found along river valleys and on lake plains primarily south of theTension Zone; also 420,000 in the two types. Forests in depressions on poorly drained moraine; known as bottomland or floodplain forests along rivers and hardwood swamps on lake plains; floodplain forests present along all of the major rivers in southern Wisconsin; hardwood swamps found around the larger existing lakes and also on extinct glacial lakes. American elm was formerly important in all southern lowland forest types. Southern Dominated by boxelder, black willow, cottonwood, silver maple and river birch. Very small, probably only 20% of Wet Forests total bottomland forest or 84,000 acres. Southern Dominated by silver maple, green ash, swamp white oak, and hackberry. Uncertain. Probably 80% of total Wet-Mesic Forests bottomland or 336,000 acres. Northern Include tamarack-black spruce bog forests, white cedar-balsam fir conifer swamps, 2,240,000 in the two types. Lowland Forests and the black ash-yellow birch-hemlock swamps; found on lake beds and along streams north of the Tension Zone. Northern Dominated by black spruce and tamarack; white cedar, balsam fir, and jack pine of Uncertain. Possibly 75% of total Wet Forest secondary importance; with an understory of mosses, sedges, and ericaceous northern lowland forest or shrubs; occurs on acid peat. 1,680,000 acres. Northern Cedar swamps are dominated by white cedar and balsam fir; with hemlock, yellow Uncertain. Possibly 25% of total Wet-Mesic birch, and black ash of secondary importance. Hardwood swamps are dominated by northern lowland forest or 560,000 Forest black ash with yellow birch birch, red maple, and white cedar. acres. Open Bog Has a continuous carpet of sphagnum moss; found in pitted outwash or kettle No information. Probably less than depressions, mostly in northern Wisconsin with a few relicts in southern Wisconsin; 5% of conifer swamps or 110,000 dominant families are the Ericaceae and Cyperaceae; bog shrubs include rosemary, acres. leatherleaf, bog laurel, and Labrador tea. Alder Thicket Common along springy areas with mineral or muck soils, along streams, and around Unknown. lakes north of the Tension Zone; dominated by tag alder. Shrub-Carr Common around lakes and ponds and invades sedge meadow south of the Tension Unknown. Zone; wet-ground community dominated by tall shrubs other than tag alder; dominated by red osier dogwood and willow species. Sedge Meadow Open community of wet soils where more than half the dominance is contributed by 1,115,000 acres in the two types. sedges rather than grasses; found in all regions of the state in extinct lake beds, around the shores and banks of lakes and streams, and in depressions in pitted outwash or moraine topography. Northern Tussock meadows (dominated by Carex stricta) occur statewide and are generally Uncertain. Probably 105 or smaller in the north. Wire-leaved sedge meadows are found mostly in northern 115,00 acres. Wisconsin and can cover thousands of acres. Southern Dominated by Carex stricta and bluejoint grass; occur along streams and lakeshores Uncertain. Possibly 90% or and in morainal lowlands. 1,000,000 acres. Calcareous Fen Shrub-herb community on a wet and springy site with an internal flow of alkaline Very small, probably only a few water; found more frequently in southeastern counties. hundred acres. Wet Prairie Grassland on wet soils; located south of the Tension Zone; dominated by bluejoint Uncertain. Possibly 5% of total grass, sloughgrass, big bluestem, and prairie muhly grass. prairie or 105,000 acres. Wet-Mesic Prairie Grassland on seasonally wet soils; located south of the Tension Zone; dominated by Uncertain. Possibly 20% of big bluestem, bluejoint grass, sloughgrass, and wild rye. total prairie or 420,000 acres. Emergent Aquatic Group of wetland communities along the dividing line between true aquatic and true Unknown. Communities terrestrial communities. Includes deep and shallow marshes. Found along streams and streamside marshes throughout Wisconsin and along lakes in glaciated Wisconsin.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 131 WETLAND COMMUNITIES

STATUS

PAST Wisconsin’s topography was shaped largely by glacial activity. As a result, wetland communities were abundant in Wisconsin before Euro-American settlement and occupied an estimated ten million of the state’s 35 million acres (see Table 9). This estimate is based on the original government land surveys of the early 1800s and modern soil surveys; it Wetlands are part of many species of plants and animals (Stearns the water cycle, and may be a low estimate, since the data used their location in the 1978). by surveyors in the 1800s were based on a landscape allows them Wetlands are interrelated and inter- wetland definition that is conservative to function as a buffer spersed among all the other community compared to our current definitions and between upland areas types described in this report. Many and surface waters. because few soils have been mapped in This ecosystem, which wetlands are forested (e.g., wet forests and northwestern Wisconsin (Wis. Dep. Nat. includes pond, ridge, wet mesic forests) and must be considered Resour. 1990). fen, open bog, and as part of the continuum of northern or upland along Lake In the Driftless Area of the state, Superior, serves as a southern forest ecosystems. Wetlands are which was not affected by the most recent natural buffer that traps also interspersed among the prairie and oak glaciation, forested and nonforested wet- sediments, removes savanna areas of southern and east-central nutrients, and protects lands existed primarily along streams and the shoreline. Photo by Wisconsin. The rivers or as spring Cliff Germain. spatial connections seeps. In other between wetlands, Unique to wetland communities and regions of the state, lakes, rivers, and aquatic communities are the state and wetlands occurred on streams are obvious vast areas of peat soils federal laws that govern their use. These to anyone who has occupying former spent time in wetland are the only community types in glacial lake beds, as communities. Wisconsin for which a body of law potholes and fens; Unique to regulating use has developed. along streams and wetland communities rivers; on the borders and aquatic commu- of lakes; as bogs, nities are the state forested swamps, and bottomlands; and as and federal laws that govern their use. estuaries and coastal wetlands along Lake These are the only community types in Michigan and Lake Superior. Wisconsin for which a body of law regulat- Wetlands have been subjected to ing use has developed. As discussed later, intense modification and use and have these laws developed over many decades as greatly decreased in number since Euro- these communities suffered continued American settlement. Nearly all remaining destruction. The direct positive effects wetlands have suffered from the effects of wetlands exert on water and water quality simplification and fragmentation. From the served as the driving force behind the beginning of Euro-American settlement in development of these regulations. In the early 1800s until relatively recently, contrast, no regulations to protect terres- wetlands were viewed as wastelands and trial communities from permanent loss and were given economic value only when alteration have been developed. drained or filled (McCormick 1978). The 1850 Federal Swamp Land Act officially set national policy as one of wholesale wetland

132 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE “reclamation.” In Wisconsin, wetland loss was also accelerated by the 1925 Wisconsin Drainage Law (Kabat 1972). Historically, the greatest threat to wetlands in Wisconsin has been from agricultural drainage and urban development. Nationally, more than 87% of wetland losses have been due to agricultural development (Tiner 1984). Since wetlands often occur along rivers and lakes, these sites have also been considered to be of particular value for port facilities, for industrial development that required access to water for transport, for industrial pro- and continued wetland clearing and Wetlands are cessing or cooling water, for the discharge draining, subsidized by federal programs interrelated and interspersed among all of wastes, for marinas, for residential and tax incentives. However, beginning in other community developments with access to water, and for the late 1940s, with continued momentum types. Many wetlands deposition of dredge materials during into the 1970s, wetlands achieved ever- are forested and are construction of channels and wharf facili- part of the continuum wider recognition as valuable natural of northern and ties (McCormick 1978). resources. Increasingly, land-use plans southern forest Many thousands of acres of wetlands recommended various levels of wetland ecosystems, as shown were eliminated by shoreline development here by the concentric preservation; acquisition of wetlands for bands of open bog, for homes, resorts, and commercial and both state and national waterfowl manage- forested bog, and industrial development. As many of ment steadily increased; and new research forested upland in Wisconsin’s larger began to show the Washburn County. cities expanded, Wetlands are also Wetland communities were abundant in relationship between interspersed among forested wetlands and wetlands, water the prairie and oak Wisconsin before Euro-American the marshy estuaries quality, economically savanna areas of the of rivers were cleared settlement and occupied an estimated ten state. Photo by Robin important fish and Moran. and filled to accom- million of the state’s 35 million acres . . . . wildlife species, and modate development. Wetlands have been subjected to intense the preservation of For example, most of modification and use and have greatly rare plant and animal the industrial areas in decreased in number since . . . species (Kabat 1972, Milwaukee, Superior, settlement. McCormick 1978, and Green Bay are Visser 1982). The former view built on fill deposited Gradually, as that wetlands were in coastal wetlands. Portage, La Crosse, and the ecological values of wetlands were Prairie du Chien are Wisconsin cities built recognized, changes began to occur in just wastelands partly on riverine wetlands (Visser 1982). federal and state policies towards wetlands. and impediments Historically, some appreciation for The former view that wetlands were just to progress was unaltered wetlands began to appear in the wastelands and impediments to progress replaced by a 1930s, although drainage and filling was replaced by a recognition that wetlands recognition that continued to be promoted by federal and are critical components of healthy function- wetlands are state policies. During that period, concern ing ecosystems with significant direct and for wetlands was prompted in part by the indirect economic benefits. critical catastrophic decline in North American The oldest of federal laws used to components of waterfowl populations during the droughts protect wetlands is the River and Harbor healthy functioning of the 1930s (Kabat 1972). Nevertheless, Act of 1899, which prohibited the excava- ecosystems with an era of intensified agriculture began tion or deposition of material into any significant direct following World War II, which included navigable water of the United States with- and indirect heavy applications of fertilizers and pesti- out a permit from the U.S. Army Corp of economic benefits. cides, increasingly mechanized agriculture, Engineers. Although this act had the

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 133 WETLAND COMMUNITIES

potential for major wetland protection, it gated in NR 115 Wisconsin Administrative was administered in a manner that greatly Code. (Further references to the Wisconsin limited its effectiveness (McCormick 1978). Administrative Code will be shortened to The National Environmental Policy Act of “NR.”) In 1979, the Wisconsin Legislature 1969 required that many wetland drainage approved a statewide wetland inventory and filling projects be reviewed for their program, but the program had no concur- impact on the human environment, par- rent protection authority. In 1980, the ticularly if federal agencies or federal Natural Resources Board adopted NR 1.95, money was involved. The Federal Water which required Department personnel to Pollution Control Act of 1972 required consider the effect on wetlands when permits for the disposal of dredge material granting permits and to minimize wetland and the filling of waters of the state, damage in the permitting process. Also in including wetlands. The Federal Endan- 1979, NR 115 was amended to require gered Species Act of 1973 called attention counties to protect wetlands within 1,000 to the need to protect and restore wetland feet of lakes and within 300 feet of streams. habitats for endangered plant and animal An analogous rule, NR 117, was approved species. to protect wetlands occurring within cities The 1985 and 1990 Federal Farm and villages. NR 115 and 117 prohibit Bills contained milestone provisions for wetland alteration without first obtaining a protecting wetlands by imposing penalties rezoning approval from the county, city, or for converting wetlands to agricultural uses, village. The Department can veto a rezon- thus ending the federal agricultural subsidy ing approval if the local government fails to for wetland drainage. These restrictions, consider significant environmental factors which are known as the “Swampbuster” in granting the rezoning (Dawson 1982, provisions, helped to stem the rate of Visser 1982). wetland conversion to agricultural uses. The most recent wetland protection The 1990 Farm Bill also contained a law in Wisconsin is NR 103, which estab- wetland reserve program which allows the lishes state water quality standards for U.S. Department of Agriculture to take wetlands. These narrative standards are permanent wetland easements on restored applied to all Department activities that wetlands on private lands. affect wetlands. They are also applied to Paralleling wetland protection actions federal permits through the state’s water- on the federal level during this time period, quality certification process under the a number of legislative actions occurred in Clean Water Act. Wisconsin to strengthen state authority to During the era of the most intensive protect wetlands. Prior to the 1960s, the wetland loss and modification, wildlife and state’s role consisted of buying wildlife other natural resource values associated habitat, fish spawning grounds, and public with wetlands were recognized only in hunting areas. The state also had authority passing. As a result, along with the loss of under Chapter 30 of the Wisconsin Statutes wetland acreage, there was a concomitant Navigable Water Law to control filling on loss in the numbers of species dependent the beds of navigable waters (Visser 1982). on wetlands, including waterfowl, shore- Established in 1966, Section 144.025 birds, herptiles, fish, invertebrates, and of the Wisconsin Statutes required the many species of plants. There was also a Department to protect the waters of the loss of the ecosystem services performed by state, including wetlands. In the same year, wetlands, including floodwater storage, Section 59.971 of the Wisconsin Statutes sediment and contaminant filtering, and required counties to adopt shoreland groundwater discharge and recharge. zoning ordinances for unincorporated areas The species richness of many wetland within 1,000 feet of lakes and flowages and types prior to Euro-American settlement within 300 feet of navigable streams. Rules does not appear to be well-documented. for implementing s.59.971 were promul- There are, however, indications from

134 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE historical observers that many marshes attracted large numbers of migrating waterfowl, were important for ﬁsh spawning, and produced large amounts of useful products such as lumber, sphagnum moss, wild rice, and marsh hay (Curtis 1959).

PRESENT At present, Wisconsin has lost 47% of its original ten million acres of wetlands. Many of the remaining 5.3 million acres are in the northern third of the state (Wis. Dep. Nat. Resour. 1990). In some southern Wisconsin counties, the amount of wetland loss is well over 75%. Wisconsin’s losses are reﬂective of the national status of wetlands; ing wetland loss through an aggressive Many wetlands have it is estimated that one-half of the nation’s regulatory program involving local, federal, been lost to agricul- original 221 million acres of wetlands have tural drainage, urban and state governments. In addition, there development, and been lost (Feierabend 1992). A large are a number of incentive programs, many industrial develop- amount of remaining acreage in Wisconsin rather recent in origin, that are designed to ment. Channelization exists in a partly altered state, such as with of streams, like this restore or enhance wetlands. one in the central old drainage ditches still functional enough One newer program involves the U.S. sands region, was to change the hydrology of the wetland. Department of Agriculture Farmers Home used to drain land and Much of this remaining wetland acreage resulted in a simpliﬁed Administration (FmHA). As part of the and less diverse was at one time disturbed, either by 1985 Farm Bill, FmHA was given the stream system. Photo drainage (followed by authority to place by Michael J. restoration) or by restrictive-use wet- Mossman. being cleared, land easements on Wisconsin has lost 47% of its original ten repeatedly burned, properties they offer grazed, or periodi- million acres of wetlands. for sale after foreclo- cally plowed (Curtis sure. These wetland 1959). easements are then Although there is considerably less enforced by the U.S. Fish and Wildlife drainage of wetlands today due to the Service. FmHA can also allow borrowers to “Swampbuster” requirements of the 1985 reduce their debt by granting an easement Farm Bill, agriculture still affects wetlands to the Fish and Wildlife Service. Under the through grazing, barnyard and feedlot 1990 Farm Bill, the U.S. Department of runoff, pesticide and fertilizer runoff, and Agriculture was authorized to take perma- sedimentation from nonpoint sources. nent wetland easements. However, Con- Sedimentation of wetlands leads to the gress funded the program in 1992 only, so gradual loss of open-water areas and its effects have been very limited. Addi- development of monotypic stands of tional wetland protection opportunities vegetation that have less habitat value to occur in other federal laws such as the wildlife. Coastal Zone Management Act of 1972 and Currently, the collective use of Section the Fish and Wildlife Coordination Act. 404 of the Clean Water Act, the Overall, these state and federal “Swampbuster” provisions, NR 115, NR regulatory programs have the capability to 117, and NR 103 have controlled major substantially reduce wetland losses in wetland losses in Wisconsin (Dale Simon, Wisconsin. However, contained in these Wis. Dep. Nat. Resour., pers. comm.). The regulations are some exemptions for Department plays the lead role in prevent- agriculture, forestry, and various types of

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 135 WETLAND COMMUNITIES

commercial navigation activities. These wetlands that had been drained for agricul- exemptions have been criticized by ecolo- ture. Many of the drained wetlands were gists and wetland protection advocates as originally sedge meadows, shrub-carr, being unnecessary, and efforts continue to tamarack swamps, and wet prairie; how- bring all activities affecting wetlands under ever, shallow lakes were also drained. regulatory review. Restorations for waterfowl habitat often In addition to protecting wetlands resulted in shallow and deep-water marshes through regulations, the Department and that may not have been the condition of the the U.S. Fish and Wildlife Service have a wetland before it was drained. Some habitat commendable record of acquiring wetlands improvement projects also purposefully for wildlife and fishery management, converted sedge meadows, shrub-carr, and natural areas, and other purposes in the wet prairie into shallow and deep-water state. Between the U.S. Fish and Wildlife marshes under the justification that these Service, the Department, and nonprofit wetlands were being enhanced for water- conservation organizations, hundreds of fowl and wildlife. The result has been that thousands of acres of wildlife and plant wetlands have been species needing acquired, and many Many thousands of acres of small shallow and deep- thousands of acres of wetlands and associated uplands are in water marshes have drained wetlands greatly benefited, state and federal ownership. Much of this have been restored. while species that Notable large wet- acquisition was focused on waterfowl and preferred the pre- land acquisition and fishery management, but significant existing wetland type restoration projects benefits are provided to other wetland- suffered some habitat are Horicon Marsh dependent species such as sandhill reduction. From a National Wildlife cranes and other wading birds, furbearers, statewide perspective, Refuge, the Glacial however, the areas of herptiles, and plants. Lake Grantsburg wetland affected by Wildlife Area Com- wildlife management plex, Necedah activities are a small National Wildlife Refuge, Mead Wildlife portion of the total wetland modification or Area, Meadow Valley Wildlife Area, Green loss that occurred due to agriculture and Bay West Shores Wildlife Area, the Upper urban development. The net effect of Mississippi National Wildlife Refuge, and wildlife management wetland restoration the Mink River Estuary. Additionally, many and enhancement projects on biodiversity thousands of acres of small wetlands and appears to be positive. Current wildlife associated uplands have been purchased. management of wetlands focuses on Much of this acquisition was focused on restoring many of the original shallow and waterfowl and fishery management, but deep-water marshes that were drained for significant benefits are provided to other agriculture. wetland-dependent species such as sandhill On both national and state levels, cranes and other wading birds, furbearers, renewed emphasis was placed on the value herptiles, and plants. A number of private of wetlands for waterfowl and other aquatic organizations have also protected large wildlife with the implementation of the areas of wetlands. North American Waterfowl Management Wetland management practices Plan beginning in 1986 (U.S. Fish and conducted to improve waterfowl habitat Wildl. Serv. 1986). This program, signed by have impacted wetlands in the state. The Canada and the United States, encourages principles and techniques used and their public-private partnerships in protecting implications are discussed by Weller (1978, and restoring wetland habitats. The plan is 1981) and Payne (1992). In most cases, continental in scope, recognizing that many these activities have restored large areas of species of birds associated with wetlands

136 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE need secure habitats across the entire North American continent. Wisconsin is a direct participant in this effort through the Upper Mississippi River and Great Lakes Region Joint Venture (Wis. Dep. Nat. Resour. 1992a) The Wisconsin Wetland Inventory (Visser 1982, Wis. Dep. Nat. Resour. 1992c), authorized by the Legislature in 1979, was initially completed for all counties in 1984. Wetlands two acres or larger in size are delineated and classified on 1:24,000-scale maps. These inventory maps were supposed to be updated every ten years, but limited funding has slowed the process to a 20-year cycle (Dale Simon, Wis. Dep. Nat. Resour., pers. comm.). associated with wetlands, these species Wetlands provide habitat for many Wetlands are have suffered from the species. This forested noted for their loss of wetlands. bog in Douglas County supports abundance of plant 43% of all federally listed threatened and Currently, 43% and animal life. Of of all federally listed breeding pairs of at endangered species use wetlands at least sixteen species Wisconsin’s 370 threatened and of warblers. Photo species of birds, some point in their life cycle. 32% of the endangered species from Department of 39% live in or use state’s threatened and endangered plants use wetlands at some Natural Resources files. wetlands. According and animals are wetland-dependent. point in their life cycle to Hale (1982), no (Feierabend 1992); for other Wisconsin Wisconsin, 32% of the habitat type comes close to this avian state’s threatened and endangered plants occupancy rate. Hale also commented that although no wetland bird species has been The prairie white- fringed orchid extirpated from Wisconsin due to wetland (Platanthera destruction, the significant loss of wetland leucophaea), a showy acreage has to have caused a decline in state endangered and federally threatened wetland-dependent birds. species, occurs in Many important game birds, mam- wetlands in the mals, and fish are associated with wetlands. southern part of Waterfowl, beaver, muskrats, and northern Wisconsin. Photo by Thomas A. Meyer. pike are obvious examples. However, other species are also significantly related to wetlands. In some river systems, such as the Wolf River, walleye use seasonally flooded wetlands for spawning. Ring- necked pheasants use shrub-carr and cattail marshes during the winter. White-tailed deer thrive in wetland areas composed of shrubs and trees. Vogt (1981) identified southern lowland forests as “exceptionally rich” and aquatic communities (including open-water marshes) as “extremely rich” in herptiles, as compared with other community types in Wisconsin. Since many herptiles are

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 137 WETLAND COMMUNITIES

limited or no mobility. The preservation of wetland species that are essentially immo- bile is dependent upon land protection programs such as state, federal, and private acquisition or cooperative programs with private landowners. Currently, Wisconsin has an aggressive program of identifying and protecting lands with high natural-area value using the Natural Heritage Inventory. The highest ranking examples of all wetland types are considered priorities for permanent protection in the Department’s Natural Areas program.

PROJECTED

The hardwood swamp and animals are wetland dependent The enforcement of existing wetland- has recently been use regulations should prevent further identified as a discrete (Charles Pils, Wis. Dep. Nat. Resour., pers. community in comm.). Tables 10 and 11 show the major loss of wetlands in Wisconsin. It is Wisconsin. The ash wetland species currently on Wisconsin’s not possible for every remaining acre of swamp, including the endangered and threatened species lists. wetland to be preserved. In our society, Ashland County site some wetland loss will be unavoidable, but pictured here, is very Considering the vast acreage of wet and is dominated northern and southern wetlands that have rigorous planning and analysis of alterna- by black ash and alder been drained, cleared, intensively grazed, tives should help to minimize losses and with mineral rich avoid negative impacts from the perspective groundwater. Photo by repeatedly burned, plowed, flooded to Eric Epstein. create recreational lakes, or filled, it is of concern for biodiversity. A major threat possible to appreciate how local popula- would result if the federal government tions of species became disjunct from one changed its definition of wetlands, thus another and eventually extirpated because eliminating protection under Section 404 of they could not adapt to the changes in the Clean Water Act for millions of acres of Table 10 plant succession or were unable to with- wetlands (Wis. Dep. Nat. Resour. 1992b). stand the changes in their microclimates. This definition is a major issue in the Endangered and threatened wetland Migratory species and more mobile species upcoming reauthorization of the Clean animal species. probably suffered less than those with Water Act in 1995.

Herptiles Birds Lepidopterans

Endangered Threatened Endangered Threatened Endangered Threatened

Blanchard’s Blanding’s turtle yellow-throated red-shouldered Powesheik swamp metalmark cricket frog warbler hawk skipper butterﬂy butterﬂy massausauga wood turtle trumpeter swan cerulean silphium borer rattlesnake warbler moth western ribbon Caspian tern hooded warbler snake northern ribbon Forster’s tern yellow-crowned snake heron queen snake common tern great egret red-necked grebe

138 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Existing state, federal, and private could increase somewhat; at least many wetland acquisition and easement programs wetland areas not being farmed or other- will continue for the foreseeable future. wise drained will be restored. These efforts There are also plans will also result in covering the Missis- better protection of sippi River (U.S. Since wetlands are so interspersed undisturbed wetlands Army Corps Eng. among the other major community types and improved 1991) and the Lake in the state, the benefits of protecting, management. The Winnebago Pool cumulative result restoring, and enhancing wetlands will Lakes (Wis. Dep. should be a better Nat. Resour. 1989) to contribute to the ecological health of these assurance that the enhance wetland communities, also. biodiversity of areas that have been wetlands statewide severely degraded. will be protected and The recent surge of effort to restore wet- enhanced wherever possible. Since wetlands on private lands enrolled in the lands are so interspersed among the other Table 11 Conservation Reserve Program will con- major community types in the state, the Endangered and tinue as long as the program continues. biodiversity benefits of protecting, restor- threatened wetland This work could be considerably enhanced ing, and enhancing wetlands will contrib- plant species. if the U.S. Congress and Department of Agriculture would commit to long-term funding of the Wetland Reserve Program; to Endangered Threatened date, Congress provided funding for 1992 only. auricled twayblade beaked spike rush Watershed-based, nonpoint pollution control programs will continue to expand. angle-stemmed spikerush false asphodel These activities will afford major opportu- prairie white-fringed orchid English sundew nities to work with private landowners to netted nut-rush lenticular sedge achieve water quality benefits and wetland floating marsh marigold coast sedge preservation and restoration goals. In many watersheds, restoring wetlands will be a hop-like sedge Michaux’s sedge major technique for achieving nonpoint chestnut sedge bald rush pollution objectives. bog rush calypso orchid A continuing driving force behind wetland acquisition, management, and pink milkwort round-fruited St. John’s wort protection in Wisconsin will be the desire tussock bulrush bog bluegrass to enhance hunting and fishing opportuni- lake cress white lady’s slipper ties. Organizations such as Ducks Unlim- ited, Wisconsin Waterfowl Association, alpine milk vetch marsh valerian Trout Unlimited, The Nature Conservancy, crow-spur sedge linear leaved sundew the Wisconsin Wildlife Federation, the brook grass marsh grass-of-parnassus Conservation Congress, and others will continue to be very active in assuring that hemlock-parsley ramshead lady-slipper orchid state policies protect wetlands. These beak grass small round-leaved orchid groups now are also working with other chestnut sedge Garber’s sedge conservation groups to integrate their specific interests with broader goals for umbrella sedge sweet coltsfoot water quality, nongame wildlife, soil Fassett’s locoweed algal-leaved pondweed erosion control, and aesthetics, all of which heart-leaved plantain sheathed pondweed contribute to the protection of biodiversity. seaside crowfoot If all of these programs and efforts are continued, wetland acreage in Wisconsin small yellow water crowfoot

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ute to the ecological health of these com- highway safety—lead to compromises to munities, also. mitigate wetland losses. While the goal is to achieve no-net-loss of wetlands, it is very difficult to replace all the functions and ACTIONS CAUSING CONCERN values of wetlands that are lost to highway development, particularly in the immediate Although the era of large-scale, area of the loss. federally subsidized wetland drainage and Harvesting of forest products can filling has been assumed to be over, affect forested wetlands, mainly through changes in federal policy could dramatically changes in the microclimate when over- alter the current condition. Nationally, story trees are removed and soil is com- wetlands are afforded major protection due pacted by equipment. to the provisions of sections 404 and 401 Wetlands will continue to be affected of the Clean Water Act and the 1985 and by agriculture through grazing, barnyard 1990 Farm Bills. These provisions were and feedlot runoff, pesticide and fertilizer recently under serious threat by attempts to runoff, sedimentation from nonpoint redefine what a wetland is in the federal sources, and drainage. Landowners not manual for identifying and delineating participating in federal commodity support wetlands (Wis. Dep. Nat. Resour. 1992b). programs may still drain wetlands. Cran- The proposed changes would have more berry operations have the potential to affect narrowly defined wetlands, and the impact wetlands by converting existing wetlands to would be highly significant. In Wisconsin, cranberry beds, through the application of for instance, under the proposed 1991 pesticides and through the development of Wetland Delineation Manual, as much as water storage reservoirs. 80% of the state’s wetlands would not fall Agriculture in the United States and under the new definition and thus not be the world is undergoing major change. Free afforded the protection they now have trade agreements, the changes in Eastern (Dale Simon, Wis. Dep. Nat. Resour., pers. Europe, the demise of the Soviet Union, comm.). and the national deficit all affect U.S. In coming years, wetland-filling will agricultural trends. If remaining wetlands continue to be an increasing threat to are to be preserved, it will be necessary to wetland areas, as pressures for nonagricul- incorporate their protection into sustain- tural land use become more intense. able-agricultural policies that recognize the Shoreline development on inland lakes is need to be sensitive to ecological values continuing but is subject to county regula- while producing the food, fiber, and other tion. Since most of the best lakeshore products needed by society. In the U.S., the properties have already been developed, Clean Water Act and the Farm Bill are due those that remain are less desirable; some- for reauthorization in 1995. Wetland- times these are wetland areas that the agricultural issues will be major consider- owner wants to fill for development. The ations in both acts. loss of these wetlands would have negative The invasion of wetlands by exotic implications for water quality and wetland plant and animal species is a significant species habitat. Application of existing problem. For example, reed canary grass regulations will be required to prevent has been an extremely aggressive invader of negative impacts. sedge meadows. It has significantly dis- Highway construction also continues placed native species on many thousands of to affect wetlands. Wetlands often cannot acres of sedge meadows and shrub carr in be avoided during highway corridor the southern parts of the state. When this selection due to concerns for human safety, plant dominates a site, other species are farm operations, industry, and historical excluded and the community becomes use patterns. Thus competing public highly simplified. Controlling reed canary purposes—i.e., wetlands protection and grass is very difficult and expensive.

140 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Similarly, purple loosestrife is rapidly originally, which favors shrub and tree invading many wetlands throughout the growth over herbaceous vegetation. Restor- state. It is an exotic species, first released in ing fire as a natural process in wetland this country by nurseries and gardeners, communities can be highly beneficial and it crowds out native species. Control of (Payne 1992). this plant is very difficult, labor-intensive, Beaver can have major effects on costly, and controversial, since herbicides wetlands. From a positive standpoint they may be necessary (Payne 1992). Research can help maintain water levels and set back has been conducted on importing weevil succession into herbaceous wetlands. On species from their original habitats in the negative side, their dam building Europe as a biological control; results show activity can severely affect communities promise. So far a dozen states have released such as fens and bogs when associated This Wisconsin River weevils for loosestrife control. Wisconsin plant and animal life is replaced by persis- floodplain forest is did its first release in 1994. tent high water levels. In recent years high dominated by silver maple. Many wetland Common carp, another exotic species beaver populations in many parts of communities, including imported from Europe, has had serious Wisconsin have undoubtedly had a wide this southern wet- negative effects on many wetlands associ- variety of effects on wetland communities. mesic forest, are dependent upon ated with lakes and rivers. During feeding, The long-range effect of elevated beaver seasonal flooding. carp root out aquatic plants, causing populations on wetland community Elimination of this turbidity that prevents the regrowth of biodiversity is unknown even though local water recharge can plants and greatly reducing aquatic inverte- effects may appear quite severe. change the character of a wetland over time. brate diversity and abundance. Wetlands Many wetlands are dependent upon Photo by William Tans. with high carp populations have noticeably seasonal flooding. Elimination of this water less abundant wildlife populations than similar types of wetlands without carp. Control is difficult; the best that may occur would be periodic population reductions using intensive harvesting or chemical treatment (Payne 1992). The aquatic plant and wildlife response following a major reduction in carp populations in a wetland is very dramatic. The lack of fire in some wetland communities results in gradual invasion of woody shrubs and trees, eventually leading to a change in the wetland type. This is most significant for sedge meadows, fens, and shrub-carrs (Curtis 1959). As sedge meadows and other seasonally flooded wetlands convert to dense shrub and forested wetlands, the wildlife species needing open, herbaceous habitat are replaced by those preferring forest and dense shrubs. In Wisconsin prior to Euro- American settlement, this condition was dynamic. During drought periods, wetlands often burned—which set back succession—and often “peated in,” creating shallow open-water depressions. Many wetlands that have had their hydrology permanently disrupted by drainage systems are now generally drier than they were

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recharge can drastically change the charac- wetland restoration because of the growing ter of a wetland over time. Drought cycles public recognition of wetland values to can be beneﬁcial to wetlands through society and the economy. enhanced recycling of nutrients (Sloey et al. Despite this trend towards greater 1978), but prolonged drought can result in protection, wetlands will continue to be substantial vegetational changes. For affected by agriculture, highway construc- example, cattail and shrub invasion during tion, commercial navigation, and urban/ dry periods can be so dense as to exclude suburban development. In our society, it is wetland species that require a major open- probably not realistic to assume every water component (van der Valk and Davis remaining acre of wetland can or should be 1978, Weller 1981). During drought preserved. However, enough is now known periods many wetlands can also be farmed, about wetlands, their values, and their resulting in disturbance from cropping or functions that any proposed permanent loss grazing. During the period prior to Euro- must be very carefully considered. American settlement, buffalo and, perhaps, Wetlands are also important for herds of elk had signiﬁcant impacts on recreation, aesthetics, and education. They wetland vegetation in some parts of the provide open spaces in landscapes that are state. Grazing and trampling probably becoming increasingly rare as development helped maintain a herbaceous cover. continues. Hunters and anglers use them However, this use was seasonal and tempo- for recreational pursuits. They can be used rary, unlike the continuous grazing and seasonally for canoeing, hiking, and cross- trampling by domestic livestock that occurs country skiing. Viewing and listening to today in some wetlands. Thus controlled, wildlife are also popular wetland activities. periodic grazing can be used to maintain The bird life in wetlands is often particu- some types of wetlands (Payne 1992). larly easy to observe, making wetlands favorite bird-watching and photography areas. SOCIO-ECONOMIC ISSUES

The role of wetlands as essential POTENTIAL FOR COMMUNITY components in the healthy functioning of RESTORATION ecosystems has gained broad recognition in the last 20 years. Because so much wetland In assessing the potential for and acreage has already been lost, many regula- possible effects of restoring wetlands, the tory programs have been developed to specific characteristics of the types of protect remaining wetlands for the myriad wetlands and the types of disturbance of values they provide to society. While involved must be considered. Most perma- their value for wildlife and plant life has nently lost wetlands are those that have been most promoted, there is ever-growing been filled or excavated. Some disturbed awareness among ecologists and land-use wetland communities will readily respond planners that protecting wetlands for their to protection, restoration, and management flood storage, sediment and nutrient techniques but others may need many filtering, and groundwater recharge/ decades to return to a pre-altered state. discharge capabilities provides services to Because wetland communities differ, some our human communities that cannot be thrive with periodic disturbance while simply duplicated with engineered facilities others need long-term stability. Wetlands (Stearns 1978, Weller 1981). Thus, in the drained for agriculture often quickly future there will be more land-use planning respond to restoration efforts, since seed that avoids impacting existing wetlands and banks can lie dormant for many years (even more proposals that call for the restoration decades) waiting for the right conditions to of wetlands where possible. It appears we flourish (Weller 1981). Many wildlife may be at the beginning of an era of major species will re-inhabit wetlands within a

142 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE few years, some within days or months. 2. The effectiveness of existing federal, Once drainage has been stopped, the state, and local regulatory programs hydrological functions of a wetland may needs to be continually evaluated. The return somewhat to protection of exist- pre-drained condi- ing wetlands and the tions. The ability for While their value for wildlife and plant life restoration of drained and partially wetlands depends has been heavily promoted, there is ever- drained wetlands to upon the combined be restored to an growing awareness among ecologists and efforts and support ecologically functional land-use planners that protecting wetlands of many levels of level allows decisions for their flood storage, sediment and government interact- to be made regarding nutrient filtering, and groundwater ing with agricultural, how much wetland recharge/discharge capabilities provides business, industrial, acreage should be services to our human communities that and other interests. restored and where. Good communica- cannot be simply duplicated with Since much wetland tion and the creation loss has been due to engineered facilities. of shared goals and agriculture, it is highly values is essential to feasible to design prevent attempts at wetland restoration programs that fully weakening regulations to serve special integrate with water-quality and sustain- interests. able-agriculture programs. 3. State, federal, and local land acquisition of wetlands needs to occur in an Despite this trend POSSIBLE ACTIONS ecoregion context. Wetland complexes, towards greater rather than individual wetlands, have protection, The following possible actions are been and should continue to be the wetlands will consistent with ecosystem management, focus of acquisition. Wetland acquisition but require more analysis and discussion. programs should be integrated with continue to be How priorities are set within this list will be prairie and oak savanna acquisition affected by based on ecoregion goals, staff workload, programs, as these communities were agriculture, fiscal resources, public input and support, originally highly interspersed with highway and legal authority. We will work with our wetlands and have been the most construction, customers and clients to set priorities and severely reduced in acreage. Current commercial bring recommendations to the Natural public wetland acquisition efforts by the navigation, and Resources Board for consideration begin- Department, the U.S. Fish and Wildlife ning in the 1995-97 biennium. Service, or other public agencies should urban/suburban development. 1. Federal legislation and programs encour- be continued. The Natural Heritage aging wetland protection and restoration Inventory is capable of identifying high- need to be supported. U.S. Department quality undisturbed wetlands which of Agriculture policies linking participa- should be given protection from distur- tion in commodity support programs to bance. wetland protection need to be continued and enforced. For example, the Conser- 4. Better integration should occur among vation Reserve Program and the Wetland the goals and objectives of the many Reserve Program will need to be reau- interests in wetland restoration and thorized in the 1995 Farm Bill. The management involving Department Clean Water Act Sections 401 and 404 programs such as Wildlife, Fisheries, are also due for reauthorization in 1995. Water Resources Management, Forestry, Attempts to define wetlands politically Environmental Analysis and Review, rather than scientifically should be Water Regulations and Zoning, and opposed. Endangered Resources; federal agencies;

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city and county governments; and the restoration alternatives, including the many private organizations contributing use of local genotypes, and resulting money and time, such as Ducks Unlim- benefits to a wide variety of wildlife and ited, Wisconsin Waterfowl Association, plant life in a local area and region. and The Nature Conservancy. Ideally, wetland acquisition, protection, manage- 8. Riverine-floodplain wetlands along large ment, and restoration plans would be rivers in the state should receive addi- developed in partnership for each tional attention. These lowland and ecoregion of the state. A wetlands bottomland hardwood forest areas have management plan has already been diminished significantly in the state, and developed for Wisconsin as part of the the remaining acreage of these types North American Waterfowl Plan Joint should receive additional protection. Venture. This plan focuses primarily on Studies should be conducted to assess waterfowl, but it is an excellent docu- the feasibility of restoring these lowland ment with which to begin integrating forest wetland types. other wetland protection needs. 9. Coastal wetlands along Lake Michigan 5. Continued education and information and Lake Superior have been severely programs are needed to develop in- reduced in acreage. The remaining creased public support and understand- wetlands should be protected from ing of wetland protection and manage- development through regulation or, if ment activities. Wetland values, func- necessary, through easement or fee title tions, and protection and management acquisition. needs should be emphasized in primary and secondary environmental education 10.The issue of mitigation will have to be curriculums. Public-attitude surveys addressed. Currently, the Department should be conducted to assess knowl- has authority to mitigate only for edge of, use of, and interest in wetlands. Department of Transportation highway projects. Pressures to apply mitigation 6. The current 20-year cycle for updating for other types of development will Department wetland inventory maps is likely increase. The Department must inadequate for effective monitoring for assess the scientific and public policy state wetland protection and regulatory implications of mitigation to prevent the needs. A ten-year update cycle is desir- misuse of this concept, which can able but will require additional staff and contribute to the decline of biodiversity funding. The inventory mapping pro- of wetland communities. gram should continue to be integrated with the Department’s overall Geo- 11. Additional research should be con- graphic Information System program ducted to understand the long-term and the Department’s proposed Aquatic effects of using wetlands for stormwater and Terrestrial Inventory. and wastewater disposal. Additional research is also needed to better under- 7. Wetland restoration, development, and stand how nutrients, heavy metals, and enhancement projects should consider pesticides are cycled in wetland systems. the full range of biodiversity concerns. There is also a need to continue to Wetland restoration projects need to improve the Department’s knowledge assess the biological aspects of restoring base on how to best achieve wetland a wetland to its pre-altered state versus restoration and management objectives raising the water level above that which for a wide variety of plant and animal occurred before the wetland was altered. species and communities. This analysis should take into account the type of wetland that will result from

144 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Case Study

RESTORING A PRAIRIE WETLAND LANDSCAPE IN SOUTHERN WISCONSIN

Contributed by Alan Crossley.

Land for Patrick Marsh Wildlife Area was transferred to the DNR by the Department of Transportation (DOT) in December 1991, creating the first wetland mitigation bank site in Wisconsin. The land was purchased by DOT to allow the restoration of a large wetland area known variously as “Patrick Lake,” “Brazee Lake,” “Brazee Swamp,” “Duscheck’s Marsh,” “Phantom Lake,” “The Old Lake,” and more recently “Lake Sun Prairie.” The goal of the project is to recreate a microcosm of what Patrick Marsh and the surrounding landscape looked like when William Patrick first came upon it in 1841—a large, thriving wetland community surrounded on the uplands by oak openings and tallgrass prairie. The wetland restoration itself is different from most in that rarely do restorationists have a benchmark from which to evaluate the success or failure of the restoration, especially wetland restorations. Most of the time a wetland restoration merely attempts to restore the hydrology of a site, with no clear picture of what the wetland being restored looked like prior to drainage. Fortunately, we have lots of information about this site. From the original land survey notes of Orson Lyon in 1834 to the reconstruction of the history of the marsh (beginning in 1841) by Effa Duscheck as part of her address to the Twentieth Century Club of Sun Prairie in 1925, much is known about the marsh. Because of its importance to Sun Prairie life, pictures dating back to the late 1800s show it in various stages of inundation and drawdown. Aerial photographs beginning in 1937 again give a picture of the changing character of this dynamic wetland. And Dr. Robert A. McCabe’s study of the nesting ecology of water-obligate birds using the marsh from 1947 to 1951 describes bird use of the marsh and in particular notes the presence of the largest nesting colony of yellow-headed blackbirds in southern Wisconsin. His study also gives a glimpse into the species composition of the aquatic plant community. The marsh was drained in 1965 after a court battle in which the DNR tried, unsuccess- fully, to stop the drainage. But the recent expansion of State Trunk Highway 151 from two lanes to four lanes from Sun Prairie to Columbus set the stage for the cooperative restoration of the marsh as part of a wetland mitigation agreement between DOT and DNR. Soon after DOT removed the pumping system in the winter of 1991-1992, the marsh began to fill with water. By April of 1992 there were close to 100 acres of water on the marsh with an average depth of about 18 inches and a maximum depth of about three feet. More than 5,000 ducks and 200 tundra swans were observed on the marsh during spring migration. Surveys that year found 13 species of breeding birds using the marsh itself and an additional 26 species using in the uplands. Twenty-eight different species of aquatic plants were already found in the marsh, just six months after it began to fill with water. A survey of frogs and toads found only the American toad present in the marsh.

Continued on next page

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This 1937 air photo By the spring of 1993, the marsh shows Patrick Marsh as it was—a shallow filled to its normal level of about 160 acres marsh and wet of water with an average depth of almost meadow that five feet and a maximum depth of nearly supported a wide eight feet. Sixteen species of breeding birds diversity of plants and animals, including the were found using the marsh and about the largest breeding same number in the uplands. Aquatic plant population of yellow- diversity appeared to decrease slightly, headed blackbirds in southern Wisconsin. perhaps as a result of the deepening water Photo from Agricul- levels. But instead of hearing only the tural Stabilization and American toad, biologists heard six addi- Conservation Service. tional species of frogs. A graduate student working in the marsh found dozens of coot nests, as well as those of pied-billed grebe, sora rail, redhead, mallard, and blue-wing teal, to name a few. Several yellow-headed blackbirds returned to the marsh in 1993, although none were known to have nested. In 1994, water levels in the marsh In 1991, when this stabilized at their maximum level. Bird nest photo was taken, the density seemed to be reduced, although marsh was being nest success seemed to increase. At least drained and crops two pairs of yellow-headed blackbirds were being grown in it. The outline of the probably nested on the marsh. Tiger marsh, though, is still salamanders were also caught at the marsh clear. Photo from for the first time. Agricultural Stabiliza- tion and Conservation Service.

146 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE On the uplands, some progress has been made in restoring a few acres of prairie using locally collected native seed, thanks to funding support from DOT, lots of work by DNR wildlife managers, and great volunteer support from local citizens and Madison Audubon Society. During the winter of 1993-1994, many of the weedy tree species in the small wooded areas of the property were removed in favor of oaks and the native shrub understory. Every day, one can see a car or two parked outside the gates as people walk along the road or stop to watch birds. A Sun Prairie middle school teacher has been working with DNR wildlife managers to use the marsh as an outdoor classroom. During spring and fall, small groups of students come out to the marsh for an hour or two at a time to learn about the wetland, its unique history, and the plants and animals that live in it. A Wisconsin Environmental Education Board grant is also being used to develop an education program at the marsh for Sun Prairie elementary, middle school, and high school classes.

[Top] The pumping system was removed in the winter of 1991Ð LITERATURE CITED 1992, and the marsh began to fill with water. By April 1992, when Cowardin, L.M., V. Carter, F.C. Golet and Hale, J.B. 1982. Birds and wetlands. Wis. this photo was taken, E.T. LaRoe. 1979. Classification of Acad. Rev. 29(1):44-45. there were almost 100 wetlands and deepwater habitats of the acres of water. More Kabat, C. 1972. A capsule history of than 5,000 ducks and United States. U.S. Fish and Wildlife Wisconsin wetlands. Wis. Dep. Nat. 200 tundra swans Service, Washington, D.C. 103 pp. Resour., Madison 6 pp. [unpubl. rep.] were observed during that spring’s migration. Curtis, J.T. 1959. Vegetation of Wisconsin. McCormick, J. 1978. Management Photo from DNR files. Univ. Wis. Press, Madison. 657 pp. potential: summary and [Bottom] By June Dawson, T.J. 1982. Plugging the leaks in recommendations. pp. 341-55 in Good, 1993, when this photo Wisconsin wetlands protection law. R.E., D.F. Whigham, and R.L. Simpson, was taken, the marsh was filled to its normal Wis. Acad. Rev. 29(1):46-49.. eds. Freshwater wetlands: ecological level of about 160 Feierabend, J.S. 1992. Endangered species processes and management potential. acres. Sixteen species Academic Press, New York. of breeding birds were endangered wetlands: life on the edge. found using the same Natl. Wildl. Fed., Washington, D.C. 49 Payne, F.G. 1992. Techniques for wildlife marsh, and about that pp. habitat management of wetlands. same number were in McGraw-Hill, New York. 271 pp. the uplands. Photo from DNR files.

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Shaw, S.P. and C.G. Fredine. 1956. Vogt, R.C. 1981. Natural history of Wetlands of the United States. U.S. Fish amphibians and reptiles in Wisconsin. and Wildl. Serv. Circ. No. 39. 67 pp. Milw. Public Mus., Milwaukee. 205 pp. Sloey, W.E., F.L. Spangler, and C.W. Fetter, Weller, M.W. 1978. Management of Jr. 1978. Management of freshwater freshwater marshes for wildlife. pp. wetlands for nutrient assimilation. pp. 267-84 in Good, R.E., D.F. Whigham, 341-55 in Good, R.E., D.F. Whigham and R.L. Simpson, eds. Freshwater and R.L. Simpson, eds. Freshwater wetlands: ecological processes and wetlands: ecological processes and management potential. Academic Press, management potential. Academic Press, New York. New York. Weller, M.W. 1981. Freshwater marshes: Stearns, F. 1978. Management potential: ecology and wildlife management. Univ. summary and recommendations. pp. Minn. Press, Minneapolis. 146 pp. 357-63 in Good, R.E., D.F. Whigham, Wisconsin Department of Natural and R.L. Simpson, eds. Freshwater Resources. 1989. Winnebago wetlands: ecological processes and Comprehensive Management Plan. Wis. management potential. Academic Press, Dep. Nat. Resour., Oshkosh. 82 pp. New York. Wisconsin Department of Natural Tiner, R.W. 1984. Wetlands of the United Resources. 1990. Wetland losses in States: current status and recent trends. Wisconsin. Wis. Dep. Nat. Resour., U.S. Fish and Wildl. Serv., Washington, Madison. 1 pp. D.C. Wisconsin Department of Natural U.S. Army Corps of Engineers. 1991. Upper Resources. 1992a. Upper Mississippi Mississippi River system environmental River and Great Lakes Region joint management program, sixth annual venture—Wisconsin plan. Wis. Dep. addendum. U.S. Army Corps of Eng., Nat. Resour., Madison. 99 pp. Chicago. 44 pp. plus appendices. Wisconsin Department of Natural U.S. Fish and Wildlife Service. 1986. North Resources. 1992b. Wisconsin water American waterfowl management plan. quality assessment, report to Congress. U.S. Dep. Int., Washington, D.C. 19 pp. Wis. Dep. Nat. Resour., Madison. 220 van der Valk, A.G. and C.B. Davis. 1978. pp. Primary production of prairie glacial Wisconsin Department of Natural marshes. pp. 21-37 in Good, R.E., D.F. Resources. 1992c. Wisconsin wetland Whigham, and R.L. Simpson, eds. inventory classiﬁcation guide. Wis. Freshwater wetlands: ecological Dep. Nat. Resour., Madison. 4 pp. DNR processes and management potential. Pub WZ 023, Feb. 1992. Academic Press, New York. Visser, K. 1982. DNR wetland regulations. Wis. Acad. Rev. 29(1):44-45.

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one integral piece of a larger continuum that includes upland terrestrial systems and CHAPTER 10 transitional wetland areas. The location of a species or community along this continuum is critical to understanding its role in the landscape ecosystem. Wisconsin waters have been classified based on geographic locations. Frey (1963) Aquatic identified four major geographic regions: driftless area, northwestern lakes district, northeastern lakes district, and southeastern lakes district. A classification based on Communities nationally identifiable ecoregions was Compiled by DuWayne Gebken proposed by Omernik and Gallant (1988). Most of Wisconsin lies in four of these Bureau of Environmental Analysis and Review ecoregions: northern lakes and forests Department of Natural Resources (NOLF), north-central hardwood forest and (NCHF), driftless area (DRFT), and south- Michael Staggs eastern Wisconsin till plain (SETP) (Fig. Bureau of Research 17). Lyons (1989a) demonstrated that Department of Natural Resources Wisconsin stream fish communities show a general correspondence with these and ecoregions. Other ecoregion classifications Wendy McCown have been developed (e.g., Bailey 1989a, Bureau of Research 1989b) and will be used by the Department Department of Natural Resources to develop a classification system for the state. With significant contributions from Most classifications agree that the Robert DuBois, John Lyons, Don Fago, Richard Narf, Tom Pellett, driftless area is the dominant Wisconsin and Martin Jennings from the Bureau of Research; Dave Siebert geologic aquatic boundary. Covering an and Dreux Watermolen from the Bureau of Environmental area missed during the last glaciation, the Analysis and Review; Robert Hay and William Smith from the driftless area is distinguished by classic Bureau of Endangered Resources; David Heath from the North dendritic stream patterns, few natural lakes, and sharper, more eroded terrain (Becker Central District; Beth Goodman from the Bureau of Water 1983). In contrast, the remainder of the Resources Management; Kathy Patnode from the Bureau of state was smoothed by glaciation and has Wildlife Management; and David Ives from the Bureau of less topographic relief. Rivers are sinuous Fisheries Management. and have less average elevation drop. Glaciers also left substantial numbers of natural lakes. Lakes in northern Wisconsin tend to be cooler, more oligotrophic, and DESCRIPTION less productive than southern Wisconsin lakes. North-central Wisconsin also has one isconsin has a large and of the highest concentrations of spring-fed diverse aquatic resource lakes and streams in the world. which supports numerous Understanding the issues affecting species, communities, aquatic biological diversity in Wisconsin ecological processes, and must involve some generalization of aquatic human uses. In addition, ecosystem types. A general physical classifi- many terrestrial species and processes are cation includes drainage lakes—im- dependent on neighboring aquatic systems. pounded or natural lakes whose main water On a landscape scale, aquatic systems are source is from stream drainage and have at

150 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE least one inlet and one outlet; seepage lakes—landlocked natural lakes whose main water source is the groundwater table and with no inlet or permanent outlet; spring lakes—natural lakes for which the main water source is the groundwater table (springs) but always have an outlet of substantial flow; streams—smaller, low- order flowing waters which form the headwaters of river systems and which usually have a high-moderate gradient; and rivers—larger flowing waters formed by the confluence of several streams and that usually have a low gradient. The lake Northern Lakes & Forests classifications are derived from the Wiscon- (NOLF) sin Department of Natural Resources North Central Hardwood Surface Water Resources program (e.g., Forests (NCHF) Carlson and Andrews 1977). Southeastern Wisconsin Till Plains (SETP) LAKES Driftless Area (DRFT) Lake communities often vary dramatically based on limnological characteristics. natural acidity despite having typical ranges Figure 17 Lakes are often classified according to of nutrient input. These dystrophic lakes Ecoregions of trophic status. Lakes with very low nutrient contain unique communities that have very Wisconsin as input and abundant dissolved oxygen levels low species diversity and are among the developed by Omernik and Gallant (1988). throughout the water column are termed simplest of ecological systems. This system is used in “oligotrophic.” Oligotrophic lakes, like Lake Lakes normally undergo a natural this chapter and is one Superior, are often succession from example of how ecoregions could be considered to be the oligotrophic to defined for Wisconsin. epitome of desirable Wisconsin has a large and diverse aquatic eutrophic although water quality condi- the time span may be resource which supports numerous tions but have low thousands of years. overall productivity, species, biological communities, Human intervention ecological processes, and human uses. few species, and can shorten this On a landscape relatively simple process to a few scale, aquatic ecological systems. decades. Lakes systems are one Conversely, lakes with high nutrient input receiving unnaturally high nutrient in- or high rates of nutrient recycling are puts—termed “hyper-eutrophic”—have integral piece of a termed “eutrophic.” Eutrophic lakes that degraded habitat that results in simplified larger continuum thermally stratify may become devoid of communities, altered species compositions, that includes oxygen below the summer thermocline, and dysfunctional ecological processes. upland terrestrial precluding the production of many species. systems and Eutrophic lakes have high overall produc- transitional tivity and typically support high species GREAT LAKES wetland areas. diversities and more complex ecological Wisconsin waters include 1.7 million systems. Intermediate lakes with moderate acres of Lake Superior (Wis. Dep. Nat. nutrient levels and occasional oxygen Resour. 1988) and 4.7 million acres of Lake depletion are sometimes termed “me- Michigan (Wis. Dep. Nat. Resour. 1986) sotrophic.” Wisconsin also has a special including most of Green Bay. Wisconsin has class of lakes termed “dystrophic” or bog 156 miles of shoreline along Lake Superior lakes, which are primarily affected by and 407 miles of coastline along Lake

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The fish communities of Wisconsin’s Great Lakes are characteristic of north temperate oligotrophic and mesotrophic lakes.4 Cold-water communities with lake trout, rainbow trout, brown trout, and coho and chinook salmon as the top predators dominate, but warm-water communities featuring walleye, smallmouth bass, and northern pike exist in littoral and estuarine areas. Cold-water communities contain panfish and non-game species such as deepwater sculpins, bloater, cisco, lake, round whitefish, ninespine stickleback, longnose suckers, rainbow smelt, alewives, and sea lamprey. Warm-water communities contain yellow perch, burbot, white suck- With 6.4 million acres Michigan (Napoli 1975). Features of of surface water and ers, lake sturgeon, emerald shiners, and 563 miles of shoreline, national significance include the cobble carp. Both communities contain a mix of Wisconsin’s Great beach found along only the shoreline of the native and introduced species (Downs Lakes represent an Door County peninsula; Lake Superior immense resource. 1984, 1986). Wisconsin waters of the Great Geologic features, drowned bay mouth estuaries (e.g., St. Lakes at one time supported a complex of such as this exposed Louis River, Kakagon seven different cisco dolomite along the Sloughs, and Port species, four of Lake Michigan shore in Wing) found only Door County, add Understanding the issues affecting which were endemic structural and along Wisconsin’s aquatic biological diversity in Wisconsin to the Great Lakes functional diversity. shore; Lake Michigan must involve some generalization of (Becker 1983, Robins Photo by Robert H. drowned bay estuar- Read. aquatic ecosystem types. A general et al. 1991). ies (e.g., Marinette, The physical classification includes drainage Peshtigo, Green Bay’s macroinvertebrate Most of Wisconsin’s Atkinson’s Marsh) are lakes, seepage lakes, spring lakes, inland lakes are fauna of Lakes located in northern found primarily along streams, and rivers. Michigan and Wisconsin. Some, Wisconsin’s shoreline; Superior is domi- such as this lake in the Apostle Islands nated by amphipods Vilas County, are National Seashore located in Lake Superior remote and largely (especially Pontoporeia), oligochaetes, undisturbed. Photo by near Ashland; and Lake Superior itself—the nematodes, sphaeriids, and chironomids Michael Mossman. second largest freshwater lake in the world. (Cook and Johnson 1974, Dermott 1978, Nalepa 1989). Over 90 taxa of Chironomidae have been collected from southeastern Lake Michigan alone (Winnell and White 1985). A few types of typically lotic water forms such as heptageniids and hydropsychids are common in near shore areas (Barton and Hynes 1978) as well as being present in deeper water (Selgeby 1974). During the mid-1980s the European cladoceran Bythotrephes cederstroemi (BC) became established in Lake Huron and quickly spread to the other four Great

4 Fish and herptile species, for which data are plentiful, are well described in this discussion; other taxa are mentioned throughout the chapter wherever information was made available by contributors.

152 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Lakes (Garton and Berg 1990). Its impact Ecoregion on native zooplankton communities is unknown. It appears inevitable that BC will N. Central Northern SE WI Driftless Hardwood Lakes and Till All eventually spread to inland lakes in the Lake Type Area Forest Forest Plains Great Lakes region. Seepage

INLAND LAKES Number 164 1,837 5,966 404 8,371 Total Acres 1,106 28,253 95,864 8,790 134,013 Wisconsin has more than 14,000 inland lakes covering a million-plus acres Drainage (Table 12). Most lakes are located in the Number 132 922 2,715 255 4,024 northern part of the state. Using the Total Acres 27,548 34,375 146,316 10,494 218,733 Omernik and Gallant (1988) system of ecoregions, the NOLF ecoregion contains Impoundment 9,300 lakes covering 455,000 acres, but Number 261 447 601 239 1,548 85% are glacial or bog lakes of less than ten Total Acres 39,249 159,974 213,043 253,749 666,015 acres. The NCHF ecoregion contains another 3,200 lakes covering 223,000 All acres. In contrast, the DRFT ecoregion, Number 557 3,206 9,282 898 13,943 because of its steep topography, contains Total Acres 67,903 222,602 455,223 273,033 1,018,761 very few lakes—only 557 covering 68,000 acres. The SETP ecoregion contains only 6% of Wisconsin’s lakes but the region constructed on the Chippewa-Flambeau Table 12 includes Lake Winnebago, at 137,708 river system including Lake Wissota (6,300 Number and area of acres, the state’s largest inland lake. The acres), Lake Chippewa (15,300 acres), and Wisconsin lakes, by largest concentration the Turtle-Flambeau ecoregion as deﬁned by Omernik and of glacier kettle lakes Flowage (13,545 Gallant (1988), based in the world occurs acres). The Missis- on nearest county in the Vilas and Wisconsin has more than 14,000 inland sippi River in Wis- boundary. Oneida county area lakes covering more than a million acres. consin has a series of (Tonn and Magnuson The largest concentration of glacier kettle navigation dams 1982), and a high lakes in the world occurs in the Vilas and which have made existing riverine concentration of Oneida counties, and a high concentration NOLF Northern spring ponds occurs habitat and backwa- Lakes and of spring ponds occurs in the Forest, in the Forest, ter areas more Forest Langlade and Oneida Langlade and Oneida counties. lacustrine in charac- NCHF North county area (Carline ter. Smaller reservoirs Central and Brynildson occur on nearly every Hardwood 1977). river and stream system in the state. Dams Forest Most of these lakes are naturally have also been built on many natural lakes DRFT Driftless occurring and of glacial origin. However to control water levels. Area there are 1,550 dams on state waterways Fish communities in Wisconsin’s lakes which affect water levels on 666,000 acres are generally typical of warm-water me- SETP Southeast (65%) of Wisconsin’s inland lakes. A series sotrophic or eutrophic systems. They are Wisconsin Till Plains of hydropower reservoirs on the Wisconsin dominated by native species, including River system dominate central Wisconsin. largemouth bass, black crappie, northern The largest reservoirs are Petenwell Flow- pike, rock bass, and smallmouth bass. age (23,040 acres), Castle Rock Flowage Common insectivores include bluegill, (13,955 acres), Big Eau Pleine Reservoir yellow perch, pumpkinseed, and johnny (6,830 acres), Lake DuBay (6,700 acres), darter (Table 13). The most abundant and Lake Wisconsin (9,000 acres). Large omnivores are bluntnose minnow, golden hydropower reservoirs have also been shiners, white sucker, and common carp.

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Table 13 Percent of Sampled Lake Stations, by Ecoregion Comparison of percent fish species occurrences at stations in *** * * * Wisconsin lakes. Trophic Level NOLF NCHF DRFT SETP* Average Includes only fish and Species species found at > 10% of stations in at Top Piscivores least one region, as defined by Omernik Largemouth bass 59.5 66.5 95.5 49.8 67.8 and Gallant (1988). Black crappie 17.3 38.3 31.8 27.3 28.7 Northern pike 21.7 20.8 22.7 13.0 19.6 Rock bass** 21.4 20.8 9.1 11.4 15.7 Smallmouth bass** 12.6 16.9 18.2 6.4 13.5 Insectivores Bluegill 66.7 79.2 68.2 72.2 71.5 Yellow perch 72.0 70.7 36.4 56.7 58.9 Pumpkinseed 43.9 53.0 27.3 51.1 43.8 Johnny darter 44.1 38.9 40.9 20.4 36.1 Logperch 11.5 16.6 40.9 10.2 19.8 Spotfin shiner 2.3 7.9 45.5 11.9 16.9 Iowa darter** 32.3 21.7 0.0 13.3 16.8 Blacknose shiner** 23.3 9.0 0.0 15.2 11.9 Green sunfish 2.1 8.5 9.1 26.9 11.6 Spottail shiner** 7.6 3.1 27.3 7.2 11.3 Common shiner 14.7 15.8 4.5 5.6 10.2 Banded killifish 8.2 11.5 0.0 18.6 9.6 Brook silverside 5.3 1.7 9.1 21.3 9.3 Blackchin shiner** 15.2 12.1 0.0 9.2 9.1 Black bullhead 8.8 7.3 4.5 13.5 8.5 Mimic shiner 14.8 4.5 0.0 10.4 7.4 Orangespotted sunfish 0.0 0.0 27.3 1.5 7.2 Brown bullhead 3.9 11.3 0.0 7.2 5.6 Emerald shiner 0.2 2.5 4.5 13.2 5.1 Continued on next page A variety of herptiles inhabit lakes Wisconsin amphibians rely on ephemeral throughout the state (Table 14). Some waters for primary production. Five species amphibians use lakes, particularly their of turtles occupy natural lakes including shallow bays, for reproduction. In many the state-threatened Blanding’s turtle. While instances these are marginal breeding all five also occupy streams and rivers, all habitats with the exception of species but the eastern spiny softshell are most dependent on permanent water, such as the productive in lake environments. All but bull, green, mink, and Blanchard’s cricket the common musk turtle, which is limited frogs. The totally aquatic mudpuppy lives to the SETP and DRFT ecoregions, are its entire life on the bottom of lakes, usually found in all ecoregions of the state. in deep water (Vogt 1981). All other

154 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Table 13 (cont’d) Percent of Sampled Lake Stations, by Ecoregion Comparison of percent fish species occur- *** * * * rences at stations in Trophic Level NOLF NCHF DRFT SETP* Average Wisconsin lakes. and Species Includes only fish species found at > Omnivores 10% of stations in at least one region, as Bluntnose minnow 55.3 55.5 45.5 50.9 51.8 defined by Omernik and Gallant (1988). Golden shiner 24.8 21.4 27.3 23.1 24.1 White sucker 25.8 21.4 13.6 11.4 18.0 Table 14 Common carp 0.2 6.2 31.8 12.2 12.6 Herptile species Fathead minnow 11.2 15.8 9.1 8.7 11.2 occurring in Wisconsin lakes, classified by Bullhead minnow 0.0 0.6 13.6 0.2 3.6 ecoregions as defined Total stations sampled 660 357 22 624 1,644 by Omernik and Gallant (1988). *NOLF = Northern Lakes and Forest NCHF = North Central Hardwood Forest * * * * DRFT = Driftless Area Species Name NOLF NCHF DRFT SETP SETP = Southeast Wisconsin Till Plains Blue-spotted salamander** **Italics indicate a fish species intolerant of environmental degradation, as defined by Lyons (1992) Central newt ***Trophic level as defined by Lyons (1992) Eastern tiger salamander** Mudpuppy ** The macroinvertebrates of Wisconsin’s Spotted salamander inland lakes have not been intensively Blanchard’s cricket frogE H studied at a statewide scale. Preliminary BullfrogSC indications suggest that species of Cope’s gray treefrog** Chironomidae would make up 75% or ** more of the taxa for most lakes (Richard Eastern American toad Narf, Wis. Dep. Nat. Resour., pers. comm.). Eastern gray treefrog** Most species of Hemiptera, Coleoptera, and Green frog Diptera occur solely or predominantly in Mink frog inland lakes. There are no federal or state Northern leopard frog** endangered or threatened aquatic insects for which inland lakes form primary Spring peeper** habitat. Western chorus frog** Blanding’s turtleT IVERS AND TREAMS R S Common snapping turtle Wisconsin’s rivers and streams do not Common Map turtle form distinct trophic states. Energy systems Common musk turtle and species assemblages typically form a Eastern spiny softshell turtle continuum from smaller, upstream headwaters to larger, downstream rivers (Vannote Western/Midland painted turtle et al. 1980, Minshall et al. 1985). Rivers Northern water snake and streams may be classified into orders according to the number of branches or *NOLF = Northern Lakes and Forest ** = Breeding Habitat Only divisions from their mouth to their source NCHF = North Central Hardwood Forest E = State Endangered (Strahler 1957). Lyons et al. (1988) showed DRFT = Driftless Area T = State Threatened that there is considerable gradation of fish SETP = Southeast Wisconsin Till Plains SC = Special Concern H = Historic

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 155 AQUATIC COMMUNITIES

streams and some rivers in the northern part of the state can support a fish community with trout or salmon as the top fish predator. Smaller streams fed by surface water or located in the southern part of the state are typically warmer and support fish communities with smallmouth bass as the top fish predator. Larger rivers support only warm-water fish communities with smallmouth bass, walleye, largemouth bass, northern pike, or muskellunge as the top fish predators. Rivers and streams with trout or salmon are often classed as “cold- water” systems, while the other streams and rivers are often classed as “warm-water” systems. Cold-water systems are afforded special protection under state law.

STREAMS This category includes rivers and streams with mean annual flows of 40 cms A diverse system of species along Wisconsin’s flowing water or less (Lyons 1992). A definitive inventory headwater streams and of Wisconsin’s streams is not available, but tributaries feed into habitats. Rivers and streams may also be larger streams and classified by water temperature into warm- Becker (1983) indicates that of the 33,000 rivers throughout the water, cool-water, and cold-water systems. miles of rivers and streams in the state, state. Shown here is Species inhabiting these systems usually 9,561 miles are cold-water trout streams Mecan River in central Wisconsin, which reflect the maximum tolerable temperature (Wis. Dep. Nat. Resour. 1980). Adequate supports a trout fishery limiting the presence of various aquatic natural trout reproduction occurs in only and diverse species. 37% of the state’s cold-water streams. The macroinvertebrate community. Photo by In Wisconsin, rivers and streams are status of warm-water fish populations on Staber Reese. commonly classified by fish community most warm-water streams is not well types. Smaller, spring-fed headwater known.

Table 15 Percent of Sampled Stream Stations, by Ecoregion Comparison of percent ﬁsh species occur-

rences at stations in *** * * * Wisconsin streams. Trophic Level NOLF NCHF DRFT SETP* Average Includes only ﬁsh and Species species found at > 10% of stations in at least Top Piscivores one region, as deﬁned by Omernik and Gallant Brook trout** 46.5 32.0 15.9 3.4 24.5 (1988). Northern pike 11.5 18.9 7.6 28.7 16.7 Brown trout 13.4 21.4 22.0 7.9 16.2 Rock bass** 11.3 18.7 3.7 14.0 11.9 Largemouth bass 6.8 12.3 6.8 18.7 11.2 Smallmouth bass** 4.3 13.6 11.5 10.8 10.0 Burbot 14.0 10.8 3.3 0.5 7.1 Black crappie 2.5 7.1 2.8 11.4 6.0 Continued on next page

156 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Table 15 (cont’d) Percent of Sampled Stream Stations, by Ecoregion Comparison of percent fish species occur- *** * * * rences at stations in Trophic Level NOLF NCHF DRFT SETP* Average Wisconsin streams. and Species Includes only fish species found at > Insectivores 10% of stations in at least one region, as Creek chub 62.1 57.6 62.4 49.1 57.8 defined by Omernik Johnny darter 33.7 50.1 65.3 40.3 47.4 and Gallant (1988). Common shiner 44.2 48.9 29.1 38.7 40.2 Central mudminnow 48.3 52.4 16.8 40.5 39.5 Brook stickleback 40.4 31.4 39.0 32.8 35.9 Blacknose dace 45.0 38.6 40.3 15.8 34.9 Mottled sculpin** 45.1 29.7 7.1 10.6 23.1 Hornyhead chub 22.0 26.2 19.0 18.4 21.4 Fantail darter 5.4 19.1 30.4 14.6 17.4 Longnose dace 20.4 21.6 22.8 4.5 17.3 Black bullhead 6.8 18.8 5.2 35.5 16.6 Pearl dace 25.2 25.5 2.0 6.5 14.8 Blackside darter 9.3 26.6 10.8 10.1 14.2 Green sunfish 0.3 6.0 10.7 39.4 14.1 Pumpkinseed 6.2 19.7 4.1 21.4 12.9 Bigmouth shiner 0.8 8.8 26.0 12.9 12.1 Spotfin shiner 0.9 7.9 21.1 18.5 12.1 Bluegill 6.8 12.6 6.7 21.4 11.9 Northern hog sucker** 8.3 22.1 10.5 6.5 11.8 Yellow perch 15.0 12.7 1.6 13.0 10.6 Shorthead redhorse 5.6 7.1 14.8 8.9 9.1 Yellow bullhead 2.8 6.6 2.3 18.2 7.5 Stonecat 1.0 7.9 8.7 11.4 7.2 Sand shiner 1.1 2.7 9.5 14.5 6.9 Rosyface shiner** 1.1 10.4 9.0 4.0 6.1 Blacknose shiner** 10.3 10.2 0.4 2.9 5.9 Banded darter** 0.4 10.9 6.2 5.8 5.8 Rainbow darter** 0.8 11.9 1.9 4.8 4.9 Finescale dace 12.5 2.3 0.0 0.1 3.7 Suckermouth minnow 0.0 0.1 10.2 3.9 3.5 Continued on next page

A wide variety of warm- and cold- dace, white sucker, bluntnose minnow, and water ﬁsh species are found in Wisconsin fathead minnow. streams (Table 15). Common species Knowledge about macroinvertebrates include brook trout, creek chub, johnny of Wisconsin’s streams is still at the descrip- darter, common shiner, central tive stage where distributions of species are mudminnow, brook stickleback, blacknose becoming reasonably well known for many

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 157 AQUATIC COMMUNITIES

Table 15 (cont’d) Percent of Sampled Stream Stations, by Ecoregion Comparison of percent fish species occurrences at stations in *** * * * Wisconsin streams. Trophic Level NOLF NCHF DRFT SETP* Average Includes only fish and Species species found at > 10% of stations in at Omnivores least one region, as defined by Omernik White sucker 60.8 70.3 70.3 70.8 68.0 and Gallant (1988). Bluntnose minnow 12.0 29.6 42.1 42.2 31.5 Fathead minnow 16.0 30.2 30.3 40.4 29.2 Common carp 0.2 6.4 10.7 31.8 12.3 Golden shiner 7.3 10.6 4.0 13.4 8.8 Herbivores Central stoneroller 0.2 6.9 33.4 15.4 13.9 Brassy minnow 20.6 15.9 9.2 8.4 13.5 Northern redbelly dace 27.7 18.3 0.8 5.0 12.9 Southern redbelly Dace 0.0 1.9 20.3 13.7 9.0 American brook lamprey** 2.6 7.0 16.8 2.2 7.2 Largescale stoneroller 5.9 15.4 2.5 3.8 6.9 Total stations sampled 1,317 1,079 1,586 1,433 5,415

*NOLF = Northern Lakes and Forest ** Italics indicate a species intolerant of environmental NCHF = North Central Hardwood Forest degradation, as deﬁned by Lyons (1992) *** DRFT = Driftless Area Trophic levels as deﬁned by Lyons (1992) SETP = Southeast Wisconsin Till Plains

orders but significant gaps in knowledge these species. They are riffle species prefer- remain. Overall, the number of streams that ring clear, small, warm-water streams and have been studied in detail is small. No have been negatively affected by sedimenta- effort has been made to compare tion, dam construction, fish community macroinvertebrate faunas among manipulations, and point pollution dis- ecoregions. Aquatic arthropods can be used charges. They are now restricted to small to evaluate the water quality of streams reaches in watersheds where these effects based on the tolerance of the taxa to have been minimal. The rainbow shell organic and nutrient pollution (Hilsenhoff remains only in one five-mile reach of one 1987). Most species of Plecoptera, of the most well preserved SETP streams Ephemeroptera, and Trichoptera are found and is in immediate danger of extirpation solely or predominantly in streams. Two from effects of urban sprawl. dragonflies, two mayflies, and one riffle Several herptile species occupy beetle that inhabit streams and rivers are streams in Wisconsin (Table 16). The listed as state-endangered. Additionally, queen snake exclusively inhabits streams three dragonflies are listed as state-histori- and their riparian corridors in the SETP cal, suggesting they have been extirpated ecoregion. This state-endangered snake, from Wisconsin waters. while on the northern fringe of its range, Three state listed species of stream has declined in recent history as a result of freshwater mussels, ellipse, rainbow shell, water quality degradation including sedi- and slippershell, were once widespread in mentation and turbidity. The specific the DRFT and SETP ecoregions. Geo- microhabitat of this species in the stream, graphic ranges have decreased over 90% for flat rocky substrate, has been inundated by

158 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE sediments throughout much of its former range in southeastern Wisconsin. The Blanchard’s cricket frog, dependent on stream habitat and Wisconsin’s most endangered herptile, has seen a marked reduction in its range in Wisconsin and elsewhere throughout the northern limits of its distribution.

LARGE RIVERS Large rivers are those having a mean annual flow of 40 cms or larger (Lyons 1992). Wisconsin has 11 stretches of large rivers: the Mississippi River, the Wisconsin River below Tomahawk, the Chippewa The Mississippi River and Wisconsin River, shown here at River below the mouth of the Flambeau their confluence, are the most dominant riverine features Table 16 River, the St. Croix River below the mouth in the state. They are biologically rich and provide a major of the Clam River, the Fox River below the corridor for movement of species throughout the Herptile species watershed and region. Photo by Ken Beghin occurring in Wisconsin mouth of the Puchyan River and between streams, classified by Lake Winnebago and Green Bay, the ecoregions as defined Menominee River below the Highway 2/ by Omernik and 141 bridge, the Rock River below Lake Gallant (1988). Koshkonong, the Flambeau River below the * * * * confluence of the north and south forks, Species Name NOLF NCHF DRFT SETP the Wolf River below Shiocton, the Black River in LaCrosse County, and the Red Four-toed salamander** Cedar River below Menomonie. Most of Mudpuppy these river stretches have been dammed to produce hydropower. Blanchard’s cricket frogE H These large rivers support only warm- BullfrogSC water fish communities. The most abun- Green frog dant large predators are northern pike, walleye, smallmouth bass, largemouth bass, Mink frog channel catfish, and burbot (Table 17). Pickerel frogs Common middle trophic level species are Blanding’s turtleT bluegill, black crappie, yellow perch, rock bass, pumpkinseed, freshwater drum, and Common snapping turtle white bass. A large number of lower trophic Common musk turtleM level species have been found at sampled Eastern spiny softshell turtle river stations, but the most common are M spotfin shiner, shorthead redhorse, golden Western/Midland painted turtle redhorse, sand shiner, emerald shiner, Wood turtle common carp, johnny darter, logperch, Northern water snake northern hog sucker, white sucker, silver Queen snakeE redhorse, and bluntnose minnow. Wisconsin’s large rivers contain some *NOLF = Northern Lakes and Forest ** = Breeding Habitat Only of the highest freshwater mussel species NCHF = North Central Hardwood Forest E = State Endangered richness remaining in North America. The DRFT = Driftless Area T = State Threatened Wisconsin River contains 42 taxa, and the SETP = Southeast Wisconsin Till Plains SC = Special Concern St. Croix has 39. Some southern United M = Marginal Habitat States rivers contained more species but H = Historic

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Table 17 Percent of Sampled Stream Stations, by Ecoregion Comparison of percent fish species occurrences at Wisconsin *** * * * river stations. Includes Trophic Level NOLF NCHF DRFT SETP* Average only fish species found and Species at > 10% of stations in at least one region, as Top Piscivores defined by Omernik and Gallant (1988). Northern pike 54.4 36.0 23.0 38.5 38.0 Walleye 45.6 26.7 31.7 46.2 37.5 Smallmouth bass** 39.7 37.3 28.1 42.3 36.9 Black crappie 22.1 21.3 42.0 34.6 30.0 Rock bass** 30.9 24.0 29.9 7.7 23.1 Largemouth bass 4.4 13.3 42.9 19.2 20.0 Channel catfish 17.6 12.0 11.5 23.1 16.1 White bass 0.0 4.0 31.4 15.4 12.7 Burbot 32.4 9.3 0.6 3.8 11.5 White crappie 1.5 1.3 19.3 23.1 11.3 Sauger 0.0 5.3 22.4 11.5 9.8 Bowfin 2.9 2.7 11.2 7.7 6.1 Longnose gar 0.0 2.7 21.8 0.0 6.1 Yellow bass 0.0 0.0 6.6 15.4 5.5 Shortnose gar 0.0 4.0 10.9 0.0 3.7 Insectivores Spotfin shiner 51.5 70.7 67.4 50.0 59.9 Shorthead redhorse 52.9 44.0 37.8 42.3 44.3 Bluegill 16.2 37.3 58.6 42.3 38.6 Golden redhorse 63.2 40.0 16.3 15.4 33.7 Sand shiner 29.4 42.7 27.8 34.6 33.6 Emerald shiner 0.0 38.7 66.8 26.9 33.1 Johnny darter 29.4 32.0 41.4 15.4 29.5 Logperch 30.9 32.0 30.5 23.1 29.1 Northern hog sucker** 54.4 33.3 5.4 23.1 29.1 Silver redhorse 52.9 34.7 13.0 11.5 28.0 Yellow perch 39.7 24.0 38.1 7.7 27.4 Common shiner 69.1 21.3 3.6 0.0 23.5 Mimic shiner 27.9 38.7 12.4 0.0 19.7 River shiner 0.0 6.7 53.2 7.7 16.9 Brook silverside 7.4 17.3 31.1 7.7 15.9 Pumpkinseed 2.9 6.7 29.3 23.1 15.5 Spottail shiner** 2.9 2.7 43.8 11.5 15.2 Freshwater drum 0.0 8.0 33.8 19.2 15.3 River redhorse 29.4 24.0 1.5 0.0 13.7 Western sand darter 0.0 17.3 20.2 11.5 12.3 Continued on next page

160 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Table 17 (cont’d) Percent of Sampled Stream Stations, by Ecoregion Comparison of percent fish species occur- *** * * * rences at Wisconsin Trophic Level NOLF NCHF DRFT SETP* Average river stations. Includes and Species only fish species found at > 10% of stations in Hornyhead chub 41.2 2.7 0.9 0.0 11.2 at least one region, as defined by Omernik Blackside darter 29.4 12.0 2.7 0.0 11.0 and Gallant (1988). Greater redhorse** 25.0 10.7 0.0 3.8 9.9 Gilt darter** 32.4 6.7 0.0 0.0 9.8 Mooneye 0.0 14.7 16.3 7.7 9.7 Pugnose minnow 0.0 6.7 27.8 3.8 9.6 River darter 0.0 16.0 13.0 7.7 9.2 Smallmouth buffalo 0.0 8.0 13.3 15.4 9.2 Green sunfish 0.0 2.7 2.4 30.8 9.0 Spotted sucker** 0.0 8.0 23.3 3.8 8.8 Bigmouth buffalo 0.0 4.0 6.3 23.1 8.4 Blue sucker** 1.5 9.3 7.6 11.5 7.5 Black bullhead 4.4 4.0 0.9 19.2 7.1 Yellow bullhead 1.5 2.7 8.8 15.4 7.1 Bigmouth shiner 1.5 2.7 11.5 11.5 6.8 Slenderhead darter** 10.3 5.3 3.3 7.7 6.7 Tadpole madtom 1.5 1.3 18.1 3.8 6.2 Speckled chub** 0.0 2.7 10.0 11.5 6.0 Stonecat 5.9 0.0 1.8 15.4 5.8 Banded darter** 0.0 6.7 3.0 11.5 5.3 Silver chub 0.0 0.0 17.8 0.0 4.5 Orangespotted sunfish 0.0 0.0 17.5 0.0 4.4 Paddlefish 0.0 2.7 1.2 11.5 3.9 Creek chub 10.3 2.7 1.8 0.0 3.7 Longnose dace 14.7 0.0 0.0 0.0 3.7 Mud darter 0.0 0.0 10.3 0.0 2.6 Omnivores Common carp 8.8 33.3 40.5 46.2 32.2 White sucker 32.4 36.0 12.7 34.6 28.9 Bluntnose minnow 23.5 37.3 16.6 26.9 26.1 Quillback 8.8 25.3 37.8 23.1 23.7 Bullhead minnow 0.0 4.06 2.2 7.7 18.5 Golden shiner 7.4 9.3 29.0 3.8 12.4 Highfin carpsucker** 0.0 13.3 8.5 15.4 9.3 Gizzard shad 0.0 0.0 28.4 7.7 9.0 Fathead minnow 7.4 5.3 4.8 11.5 7.3 River carpsucker 0.0 2.7 10.9 7.7 5.3 Continued on next page

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Table 17 (cont’d) Percent of Sampled Stream Stations, by Ecoregion Comparison of percent fish species occurrences at Wisconsin *** * * * river stations. Includes Trophic Level NOLF NCHF DRFT SETP* Average only fish species found and Species at > 10% of stations in at least one region, as Herbivores defined by Omernik and Gallant (1988). Brassy minnow 17.6 8.0 5.7 3.8 8.8 Mississippi 0.0 2.7 18.1 0.0 5.2 silvery minnow** Parasites Chestnut lamprey 48.5 14.7 3.9 3.8 17.7 Silver lamprey 1.5 10.7 3.6 7.7 5.9 Total stations sampled 68 75 331 26 500 *NOLF = Northern Lakes and Forest ** Italics indicate a species intolerant of environmental NCHF = North Central Hardwood Forest degradation, as defined by Lyons (1992) *** DRFT = Driftless Area Trophic levels as defined by Lyons (1992) SETP = Southeast Wisconsin Till Plains

many of these species have been eliminated. Many of Wisconsin’s listed mussel PAST STATUS species have been eliminated or reduced by Wisconsin’s aquatic communities were water level manipulations, commercial shaped by the last glaciation. About 11,000 harvest, chemical treatments, fish commu- years ago, ice covered most of what is now nity manipulations, competition from Wisconsin, precluding the existence of exotics, channelization, dam construction, aquatic communities (Bailey and Smith and point and nonpoint-source pollution. A number of state 1981). The cold and turbid glacial meltwa- A variety of herptiles are found in ters draining through and federally listed Wisconsin’s rivers the DRFT would have plants are aquatic (Table 18) including eliminated all but the or riparian, and are the endangered Wisconsin’s large rivers contain some of simplest cold-water Blanchard’s cricket associated with the most diverse freshwater mussel communities. As the frog found in rivers river ecosystems. species associations remaining in North glaciers retreated, in the DRFT and the Wisconsin lists ten America. The Wisconsin River contains 42 aquatic organisms threatened wood endangered, ten recolonized turtle found in rivers taxa, and the St. Croix has 39. Wisconsin’s waters threatened and 36 in the DRFT and from the Bering (Lake species of special NOLF ecoregions. Agassiz), upper Mississippi, and Atlantic concern that are Some of the larger rivers have endangered refugia (Bailey and Smith 1981, Greene species of dragonflies. At times, these supported by river 1935, Stewart and Lindsey 1983). The dragonfly species are limited to specific ecosystems. glaciers receded and crustal rebound river reaches. Thus, they are vulnerable to alternately opened and closed connections changes in habitat from riprapping, dredg- between drainages until about 6,000 years ing, and modifications of velocities due to ago, when the current physical aquatic bridge construction. landscape emerged. A number of state and federally listed Quantitative surveys of Wisconsin’s plants are aquatic or riparian, and are aquatic resources were not made until the associated with river ecosystems. Wisconsin early 1900s. Consequently, descriptions of lists ten endangered, ten threatened and 36 Wisconsin’s earlier aquatic communities species of special concern that are sup- must be deduced from knowledge of ported by river ecosystems.

162 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE current aquatic community status; the few biota of major rivers (Junk et al. 1989). early, usually anecdotal, descriptions of Channel and flow modifications have aquatic resources in resulted in simplifica- the state; the few tion of these natural existing paleological Like terrestrial systems, aquatic systems processes. studies of aquatic are subject to the effects of simplification Fragmentation organisms; informa- of aquatic communi- and fragmentation. Most major tion on the nature ties is obvious in and scope of human simplification in Wisconsin has been cases such as dam activities that have caused by human activity, but natural construction, where occurred in the state; phenomena such as drought and forest migrations of fish or and our understand- fires have temporarily simplified other organisms are ing of the impacts aquatic systems. blocked. In other such activities can cases, severe simplifi- have on aquatic cation such as systems. channelization, Table 18 The aquatic resources of the state have dredging, or areas of poor water quality Herptile species been impacted and changed to varying have effectively fragmented aquatic com- occurring in Wisconsin degrees by human activities since the area munities. Fragmentation isolates popula- rivers, classified by ecoregions as defined was repopulated after the last glaciation. tions, thereby increasing the long-term by Omernik and Major changes began in the period of probability of loss of genetic diversity or Gallant (1988). logging and rapid agricultural development in the late 1800s and early 1900s and Species Name NOLF* NCHF* DRFT* SETP* continued through the industrialization of the 1920s to the 1960s into the current residential and recreational development Mudpuppy period. Blanchard’s cricket frogE H Aquatic systems are subject to simpli- BullfrogSC fication and fragmentation impacts just as with terrestrial systems. Most major simpli- Green frog fication impacts in Wisconsin have been Mink frog caused by human activity, but natural Pickerel frogsM phenomena such as drought and forest fires have temporarily simplified aquatic sys- Blanding’s turtleT tems. The impacts of simplification have Common map turtle included extirpation of native species, Common musk turtle reduced species richness, loss of top predator species, shifts toward more Common snapping turtle generalized-feeding or more disturbance- Eastern spiny softshell turtle tolerant species, reduced community False map turtle abundance, reduced genetic diversity, and community instability. Such impacts have Smooth softshell turtle commonly been caused by direct loss or Western/Midland painted turtle degradation of habitat, but they have also Wood turtle resulted from more subtle causes such as Northern water snake well-intended management activities (like stocking or chemical treatment), invasions Queen snake of exotic species, and commercial or sport *NOLF = Northern Lakes and Forest E = State Endangered fishing. Scientists are just beginning to NCHF = North Central Hardwood Forest T = State Threatened understand the critical importance of flood DRFT = Driftless Area SC = Special Concern events and subsequent aquatic-terrestrial SETP = Southeast Wisconsin Till Plains M = Marginal Habitat interactions in floodplains in shaping the H = Historic.

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Past and Present extinction due to random events. Fragmen- (Stark 1988). Smaller dams were main- Actions Causing tation has isolated migratory species from tained or constructed to create reservoirs Concern necessary spawning, nursery, or adult and associated lakefront property, control Dam Construction habitat. Fragmentation has also interfered water levels in natural lakes, or control Point-Source with recolonization of aquatic communities floods. Water level control structures were Pollution suffering from simplification impacts, even built in low-lying areas such as Horicon Agriculture after the impacts are corrected. Marsh to create and maintain wetlands for Non-Agricultural waterfowl habitat. A series of large dams Nonpoint Source Pollution and reservoirs was constructed on the Timber Harvest PAST AND PRESENT ACTIONS CAUSING Mississippi River to maintain a navigation Channelization and CONCERN channel for barges. Clearing of Dam construction can simplify and Streams fragment river habitats in a number of DAM CONSTRUCTION Invasion of Exotic ways. Most obviously, dams change riverine Species Over 3,700 dams of varying sizes Riparian Develop- (lotic) habitat into lake or reservoir (lentic ment have been built on Wisconsin’s rivers and or lacustrine) habitat. Since dams are Fish Stocking and streams. During the logging period, perma- generally built in areas where rivers have a Poor Understand- nent and temporary dams were constructed steeper vertical drop, higher gradient riffles ing of Genetic to provide power for saw mills and in- Diversity and rapids are eliminated. Reservoirs creased water flow to float logs down- Large-Scale created by dams can increase water tem- Chemical stream. These dams were built on almost all peratures and reduce dissolved oxygen Treatment major Wisconsin rivers, including the levels in water discharged below the dam. Department Chippewa, Flambeau, Black, Wisconsin, Dramatic changes in stream flow patterns Management Peshtigo, Menominee, Oconto, and Iron Priorities can disrupt spawning of native fish, reduce Habitat Improve- rivers, and on numerous smaller streams. macroinvertebrate habitat, and increase ment Projects In the southern part of the state, dams were erosion (Tyus 1990). Meffe (1991) and Water Level constructed to operate grain mills or for Winston et al. (1991) showed losses of Manipulations navigation. native species in a river system after Estuary Habitat In later years, many of the larger dams Management impoundments were built. Martinez et al. were converted to hydroelectric generation Lack of Monitoring (1994) documented that even small-scale Bioengineering to supply power for the paper mills that impoundments that do not radically alter Recreation grew up along the rivers or to generate hydrologic or thermal regimes can still have electricity for residential or industrial use a strong negative influence on native fish by facilitating establishment and proliferation of non-native species. Dams have allowed humans to harness the power of water and have provided recreational benefits in the form of reservoirs. However, dams can simplify and Dams also interfere with the natural fragment river habitats in a number of ways. Photo by F. Albert. flooding and sediment transport patterns in a river. Natural flooding and sediment flow patterns include periods of scouring and sediment deposition that maintain the complex gravel riffle, pool, run river habitats, and seasonally provide rich nutrients to floodplain areas. Disruptions of these patterns can result in loss of riffle and pool habitat, depletion of nutrients in floodplain areas, and loss of sandbars. Sedimentation in upper reaches of reservoirs can greatly alter wetland areas. Dams interfere with the natural downstream transport of woody debris which forms important habitat for macroinvertebrates, fish, and other aquatic organisms. Logs,

164 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE brush, and other debris that naturally enter river systems from riparian sources accu- Applying the Ecosystem Management Decision Model mulate behind dams leaving downstream to Aquatic Communities areas without this habitat. The list of past and present actions causing concern for aquatic Dams are typically impassable to upstream migration and pose mortality communities is lengthy, and the items on the list are complex threats to downstream-migrating species. and interrelated. All together, they point to the many dimensions The few fish ladders which do exist are old of the human relationship to water. It is a resource that connects and largely ineffective. No Wisconsin dams us in a myriad of seen and unseen ways to the components of are equipped for downstream fish passage the ecosystems upon which we depend. How will we make so migrating fish are exposed directly to decisions that recognize the role of humans as part of aquatic mortality in turbines or spillways. ecosystems and at the same time fully protect them for future Dams alter contaminant dynamics within aquatic systems. Spring high flows generations? flush contaminated water and sediments One positive step we can take is to begin to use and refine the from basins. Blockage of this cleansing can ecosystem management decision model described in the second cause accumulation within the reservoir particularly at the dam base. Contaminants chapter. This model provides a series of questions that in the collected sediments are then avail- managers can ask to approach decision-making from three able for resuspension in the water column perspectives: the ecological, socio-economic, and institutional. or uptake by bottom-feeding species. The Our success as resource stewards is a function of our ability to upstream flooding of riverine wetlands understand, analyze, and integrate alternatives across all three. produces elevated methyl-mercury in The conservation of biological diversity is one of the threads that mercury contaminated systems (Zillioux et weaves throughout the model as it is applied to the array of al. 1993). In Wisconsin, dam construction and actions that humans take to affect aquatic communities. operation has had major impacts on fish. The questions and considerations for managers to use to Becker (1983) noted that the gilt darter has address each of the three contexts are listed in the second been affected by dams because its preferred habitat, which is the large, fast-flowing chapter. However, there are two that deserve highlighting here. sections of rivers, has often been used as First, it is important that we apply the model on the landscape dam sites. Eddy and Underhill (1974) scale so that recommendations are made using the appropriate regarded the gilt darter population in the geographic boundaries. This will help us ask and answer the Saint Croix River as a “modern relict kind of broad regional questions that will guide the management population which has been isolated in of individual lakes. For example, how many lakes of different recent times by habitat modifications in its types are present in a region; what is their past, present, and former range.” The river redhorse, a state- threatened species, is declining in much of potential future condition; and what strategies are needed to its range due to dam construction (Becker conserve biological diversity and provide for the range of human 1983). uses? Fish such as the paddlefish, lake and shovelnose sturgeon, blue sucker, and Second, it is clear that DNR is not alone in this work; success skipjack herring and several mussel species will be measured by our ability to identify and include dependent on these fish for glocidial hosts stakeholders and to foster innovative partnerships with other are examples of species whose range has agencies, local governments, and private interests. been dramatically altered by dams (Becker 1983). According to Helms (1974), populations of shovelnose sturgeon have been sturgeon are restricted to areas immediately reduced in the Mississippi River due to below navigation dams. Construction of the habitat destruction resulting from several Keokuk Dam on the Mississippi River improvements to the navigation dams and (Lock and Dam 19 near Keokuk, Iowa) channel civil works. Now shovelnose presented a barrier to extensive upstream

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migration of paddlefish, American eel, poaching. Lake sturgeon have also been skipjack, Ohio shad, buffalo, shortnose gar, killed by hydroelectric equipment (Tom freshwater drum, carp, shovelnose stur- Thuemler, Wis. Dep. Nat. Resour., pers. geon, and three species of catfish comm.) and found entrained on dam trash (Carlander 1954). The dam interfered with racks (Tim Larson, Wis. Dep. Nat. Resour., sauger movement during the winter, and pers. comm.). spawning areas were cut off for the skipjack Dams have had an even more dra- herring, the Ohio shad, and the blue matic impact on Wisconsin freshwater sucker. The skipjack herring is the glocidial mussel populations. Mussels often congre- host for the ebony shell and elephant ear gate immediately below dams. Dams act as mussels. When the herring was extirpated barriers to upstream fish movement and from Wisconsin by construction of the fish are more likely to drop the mussels’ Keokuk dam, the ebony shell and elephant parasitic glocidial stage in areas immedi- ear mussels became endangered in Wiscon- ately below the dams (Robert Martini, Wis. sin occurring now only as scattered, old- Dep. Nat. Resour., pers. comm.). The age individuals (Becker 1983). increased velocities through the reach Becker (1983) reported that the below the dams may help scour the mussel paddlefish has also been affected by the beds clean of sediments. The upstream construction of dams and flood control reservoirs probably also help to supply projects that flood its spawning areas. It algae, diatoms, and other microscopic was once abundant in Lake Pepin, where its organisms that are food for filter feeders numbers are now considerably reduced. such as mussels (e.g., Ney and Mauney Lyons (1993) noted that paddlefish could 1981), some of which are very old. The not recolonize areas above the Prairie du concentration of these fish and mussels, Sac dam on the Wisconsin River following however, makes them susceptible to water quality improvements because the exploitation. Recently, the high price of dam prevented upstream movement. Heath mussel shells in Japan has resulted in (1993a) found that at least five mussel intensive mussel harvest and subsequent species were prevented from upstream closure of the mussel season in Wisconsin recolonization through the same dam. inland waters. Becker (1983) made similar observa- Hydroelectric facilities that conduct tions about the lake sturgeon. He noted peaking operations (varying flows to hydroelectric dams act as barriers to produce electricity for peak demand movement of lake sturgeon, isolating their periods) have an effect on downstream populations. Since lake sturgeon are long- habitats. The availability of stream habitat lived but reproduce slowly, they may persist is largely a function of stream discharge in an area for a long time, but they are (Trotzky and Gregory 1974, Milhous et al. susceptible to pollution, angler exploita- 1981, Bovee 1982, Bain et al. 1988, tion, poaching, and natural morality. Thus Leonard and Orth 1988). Changes in they may gradually die out without a discharge translate into changes in sub- source of adequate natural reproduction. strate, velocity, and depth conditions. These High spring flows through the gated section flow-dependent physical habitat features of the dams tend to attract spawning lake play an important role in governing the sturgeon, inducing some to drop their eggs. distribution and abundance of mussels Flows through the gates may later be shut, (Salmon and Green 1983, Neves and trapping the larger lake sturgeon behind Widlak 1987, Way et al. 1990, McMahon boulders, in plunge pools, and behind 1991, Strayer and Ralley 1993); conse- riffles (Joseph Kurz, Wis. Dep. Nat. quently, hydroelectric peaking operations Resour., pers. comm.). Any eggs that were can influence the availability of mussel deposited are then exposed to the air and habitat by creating wide fluctuations in eventual desiccation. The adults are subject discharge. Erosion and sand and silt to eventual death due to exposure or deposition have been implicated in decima-

166 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE tion of mussel beds on the Mississippi River diversity and abundance of metamorphos- (Stansbery 1970). Recent surveys by David ing juvenile amphibians (Pechmann et al. Heath (Wis. Dep. Nat. Resour., pers. 1991). In drought years especially, the comm.) indicate the only known popula- input to ephemeral ponds from early spring tion of winged mapleleaf mussel exists in snow melt and subsequent flooding may be the St. Croix River below the St. Croix Falls essential for amphibian recruitment. Dams hydroelectric dam, where it is subjected to can and often do eliminate or minimize the periodic exposure and desiccation due to opportunity for flood water to benefit water level manipulation. amphibians. The ecological effect of re- Dams constructed to alter water levels duced amphibian reproduction may be on natural lakes can change the aquatic significant since amphibians generally plant community. Large scale changes in represent high levels of biomass in decidu- aquatic plant communities, riparian and ous forests (Burton and Likens 1975), a littoral zone habitat, and water quality have habitat often associated with floodplains. occurred at least in part because of these The creation of dams has also converted artificial water level manipulations. many seasonal wetlands to more permanent Changes in water levels following dam water within the reservoirs. This is espe- construction have destroyed wild rice beds cially evident on the Mississippi River. on some waters (Vennum 1988). The Army Although these flooded wetlands are more Corps of Engineers has attempted to productive fishery waters, amphibian maintain a stable level in the Great Lakes in populations are reduced. The magnitude of accord with an agreement with Canada; losses of amphibian populations caused by however, the wetlands, spits, and sand flooding wetlands is unknown. beaches of the Great Lakes are shaped by Extensive dam construction in Wis- natural fluctuations in water level. The consin has reduced the available habitat for coastal marshes concentrate much of the riverine reptile populations, but the total biodiversity and productivity in the Great impacts are unknown. Painted and snap- Lakes and short- and long-term lake level ping turtles, which normally occupy slow fluctuation cycles are critical for sustaining flowing or standing water environments, the plant communities (Keddy and may displace riverine species like wood or Reznicek 1986, The Nature Conservancy map turtles in reservoirs. Impacts to 1994). When the operating levels of the amphibians by damming can also have Great Lakes were set, it is unlikely that direct impact on reptile species dependent consideration was given to the environmen- on amphibians for food. For example, the tal features that would be affected. The diets of garter snakes and northern water level of Lake Winnebago, the state’s largest snake consist primarily of frogs (Vogt inland lake, is also controlled by dams. 1981). The construction of dams and the Aquatic insect communities in the associated control of flood waters may presence of dams are qualitatively different affect the reproductivity of amphibians and usually less stable than those in un- within the floodplain ecosystems of regulated stream sections. The presence of dammed rivers. In free flowing rivers, an impoundment changes the habitat and spring snow melts and rainstorms can add quantity and quality of food released in considerably to flow levels resulting in downstream areas. Hydroelectric peaking frequent flooding of lowland areas adjacent operations result in large and rapid fluctua- to the river corridor, providing added tions in flows below dams (Cushman 1985) capacity for amphibian reproduction in the which can reduce species diversity, density, form of ephemeral ponds. Most of and biomass of aquatic insects in tailwaters, Wisconsin’s amphibians require ephemeral, with certain taxa affected selectively (Fisher fishless ponds for reproduction (Vogt and LaVoy 1972, Trotzky and Gregory 1981). The hydroperiod of ephemeral 1974, Williams and Winget 1979). Specific waters has a direct influence on both the problems include increased drift rates,

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which are known to accompany extreme owners/managers to work out agreements changes in flow (Radford and Hartland- to better protect the resources affected by Rowe 1971, Beckett and Miller 1982), and their operations. Wherever possible these stranding of stream insects in “intertidal hydroelectric facilities are encouraged to go zones” as waters recede (Kroger 1973, to a run-of-the-river flow regime in an Ward 1976, Extence 1981). Additionally, attempt to reverse effects of past peaking more time is required for aquatic insects to operations. At a minimum, studies should colonize habitats in rapidly varying flows be undertaken to determine the minimum than in unregulated flows (Gersich and levels of flow necessary to protect the flora Brusven 1981). Lentic insects have replaced and fauna of these rivers while still allow- lotic insects in impoundments resulting in ing hydro facilities to utilize this public net losses of lotic forms (Neel 1963, resource. Some successes have been Hilsenhoff 1971, Ward 1976). Changes in achieved, both through the regulatory energy processing in impoundments has process and by working cooperatively with usually led to substantial densities of the hydro owners/managers. The results are collectors and collector-gatherers in expected to benefit a variety of species, tailwaters but low densities of shredders including mussels, other aquatic inverte- and predatory insects (Spence and Hynes brates, amphibians, and fish. Dam 1971, Simmons and Voshell 1978). relicensing and regulation activities rarely Few new dams are being built at this consider abandonment as meaningful time, but renovation and expansion of options, and funds to remove dams are existing dams is common. The late 1980s limited. expansion of the dam at Jim Falls in Dam operation on the Mississippi Chippewa County created the state’s largest River and associated commercial barge hydroelectric facility. Recent interest in navigation continues to have impacts on renewable energy sources has lead to an that riverine ecosystem. Potential impacts increased number of hydroelectric develop- include conversion of riverine habitat to ment applications with the Federal Energy lacustrine, modification of normal water Regulatory Commission (FERC). Hydro- levels, sediment resuspension, dredging electric power is a “clean” energy source and channelization, and increased recre- because it produces no air emissions or ational use (Holland and Huston 1984, solid wastes. However, we do not have a Smart et al. 1985, Eckblad 1986, Holland complete understanding of the impact of 1987, Fremling et al. 1989). In recognition dam construction on biological diversity in of some of these problems, the U.S. Con- the affected river, although there is substan- gress established an environmental manage- tial evidence that modifications of the ment program with the objective of restor- natural flooding and sediment transport ing and monitoring habitat in the upper cycles in river systems can dramatically Mississippi River (Lubinski and Gutreuter simplify these systems. The Department 1993). may need to prepare to deal with the Dam construction has had many well- potential influx of hydropower develop- documented negative impacts on Wiscon- ment license requests. sin aquatic ecosystems, but it has also Under current FERC regulations, created additional reservoir habitat state- hydroelectric facility owners/managers are wide. Balancing the widespread losses of required to give equal consideration to the riverine ecosystems with gains in lake resource as is given to power generation. habitat—of which Wisconsin already had a This is a boost for environmental protection natural abundance—becomes a controver- of riverine ecosystems, especially compared sial proposition. Wildlife management with past regulatory requirements for hydro activities that impound streams for water- facilities. The Department is obtaining fowl management often increase habitat for valuable information about endangered and a variety of species, and have often been threatened species and working with hydro built on degraded or channelized wetlands.

168 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE However, such dams can still affect rivers wise not well documented. Paper and pulp and streams like any other dam. They may mills concentrated along the Wisconsin and increase nutrient loading to the impound- lower Fox Rivers were the major source of ment; disrupt movement of fish; change the pollution discharging both nutrient-rich character of existing wetlands from shrub, effluents and toxics such as mercury and sedge, or wooded to predominately open polychlorinated biphenyls (PCBs). Un- water; and disrupt water and sediment treated or poorly treated municipal sewage movement. On a few was a second major lakes, the presence of source of pollution in large numbers of many river systems. Many Wisconsin waters suffered severe waterfowl leads to Discharges of toxic increased eutrophica- simplification from the effects of industrial heavy metals occurred tion through the and municipal point-source pollution from in areas of heavy deposition of their the 1800s through the 1960s. Federal and industrial develop- fecal material. Some state Clean Water legislation has led to ment such as Milwau- flora, such as Fassett’s dramatic improvements in water quality kee, Racine, and locoweed, are inti- and the restoration of these aquatic Kenosha counties, and mately associated in central Wisconsin communities. However, the accumulation with specific lakes (Konrad and Kleinert and their unique of pollutants in sediments will remain a 1974). Impacts on water level character- source of contamination to the biota for an Wisconsin’s aquatic istics. Modifications extended period. systems from point- of these fluctuations, source pollution have changes in nutrient been severe in some levels, or pesticide inputs from groundwa- areas. Aquatic life including fish and fish- ter could threaten the existence of these eating birds suffered heavy mortality and plants. reproductive impairment in the Wisconsin and lower Fox Rivers and in localized areas POINT-SOURCE POLLUTION with heavy discharges (Becker 1983, Hauber 1989, Giesy et al. 1994). Many Wisconsin waters suffered Federal and state Clean Water legisla- severe simplification from the effects of tion has led to dramatic improvements in industrial and municipal point-source water quality in these areas and major steps pollution from the 1800s through the toward restoration of these aquatic commu- 1960s. Discharge of nutrient-rich sewage nities. However, the accumulation of effluent reduces dissolved oxygen causing pollutants in sediments will remain a direct mortalities of fish and other aquatic source of contamination to the biota for an organisms (e.g., Coble 1982). Discharge of extended period. Aquatic communities of toxic chemicals can also cause direct the Great Lakes are particularly susceptible mortalities and lead to build-up of toxic to substantial bioaccumulation of contami- materials in the aquatic system. Benthic nants due to their long water-residence invertebrate communities are simplified times. The approximate flush time in Lake through loss of species sensitive to water Superior is 182 years; in Lake Michigan it is quality and increased dominance of pollu- 106 years (Arimoto 1989). tion-tolerant generalist species (Cuffney et al. 1984, Chadwick et al. 1986, Camargo AGRICULTURE 1992). Heavy metals and organic chemical pollutants can bioaccumulate in fish posing Agriculture can have a dramatic a threat to wildlife and human health impact on aquatic ecosystems. Aquatic (Kleinert et al. 1974). systems are simplified by direct habitat Becker (1983) presents a discussion of destruction, erosion and sedimentation, this problem in Wisconsin which is other- hyper-eutrophication, and water quality

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degradation (e.g., Armour et al. 1991). prairies (Pflieger 1971). This species prefers Agricultural practices of particular concern quiet oxbows, ponds, and sloughs with are livestock grazing in riparian areas, mud, clay, and mixed sand and mud plowing and tilling of erodible soils (par- bottoms. The population of mud darter, ticularly in areas of steep terrain such as the another rare fish in Wisconsin, declined in DRFT), concentrated nutrient runoff from Illinois, due to decreased river size and barnyards and feed lots, pesticide and reduced flows (Smith 1968). Decreased nutrient runoff from fields, loss of upland river size and flows in Wisconsin could vegetation when forests and prairies are occur due to groundwater pumping, brought under cultivation, dredging and pumping for agricultural irrigation, or filling of wetlands, and channelization of droughts. Greene (1935) recorded the least streams. Almost all the agricultural chemi- darter in southeastern Wisconsin but recent cals in use are water soluble, resulting in surveys (Fago 1992) have not found the high mobility by species there. Ac- water transport and cording to Becker thus a significant Agriculture can have a dramatic impact on (1983) this loss may water pollution aquatic ecosystems. Practices of be due to increased problem with the particular concern are livestock grazing in turbidity and habitat potential for chronic riparian areas, plowing and tilling of destruction caused effects on aquatic by agricultural, erodible soils, concentrated nutrient runoff organisms (Sagar domestic, and 1991). from barnyards and feed lots, pesticide industrial pollutants. Agricultural and nutrient runoff from fields, loss of Specialist fish impacts on aquatic upland vegetation when forests and have been the most organisms in Wiscon- prairies are brought under cultivation, severely impacted. sin and other Mid- dredging and filling of wetlands, and For example, Becker western states are also (1983) notes the channelization of streams. well documented. gravel chub is limited Karr et al. (1985) to the lower Rock estimated that 44% River drainage of and 67% of fish species have disappeared Wisconsin and states, “the habitat require- or become less abundant in major Ohio ments of the gravel chub are so strict that and Illinois river systems and cited agricul- populations are isolated and confined to tural pollution as having had the greatest special riffle areas with special bottom impact. Erosion and sedimentation have types.” This specialization has made it degraded many stream channels, resulting vulnerable to turbidity and siltation, which in severe impacts to these and downstream increased as a result of agricultural activi- aquatic communities. Sedimentation ties. The creek chubsucker has probably profoundly changes stream insect popula- been extirpated from the southeastern part tions (Rosenberg and Wiens 1978, of the state, where it was at the northern Newbold et al. 1980, Lemly 1982, Culp et end of its range in the Des Plaines River al. 1986). Paleolimnological evidence from (Becker 1983). Becker (1983) believes Lake Mendota suggests there was a dra- erosion and habitat destruction in the matic increase in sedimentation and watershed eliminated the remnant popula- eutrophication after 1800, when agriculture tion of the creek chubsucker by the middle began in the basin (Kitchell and Sanford part of the twentieth century. 1992). Biological communities also became The Ozark minnow is noted by more unstable, suggesting increased Becker (1983) to be absent from a number perturbation of the aquatic community. of locations where it was previously re- One of the rarest fish in the state, the ported, apparently because it is intolerant bluntnose darter, may have been affected of excessive turbidity and siltation. Most of by increased siltation due to plowing of the the streams where the Ozark minnow was

170 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE located are characterized by heavy agricul- pesticides than other frog species (Sanders tural use. Becker (1983) also reports the 1970, Birge et al. 1979). These agricultural pugnose shiner, a state-threatened species, impacts may also be magnified through does not tolerate turbid conditions. bioaccumulation in amphibian prey sources The state-endangered queen snake has (Hazelwood 1970, Sanders 1970, Birge et also been impacted by erosion and sedi- al. 1980, Hall and Kolbe 1980, Linder et al. mentation resulting from agriculture in 1990). southeastern Wisconsin. This species has a very specialized diet consisting almost NON-AGRICULTURAL NONPOINT-SOURCE exclusively of crayfish (Vogt 1981) and POLLUTION requires a micro-habitat consisting of flat rocks on the stream bed under which it The U.S. Environmental Protection forages and seeks cover (Wood 1949). Agency (EPA) estimates that 50% of water Many of the streams once utilized by queen pollution in the U.S. is from nonpoint snakes have experi- sources (Barton enced heavy sedi- 1978). A 1985 survey mentation resulting indicated that 36% of Not all nonpoint-source pollution comes in a loss of exposed all Wisconsin’s rocky stream bed and from agriculture; it also results from urban streams and rivers are an associated reduc- stormwater runoff, use of fertilizers and affected to some tion or loss of cray- chemicals in urban areas, construction degree by nonpoint- fish populations site erosion, poorly designed or leaking source pollution (Gary Casper, Mil- septic systems, and poor land (Bergquist 1986a). Not all nonpoint- waukee Public management practices in non-agricultural Museum, pers. source pollution developments. comm.). comes from agricul- Agriculture also ture; it also results affects amphibian from urban populations in more ways than just by stormwater runoff, use of fertilizers and eliminating or altering their critical breed- chemicals in urban areas, construction site Nutrients from nonpoint pollution ing and foraging habitats. Frogs and erosion, poorly designed or leaking septic enter lakes and are salamanders have very thin, permeable skin systems, and poor land management recycled during spring and are vulnerable to chemical alterations practices in non-agricultural developments. and fall turn-over. Surface nonpoint pollution can include Excessive plant growth of their terrestrial and aquatic environ- and algae are often ments. The eggs and larvae are especially nutrient runoff, erosion and sedimentation, the result. Photo from susceptible. Amphibians are considered to and toxic substances. Loss of terrestrial DNR files. be excellent indicators of environmental health. Extremely high mortality and developmental abnormalities for some species are the result of toxicity caused by agricultural chemicals in aquatic systems (Hazelwood 1970, Birge et al. 1980). The Blanchard’s cricket frog, Wisconsin’s most endangered amphibian, has seen a dramatic decline throughout its historic range (Minton 1972, Christoffel and Hay, Wis. Dep. Nat. Resour., unpubl. data). While no specific cause has been implicated, it is suspected that agricultural chemicals (e.g., atrazine) are, in part, responsible for this decline. Hylid frogs in general, such as the cricket frog, may be more susceptible to

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vegetation in urban areas increases the in new developments will help but do not amount and variability of runoff events address problems from existing develop- contributing to flooding and erosion in ment. downstream areas. Contamination of groundwater and The addition of nutrients from surface waters from abandoned landfills nonpoint sources increases the nutrient and leaking underground storage tanks loading of the lakes and artificially acceler- continues. Inventory of these sites is ates the eutrophication process. Once a lake incomplete, and their contents are often not is overloaded with nutrients, they are hard known, but many may contain hazardous to remove, since the nutrients are continu- and toxic materials. The amount of con- ally recycled during spring and fall over- tamination depends on the rate at which turn. Increased nutrients cause increased the site fails, the content of the site, its algae or macrophyte growth. Excessive proximity to the aquatic resource, and the increases in plant growth are often domi- soils and geology of the area. However, nated by a few species reducing aquatic since the groundwater gradients are gener- plant species diversity. The proliferation of ally in the direction of surface waters, it will macrophytes into the entire euphotic area only be a matter of time before the con- of the littoral zone leads to loss of small taminated groundwater reaches a surface openings for fish spawning and creates an water. extreme amount of escape cover for young- Poorly designed and leaking septic of-the-year fish, which can become over- systems can lead to water quality problems populated and stunted. The resulting in unsewered residential areas. Lakefront competition for limited food resources can development is of particular concern adversely affect fish species and benthic because of its proximity to surface waters organisms that may be either a food source and higher than normal density of septic or a competitor for food. Decay of the systems. Lakefront developments are often increased plant biomass when it dies can in rural areas where connection to sewer result in decreased dissolved oxygen levels systems is very costly. Elimination of and kills of fish and other aquatic organ- nutrient inputs to lakes often does not isms. improve water quality because previously Changes in Wisconsin’s aquatic added nutrients are concentrated in lake systems caused by non-agricultural sediments and continuously resuspended nonpoint-source pollution are less well and recycled. documented than in agricultural areas, probably because they have been isolated in TIMBER HARVEST highly urban areas and masked by point- source and agricultural pollution problems. The impacts of silvicultural activities Since intense urbanization is a relatively in Wisconsin on water quality are not well recent phenomenon in most of Wisconsin, studied. Timber harvest within watersheds it is probable that urban nonpoint-source and along riparian areas has been shown to pollution has only recently been impacting affect water quality in other regions of the aquatic ecosystems on a statewide scale. country through increased runoff, sedimen- Except in a few isolated watersheds, rural tation, and temperature, and by reducing and urban nonpoint-source problems have primary productivity and dissolved oxygen not been controlled. The state’s major (e.g., Gray and Edington 1969, Hibbert nonpoint-source abatement activity is the 1969, Fredrickson 1970, Hornbeck et al. Priority Watershed program (Bergquist 1970, Hansmann and Phinney 1973, 1986b). The effectiveness of this program in Beschta 1978, Pearce and Rowe 1979, achieving results has been questioned, and Bernath et al. 1982, Hewlett and Fortson evaluation efforts have only recently been 1982, Lynch et al. 1984, Noel et al. 1986, initiated (Simonson and Lyons 1992). New Verry 1986, Hicks et al. 1991). laws requiring storm water retention basins

172 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Large woody debris normally resulting from streamside bank erosion or blowdowns plays an important role in stream and river morphology, hydrology, and ecology. Bilby and Ward (1989) studied the relationship of woody debris to the size of streams in western Washington. Large pieces of woody debris influenced channel morphology there through bank erosion, channel scouring, deposition, sandbar formation, nutrient and organic material retention, and species composition. However, the larger the river, the larger the woody debris needed to over- come the capacity of the river to move the debris downstream. The mean diameter, While most public lands have aes- Large woody debris length, and volume of woody debris thetic management zones to maintain the such as this fallen tree increased as channel width increased. plays an important role visual appeal of an undisturbed shoreline, in stream and river Murphy and Koski (1989) studied the rate harvesting practices on the backlands can ecology. Photo by of input and depletion of large woody still lead to erosion and disruption of Betty Les. debris in Alaskan streams. They found the overland water flow. Wisconsin has devel- rate of input and depletion was inversely oped a new program, Wisconsin’s Forestry proportional to the diameter of the debris. Best Management Practices for Water The model used predicted that 90 years Quality, which will help address these after a clear cut, large woody debris would concerns. be reduced by 70%, and it would take 250 years to return to prelogging levels. They CHANNELIZATION AND CLEARING recommended a 30-m wide unlogged buffer strip next to streams to maintain OF STREAMS large woody debris for input to streams. Streams have been straightened or Benke et al. (1985) showed that although channelized in the mistaken belief that woody debris accounted for only 4% of hydraulic efficiency was better for the habitat surfaces in a low gradient Georgia conveyance of flood waters brought on by coastal stream, they supported 60% of the runoff from pastures and intensively farmed invertebrate biomass and 16% of the cropland and denuded forest lands. Re- production for a river reach. Losses of moval of natural obstructions to navigation habitat elements such as large woody debris have also been commonplace, particularly can have effects for 80 to 160 years (Sedell during the period when rivers were exten- and Frogatt 1984, Sedell and Swanson sively used to transport logs. 1984, Minckley and Rinne 1985). Channelization is known to reduce species Although many of these studies are richness and diversity in fish, aquatic not specific to Wisconsin, the relationship invertebrates, and mussels, and to impact between water quality and logging practices other organisms such as furbearers that is important. Given the historical intensity depend on aquatic systems (Schneberger of timber harvest in northern Wisconsin, it and Funk 1971, Yokley and Gooch 1976, is likely that some forestry practices have Yokley 1977, Arner et al. 1979, Schlosser had similar water quality and habitat 1982, Kanehl and Lyons 1992). Further, it reduction impacts in Wisconsin’s aquatic can often lead to the instream disposal of systems. For example, Watermolen (1993a) dredge spoils which is detrimental to lists some specific streams in the upper aquatic life. Instream disposal directly Green Bay basin that have been impacted affects fish reproduction, benthos and water by recent forestry practices. quality (Morton 1977). The channelization

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and clearing of streams has eliminated species without an overall simplification of entire reaches of valuable aquatic commu- the community. Introduced exotics are nities for warm water, cool water, and cold often disturbance-tolerant, hardy general- water species. Reduced amounts of large ists having successfully survived human woody debris in streams can alter aquatic introductory mechanisms such as overseas insect community structure, especially in shipping or passage through pre- and post- rivers with a shifting sand bed (Dudley and export chemical treatments. These hardy Anderson 1982; Benke et al. 1984). species are well adapted to exploit already Channelization stressed and over- often results in the simplified biotic reduction of natural communities. In some edge along aquatic The establishment of exotic species or cases, exotics initially corridors and also hybrids in an aquatic ecosystem may explode in numbers results in the distur- initially appear to increase species but eventually stabi- bance of shoreline richness and diversity. However in the lize at a lower level of vegetation, opening long term, invasions of exotics may result abundance. It is the door for invading difficult to predict the in the loss of native species and the exotic plant species. impact that a new Many of Wisconsin’s disruption of habitats, predator-prey exotic species will herptile species rely relationships, and energy flow processes. have on an existing on these riparian aquatic ecosystem. areas for a great deal Non-native of the active season for shelter and foraging species have frequently invaded or have (Vogt 1981). Channelization does not likely been introduced to Wisconsin’s aquatic threaten most herptile populations but it is communities. Several key exotics uninten- certain their numbers are reduced by it. tionally gained access to the Great Lakes via The extreme impacts of the St. Lawrence Seaway, either transported channelization and dredging in Wisconsin’s in ballast water, attached to vessels (Moyle waters have been well documented. While 1991), or by direct migration. Invader these activities have been curtailed, permits species include the Asiatic clam, the sea are still sometimes issued. Smaller develop- lamprey, river ruffe, white perch, ment or maintenance projects are still Bythotrephes (a predatory cladoceran), and permitted by the state when the local the zebra mussel. Other species have been regulator does not believe the environmen- intentionally stocked, including the com- tal impacts outweigh the perceived benefits. mon carp, which was brought in with the Large navigation projects such as the best of intentions in the late 1800s. The Mississippi River are under federal control. introduction of this species is the most infamous example of a management action INVASION OF EXOTIC SPECIES that was thought to be beneficial at the time—but turned out to have devastating The establishment of exotic species or consequences (Courtney and Moyle 1992) hybrids in an aquatic ecosystem may which managers are still struggling to cope initially appear to increase species richness with today. Introduction of desirable and diversity. However in the long term, species such as brown and rainbow trout invasions of exotics may result in the loss of have had unknown impacts on native native species and the disruption of habi- brook trout. The grass carp, a more recent tats, predator-prey relationships, and introduction, is now reproducing in the energy flow processes. Exotic species often lower Mississippi River. invade without the normal predators or Exotics can also be introduced parasites that control their numbers in their through releases of species used for bait. native ecosystems, and existing ecosystems There have historically been few controls or may be unable to accommodate the new monitoring of the harvest, transfer, or sale

174 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE of fish or aquatic invertebrates used for bait zebra mussels have been identified as (Threinen 1982). It is believed the rusty responsible for concentrating organochlo- crayfish was accidently introduced from rine pollutants and maintaining them in the Illinois by bait anglers. The impacts of rusty food chain (Stone 1994). crayfish on the communities in certain The impacts of exotics on Wisconsin’s waters have been great (e.g., Olsen et al. aquatic ecosystems are difficult to assess. 1991). Intentional introductions of brown and There are several well documented rainbow trout, Pacific salmon, striped bass, invasions of exotics in Wisconsin. Eurasian and grass carp are often cited as examples water milfoil was first discovered in Wis- of successful introductions of non-native consin in 1967 and has now spread to at species, but the long-term implications of least 75 lakes in 39 counties (Bode et al. these introductions are poorly understood. 1993). The explosive growth of this plant No exotic that has become established has can substantially alter native aquatic plant ever been eradicated, so the risks associated communities, interfering with recreational with introducing exotics are extremely use, impacting fish communities, and high. It is unlikely that any species have choking water intakes. been extirpated from Wisconsin because of The river ruffe is already the second exotics, but it is probable that native most abundant species in the St. Louis species such as brook trout have been River estuary, and biologists fear that it is a significantly reduced in abundance and predator of whitefish eggs and that it can distribution by competition from exotics successfully compete against yellow perch (e.g., Waters 1983, Larson and Moore (Moyle 1991). The white perch now found 1985). The invasions of carp, river ruffe, in Green Bay also has the potential to sea lamprey, alewife, zebra mussels, white overtake the native yellow perch; however, perch, rusty crayfish, purple loosestrife, several studies have not found such im- and Eurasian water milfoil have already had pacts in Oneida Lake, New York, or Lake negative impacts on native ecosystems. Erie (Forney 1974, Schaeffer and Margraf Control of these invasions is already 1987). Fuller (1974) considers the Asiatic beyond the capability of any management clam to be a form of pollution itself. This agency. Management agencies across the species is a threat due to its free-swimming country, however, continue to propose larva and its ability to exploit any available introduction of new exotics. Most states substrate though there is no evidence to around Wisconsin have already allowed indicate that the Asiatic clam can success- introduction of the grass carp for control of fully compete against other clams and aquatic macrophytes with supposed mussels in Wisconsin as it does in some safeguards against their becoming natural- southern states. ized. Despite these safeguards, grass carp The zebra mussel has become estab- have successfully reproduced in the lower lished in Lake Michigan and the Mississippi Mississippi River (Allen and Wattendorf River, and its numbers have significantly 1987). Well-intentioned introductions of increased to date. This invader poses a largemouth and smallmouth bass in Texas significant threat to native mussels. Native have led to genetic introgression with the mussel populations have already declined endemic Guadalupe bass, which is now in some areas of the Great Lakes Basin due well on the way to extirpation in some river to the impacts of zebra mussels (Hebert et systems (Morizot et al. 1991). al. 1991; Mackie 1991). The potential Introductions of supposedly infertile impacts of zebra mussels on native bivalve sauger-walleye hybrids and stocking of populations have important implications sauger into native walleye waters have led for the upper Mississippi River, which has to genetic introgression between the two one of the most rich and diverse mussel species (Billington et al. 1988). Sauger- populations in the world (Cope, U. S. Fish walleye hybrids have been stocked in some and Wildl. Serv., unpubl. data). In addition, Wisconsin waters. North Dakota has

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already stocked zander, a close European RIPARIAN DEVELOPMENT relative of the walleye, into supposedly landlocked waters in the state (Terry Riparian habitat along Wisconsin’s Steinwand, pers. comm.). Its escape and lakes and rivers has been extensively establishment in native walleye waters developed since the mid-1800s. Shoreline would undoubtedly have a devastating development in populated southern areas impact on native walleye and sauger occurred early in this period where people populations. lived and worked. Development on north- New regulations to control the ern lakes and rivers came later and was introduction of exotics through Great Lakes initially limited by the remoteness of the ballast water exchanges have been pro- area and its sparse population. However, as posed by the U.S. Coast Guard. Wisconsin wages and leisure time increased, as has adopted new laws to eliminate the transportation improved, and as the state’s importation of exotic fish species. The population grew, more people were inter- Department has not allowed the use of ested in second homes. Cottages, resorts, grass carp and has taken steps to actively shacks, trailers, and all manner of dwellings eliminate populations that were discovered. were built. Before zoning laws existed, The Department has not proposed stocking some structures were built only a few feet other exotic species in recent years. In from the shoreline; trees and logs were addition, the Department has undertaken cleared from both water and shore, and public education programs designed to privies or septic systems were put in. The minimize spread of exotic organisms such level land and sandy shorelines suitable for as Eurasian water milfoil and Zebra mussel, beaches on well-known lakes disappeared and participated in monitoring programs first, followed by development on less for species such as purple loosestrife and desirable land that was steeper, rockier, or Zebra mussel. Whether these actions will marshier. Rates of development have continued to escalate in recent years. The Left undeveloped, be sufficient to prevent the continued riparian areas provide introduction of exotic species into Wiscon- number of lake front homes in Forest food and habitat for sin waters is unknown, but recent history County, for example, has increased 700% species such as this during the last ten years. In the Brule area, great blue heron. Photo suggests far more rigorous efforts may be by Bob Queen. needed. development has increased 19% for 200- 450 acre lakes and 78% for 100-124 acre lakes over the past 10-30 years (Korth 1993). Only a few isolated lakes escaped extensive development, including some large flowages such as the Turtle-Flambeau, Chippewa, Gile, Rainbow, and Willow, as well as small lakes or lakes on land owned by paper companies or public agencies such as the U.S. Forest Service, counties, and the Department. With riparian development came extensive loss and simplification of aquatic habitats. Owners of lake property commonly modified the shoreline or littoral area adjacent to their property by using sand blankets, shoreline protection such as riprap and retaining walls, docks and piers, boat houses, dredging for access, aquatic plant nuisance control, and filling. Disrup- tion of natural shoreline changed grada- tions in water depth in lakes, thereby eliminating natural formation of plant

176 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE communities (Keddy 1983), and similar species that are threatened or endangered development along streams causes changes in Wisconsin (Nilles 1993). in the structure of the macroinvertebrate The cumulative effects of numerous communities (Cummins et al. 1984, shoreline alterations may be detrimental to Sweeney 1993). Aquatic plant communities local amphibian and reptile populations were frequently directly altered through (Watermolen 1993b). Many amphibian mechanical removals or chemical treat- species dependent on the shoreline/water ments. Many alterations were done by interface (e.g. green, mink, and bull frogs) specific riparian property owners, but some are displaced when seawall construction municipalities have operated large-scale replaces the natural shoreline. The greater aquatic plant control activities. the loss of natural shoreline the greater the Isolated cases of shoreline modifica- impact to the local frog population. Aquatic tion may have little potential for affecting turtles, which need to leave the water to lay the aquatic community, but the cumulative their eggs on land, are affected by shoreline effects of numerous alterations can have barriers. Several species of Wisconsin significant and long- turtles show strong lasting impacts due signs of nest site to habitat loss and Isolated cases of shoreline modification fidelity. When turtles simplification (Panek may have little potential for affecting the are prevented imme- 1979). Some Wiscon- aquatic community, but the effects of diate access to these sin lakes, such as numerous alterations can have significant sites because of Shawano Lake, have shoreline develop- and long-lasting impacts due to very little natural ment, they are forced shoreline left. Lyons cumulative habitat loss and simplification. to expend additional (1989b) reported a time and energy significant simplifica- searching for a new tion of the littoral zone fish community of site or travelling indirectly to their tradi- Lake Mendota since 1900 and attributed tional site. This exposes them to potentially the changes in part to increased shoreline higher mortality since most aquatic species development. Bryan and Scarnecchia have little natural defense on land. What (1992) found significantly fewer fish effect this has on populations is unclear. species and reduced abundance in devel- Despite the well-documented negative oped shoreline areas in an Iowa lake. Miller impacts of riparian development, it contin- et al. (1989) conclude that habitat alter- ues on Wisconsin’s waters at a rapid pace. ation was a factor in 73% of fish extinctions There are few legal restrictions to develop- in North America during the past 100 ment, a situation difficult to change be- years. Kapuscinski and Jacobson (1987) cause of the high demand for and value of explain that habitat alteration can impact lakeshore property. The Department genetic diversity by reducing the effective controls permitting of erosion control population sizes and changing selection structures and lake bed modifications, but pressures on previously well-adapted such decisions are increasingly being species. challenged in legal forums. Increasing Lakes were also affected by the evidence also suggests that riparian activi- draining and filling of wetlands, which ties beyond the ordinary high water mark, supported waterfowl, reptiles, and amphib- which are not controlled by the Depart- ians, northern pike, and muskellunge ment, have impacts on the systems. Re- spawning areas. Loons, ospreys, eagles, search has been done in Canada to predict otter, muskrat, and mink were all affected sustainable levels of lakeshore development by habitat degradation and harassment (Dillon and Rigler 1975), but these meth- resulting from increased use. Habitat loss ods have not been applied in Wisconsin. and urbanization have been implicated in Filling of riparian wetland areas has been reducing populations of several dragonfly dramatically curtailed, but is still a concern.

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Riparian areas are also increasingly and whitefish was done as early as 1876. In impacted by beaver activities. Wisconsin 1937, Wisconsin stocked over a billion fish beaver populations have been increasing in of various species. recent years due to Currently the elimination of natural Department annually predators, reduced Fish have been artificially propagated and stocks on average trapping pressure, widely stocked in Wisconsin for more than about 615,000 brook and habitat manage- trout, 2.2 million a century. Fish have also been routinely ment practices that brown trout, 940,000 increase aspen and moved from one water body to another, rainbow trout, willow. Beaver and new species have been widely 280,000 lake trout, activity can have a introduced . . . . While these efforts 260,000 splake significant impact on reflected the best understanding of fish (brook trout x lake riparian areas, management at the time, there is growing trout hybrids), 2.3 including damming million chinook evidence that stocking or transfers of fish of streams, cutting of salmon, 659,000 trees, and flooding of can have long-term negative impacts on coho salmon, low-lying areas. growth, survival, reproduction and even 513,000 largemouth While moderate health of both the existing fish population bass, 63,000 small- levels of beaver and the newcomers. mouth bass, 169,000 activity are probably muskellunge, 28,000 necessary to maintain hybrid muskellunge native habitat patterns along streams, (northern pike x muskellunge hybrids), excessive levels are thought to disrupt fish 60,000 northern pike, 2,300 lake sturgeon, movement, alter sedimentation patterns, 3.5 million walleyes as fingerlings, year- and increase water temperatures, adversely lings, or catchable-sized fish. Another affecting cold-water communities. 65,000 largemouth bass, 850,000 muskellunge, 344,000 hybrid muskellunge, 15 FISH STOCKING AND POOR UNDERSTANDING million northern pike, 61,000 lake stur- OF GENETIC DIVERSITY geon, and 51 million walleyes are stocked as newly hatched fry. Other species stocked Fish have been artificially propagated periodically include channel catfish, sauger and widely stocked in Wisconsin for more and several species of panfish (Dave Ives, than a century. Fish have also been rou- Wis. Dep. Nat. Resour., pers. comm.). tinely moved from one water body to Trapping and transfers of adult fish another, and new species have been widely was common from 1874 until the 1930s introduced. The magnitude of the fish- (Becker 1983). Fish were often “rescued” stocking and transfer program in Wisconsin from waters expected to experience win- since 1874 is staggering. Virtually every terkill or from flooded backwater areas of major lake and river has been stocked at the Mississippi River as they were stranded one time or another by the Department or by receding flood waters in the spring. private individuals. Becker (1983) presents Rescued fish were transferred to waters an excellent summary of the evolution of across the state. In 1936, nearly ten million the Wisconsin stocking program. By 1900, fish, including catfish, sunfish, crappies, Wisconsin had attempted introductions of bass, and buffalo, were trapped and trans- Atlantic salmon, chinook salmon, grayling, ferred among various state waters. This rainbow and brown trout, carp, and activity became less common after 1940 goldfish. Walleye propagation began in but is still used in some winterkill and 1883 with production of eight million fry. panfish management situations. Muskellunge propagation began in 1897 While these efforts reflected the best with production of one million fry. Hatch- understanding of fish management at the ing and stocking of lake trout, brook trout, time, there is growing evidence that stock-

178 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE ing or transfers of fish can have long-term ment has also recently begun a major effort negative impacts on growth, survival, to quantify the genetic differences among reproduction and even health of both the watersheds for an additional eight warm- existing fish population and the newcom- water species and native brook trout. ers. Largemouth bass moved between Fish stocking and transfers can also Texas, Florida, Wisconsin, and Illinois impact biodiversity by introducing new invariably showed significantly lower species into aquatic communities, with growth and survival in non-native waters resulting changes in the relative abun- (Philipp and Whitt 1991, Philipp 1991). dances of the native species. Walleye, for Cutthroat trout stocking in Yellowstone was example, is not native to small seepage shown to have disrupted and reduced lakes in Wisconsin (Becker 1983), but this natural reproduction (Gresswell and Varley species is now found in almost every such 1988). A comprehensive literature review lake more than 200 acres in size. The examining releases of cultured salmonids impact of these introductions on the into native populations concluded that, existing aquatic community is not well when effects were seen, they were always understood, but Colby et al. (1987) docu- detrimental to the native stocks (Hindar et ment fish community changes resulting al. 1991). Hybridization, often resulting from species introductions. In Wisconsin, from stocking of different genetic strains, for example, introduction of walleye and was a factor in 38% of fish extinctions in northern pike into Escanaba Lake resulted North America during the last 100 years in long-term declines of smallmouth bass (Miller et al. 1989). Hatchery fish can and panfish populations (Kempinger et al. introduce poorly adapted genomes into the 1975). population and through introgression Release of bait fish and macroinverte- disrupt the genome of the existing naturally brates is another source of genetic mixing. reproduced fish (Magnuson 1976). Intro- Bait species are commonly harvested from ductions of different genomes caused by naturally occurring populations and releases of bait fish can have the same transferred to other waters for sale and effect. potential release. Cultured bait species are Genetically different stocks may exist also commonly sold but suffer the same for many important fish species found in potential genetic disruption as that caused Wisconsin (Kapuscinski and Lannan 1986). by any other hatchery-reared species. The Analysis of DNA pattern variations in bait industry in Wisconsin is lightly regu- walleye suggest that current stocks evolved lated and little information is collected on from three distinct glacial refugia, and the origin of fish that are sold. different walleye genetic types show clear Some species, such as walleye and regional distribution patterns (Billington lake trout, exhibit homing instincts during and Hebert 1988). Genetic differences spawning. Walleye and lake trout also seem among walleye stocks have been docu- to exhibit spawning preferences and mented within states (McInerny et al. requirements unique to specific strains. 1991) and even within the same drainage Whether stocking of hatchery-reared fish (Todd 1990). Similar differences have been has already affected this behavior is not documented for largemouth bass (Philipp known. In addition to the genetic effects of et al. 1983) and northern pike (Seeb et al. stocking, stocking large numbers of hatch- 1987). Only limited work has been done to ery-reared fish among relatively few natu- analyze genetic variability among stocks in rally reproduced fish can subject the Wisconsin. The Department is currently natural population to increased fishing funding a study of genetic differences pressure because fishing gear is nonselec- among different spawning populations in tive. the Lake Winnebago system. Preliminary The actual impacts of fish stocking results show limited allozyme variability and transfer activities on Wisconsin’s (Treloar and Ehlinger 1991). The Depart- aquatic communities can never be fully

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known. However, given the magnitude of poorly documented (Policansky 1993). the number of fish and waters stocked, and Highly selective gillnet fisheries were the likelihood of genetic and population implicated in long-term changes in lake changes, it is probable that there have been whitefish growth rate, condition factor, and significant changes in species abundances mean age (Handford et al. 1977). High and distributions across the state. Becker exploitation rates have changed growth and (1983) suggests many anomalous distribu- mean age in lake whitefish and walleye tion records may be due to these activities. populations (Healey 1980, Reid and Certainly the distribution of some major Momot 1985, Mosindy et al. 1987). Nuhfer game species such as walleye, muskellunge, and Alexander (1991) conclude that long- brown trout, and rainbow trout have been term angling exploitation may alter the dramatically increased, but it is also likely genetic composition of wild brook trout that species such as largemouth or small- strains. mouth bass and brook trout that existed Stocking and stock transfers remain prior to the introductions have suffered a important management practices in Wis- corresponding decline in abundance and consin. Adult stock transfers of northern distribution. Changes in lower trophic level pike and various panfish species between species are undocumented. Genetic impacts waters and watersheds still occur. Current of stockings of species in waters where Wisconsin hatchery practices and stocking there were already naturally reproducing policies do not consider genetic conserva- populations of those species are unknown, tion. For example, virtually all walleye and but it is likely that some populations have muskellunge hatchery production comes suffered declines in natural reproduction. from the Spooner and Woodruff hatcher- This evidence has led to numerous ies—both located in far northern Wiscon- recommendations against mixing different sin. All spawn is taken from local lakes, but genetic stocks. After an extensive survey of the hatched fish are distributed throughout salmonid stocking effects, Hindar et al. the state. Wisconsin also periodically (1991) recommend “strong restrictions on exchanges hatchery products with other gene flow from cultured to wild popula- states and federal agencies. tions and effective monitoring of such gene Today, managers are increasingly less flow.” In a reference work on fisheries interested in stocking hatchery-reared fish genetics, Kapuscinski and Jacobson (1987) and more interested in depending on the recommend managing to avoid stocking native stock for reproduction. Where and, when necessary, stocking only locally stocking is needed, attempts are being adapted fish. Both Meffe (1987) and made to use native brood stock. Programs Kapuscinski and Philipp (1988) conclude are being initiated to study genetic diversity that significant problems in conserving of fish within the state. existing levels of genetic diversity exist and that additional cooperation and research HARVEST will be needed to determine appropriate management strategies. The American Humans have been harvesting Fisheries Society has developed a draft Wisconsin’s aquatic resources for thousands position statement entitled “Protecting of years, and this harvest has undoubtedly Native Fish Stocks: The Elimination of had impacts on biodiversity. There is Stock Transfers,” which advocates a stock considerable evidence that Native Ameri- concept of management and restrictive cans used nets and spears and even built stocking and stock transfer policies de- dams and weirs to harvest fish (Kuhm signed to protect native stocks (Philipp et 1928) and turtles (Adler 1968). European al. 1991). settlers in the mid-1800s began commer- Population changes caused by long- cially harvesting sturgeon, lake trout, term or size-selective harvest have often suckers, yellow perch, and other common been shown; however, genetic changes are fish species.

180 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Today, sport, commercial, and subsistence harvest of some aquatic organisms is substantial. Sport harvest activities are primarily directed at game fish species. During 1991, 1.47 million licensed anglers and more than 2-million anglers over-all fished 21.3 million days in Wisconsin. Over 0.5 million anglers were non-residents—second only to Florida in fishing by tourists. Anglers most frequently fish for panfish (bluegill, pumpkinseed, yellow perch, rock bass) followed by walleye and sauger, largemouth and smallmouth bass, northern pike and muskellunge, and crappie (U.S. Dep. Int. / U.S. Dep. Comm. and larger fish, lower the average age of Fishing has been a 1993). popular sport in first reproduction, increase growth rates, Wisconsin for many Subsistence harvest is practiced by increase the variability in recruitment (e.g., years. This photo from many users to some extent when fish or Spangler et al. 1977, Coble 1988, SPOF the 1930s shows a other sport harvested animals are eaten. 1983), and may alter the genetic composi- day’s catch of muskellunge from Pine However, the only significant subsistence tion of a stock (Policansky 1993). Despite Lake in Vilas County. harvest of aquatic organisms currently these changes, there is little evidence that Photographer allowed is a Native sport angling alone unknown. American walleye and can collapse sport muskellunge spear fish populations. fishery. This fishery Today, managers are increasingly less Unrestricted angling was reinitiated in interested in stocking hatchery-reared since 1946 in 1986 and has been fish and more interested in depending on Escanaba Lake has conservatively regu- native populations for reproduction. not decreased the lated and extensively Where stocking is needed, attempts are walleye population monitored (Hansen being made to use native brood stock. (Staggs et al. 1990). 1989, Staggs et al. Angling has not been 1990, Hansen et al. cited as a major 1991, U.S. Dep. Int. factor in the extirpa- 1991). tion of any fish in Wisconsin (Becker 1983) Commercial overfishing has been or extinction of any species in North directly implicated in the decline of lake America (Miller et al. 1989). Careful sturgeon, certain cisco species, Great Lakes monitoring of this activity and its impacts, brook trout, and lake trout in Wisconsin however, remains vital. waters, although dam construction, habitat The indirect effects of angling harvest losses, and introduction of exotic species on other species in the aquatic community were undoubtedly also factors in the are less well understood. Nonharvested decline of these species (Becker 1983). species are likely to be affected by changes During this century, regulated sport in predation intensity or food availability angling has been the dominant harvest when game or commercial species are method for most Wisconsin fish. The harvested. Species interactions in north- effects of angling harvest on the most temperate fish communities are complex sought-after species are fairly well under- and often affected by weather or other stood. Non-selective harvest typically unpredictable factors (Colby et al. 1987). increases total mortality and reduces Under these conditions, demonstrating that abundance, but anglers generally select for changes in nonharvested species are caused larger fish. Such size-selective harvest can by harvest of other species is difficult. also reduce the relative abundance of older Changes in relative abundance of fish

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targeted by sport angling or commercial commercially, resulting in a drastic decline harvest can have measurable impacts on in snapping turtle populations (Vogt 1981). the absolute and relative abundance of Some evidence indicates that most of the plankton species, which in turn can have large turtles are gone from the Mississippi impacts on nutrient cycling and water and Lower Wisconsin Rivers (Dan Nedrelo, quality (e.g., Brooks and Dodson 1965, Wis. Dep. Nat. Resour., pers. comm.). Shapiro et al. 1975, Kitchell 1992). Several long-term snapping turtle trappers Commercial harvest of fish is regu- have indicated that some local snapping lated and information on the harvested turtle populations have been nearly elimi- stocks suggests that impacts are similar to nated by harvest pressures. The demand for those caused by sport angling. However, softshell turtle meat (two species) has management programs for commercial increased significantly for European and fisheries on a national and international Japanese markets. Current regulations do scale have more frequently than not failed not offer any protection for these species to prevent collapsed stocks (e.g., Ludwig et and harvest numbers are unknown except al. 1993) and in past years commercial for turtles taken by commercial operations fishing here in Wisconsin has contributed as incidental catch. These records indicate to collapsed and extirpated Great Lakes fish that incidental catch of turtles for the last (Becker 1983), so these fisheries must be three years has been the highest in 40 years managed conservatively (Peterman and (Marron 1994). Bradford 1987, Peterman 1990). Modernization of wild rice harvesting There has also been a significant methods has led to significant overharvest, commercial harvest of small fish, aquatic resulting in lower yield and elimination of insects, and crayfish for bait sale. It is some stands (Bernthal et al. 1992, Vennum generally assumed that impacts on the 1988), although this problem may have harvested species have been minimal been mitigated by current harvest regula- because these species typically have high tion. fecundity or are widely distributed relative to the areas of harvest. However, these LARGE-SCALE CHEMICAL T REATMENTS fisheries are lightly regulated, and very little information exists on the number of Some water bodies which are thought organisms harvested, much less the impact to have undesirable aquatic communities or on the aquatic ecosystems. high numbers of exotics such as carp are There is growing evidence that harvest chemically treated and restocked with a of herptiles is reducing the populations of desired species mix. Chemical treatments some species in the United States (Dundee usually eliminate all fish and many other et al. 1992; James Harding, Mich. State aquatic species in a water body including Univ., pers. comm.). Changes in status of native species and can have at least regional Wisconsin herptile species caused by impacts on biodiversity. Becker (1983), in harvest are not well known. The recent noting the occurrence of a rare bullhead surge in the herptile pet trade has increased minnow population in the upper Fox River, the exploitation of some Wisconsin species stated, “the continued poisoning of por- and even the state-threatened wood turtle is tions of the Fox River and adjacent waters being smuggled as more states add this with antimycin or other fish toxicants, for species to their protection list, creating a the purpose of carp removal, may jeopar- greater demand on protected stocks (James dize or wipe out the only known Great Harding, Mich. State Univ., pers. comm.). Lakes population of the bullhead minnow.” Wood turtles are popular turtles for the pet Chemical treatments of the Rock River trade in the U.S. and abroad. Snapping system in the 1970s may have eliminated turtles have been trapped and hooked for the least darter, a species of special concern many years in Wisconsin. The pools of the from the Maunesha River system (Fago Mississippi have been extensively trapped 1992, Becker 1983).

182 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE The impacts of chemical treatments pond dredging. The impacts of these are probably confined to the waters treated. practices on local biodiversity are unclear. Most treated waters are small, but occasion- For example, Carline and Brynildson ally larger waters such as Beaver Dam Lake, (1977) studied the results of spring pond Dodge County (6,542 acres); Delavan Lake, dredging. Invertebrates usually recolonized Walworth County (2,072 acres); Horicon dredged areas provided some undisturbed Marsh; or the upper Rock River system are habitat was left. Fish populations some- treated. Many treated waters were already times did not change dramatically. Bank perturbed by exotics or hyper-eutrophica- stabilization and other channel modification, and treatments may have improved tion practices were often applied primarily conditions for native species. On a land- in streams with already disturbed commu- scape scale, the relatively small proportion nities with the intent of reestablishing of waters treated make it unlikely that healthy aquatic communities. However, chemical treatments have had a major wood turtles typically nest in areas exposed impact on Wisconsin’s biodiversity. How- by natural erosion processes along streams ever, the lack of a long-term biological (David Evenson, Wis. Dep. Nat. Resour., monitoring program makes it difficult to pers. comm.). determine what effects such treatments On a statewide scale, it is unlikely that have had. these habitat management practices have had a measurable impact on Wisconsin’s DEPARTMENT MANAGEMENT PRIORITIES biodiversity. Cold-water stream habitat improvement structures have been placed The effects of Department manage- on about 300 miles of Wisconsin’s 9,500 ment activities—such as chemical treat- miles of trout stream and 33,000 total miles ments, intensive aquatic habitat alteration, of rivers and streams. The Department has and water level manipulations—on dredged about 60 of the state’s 1,700 spring nongame or other nontarget aquatic species ponds (Carline and Brynildson 1977; Max is sometimes not considered, given a Johnson, Wis. Dep. Nat. Resour., pers. cursory look, or the nongame species are comm.); some of these were already deemed less desirable than game fish impacted by agricultural or timber harvest species that have more sport-fishing value. related sedimentation. Although these choices are not inherently wrong, they should be made with due WATER LEVEL MANIPULATIONS consideration to all components of the aquatic ecosystem, particularly on a re- Artificial water level regulation in gional and landscape scale. The need to Wisconsin’s 1,550 dammed lakes can have approach management from an ecosystem negative impacts on the aquatic system management perspective is becoming particularly when it deviates substantially increasingly evident. Integration of the from natural patterns. Water levels fluctuate appropriate programs is essential if we are widely in some reservoirs, especially those to respond to the needs of the ecosystem as used for flood control and peaking hydro- a whole. power operations (Thuemler et al. 1989). Winter drawdowns are frequently done to HABITAT IMPROVEMENT PROJECTS minimize ice damage to shoreline properties and increase spring runoff storage Intensive habitat management prac- capacity. These fluctuations have direct tices include: placement of artificial habitat impacts when fish or amphibian eggs are structures such as fish cribs; riprapping or stranded and can have indirect impacts other bank stabilization; construction of when changing water levels favor certain channel modification or maintenance species over others. For example, water structures such as the LUNKER structure level fluctuations apparently favor carp and (Vetrano 1988); placement of artificially inhibit reproduction of native northern constructed spawning reefs; and spring

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pike in Petenwell and Castle Rock reser- in two reservoirs on the Peshtigo River voirs (Jim Kreitlow, Wis. Dep. Nat. Resour., were significantly less than at other reser- pers. comm.), and low or fluctuating early voirs on the same river where winter spring water levels disrupt spawning of drawdown had not occurred. Correspond- northern pike and walleyes (e.g., Johnson ingly, turtle populations in these impacted 1961, Johnson 1971, McCarraher and reservoirs were markedly depressed (R. Thomas 1972, Holland and Huston 1984, Hay, Wis. Dep. Nat. Resour., unpubl. data). Kallemeyn 1987). Hibernating turtles are Water levels are often held higher susceptible to freez- during summer to ing and desiccation if facilitate recreational reservoirs are lowered All of the drowned bay mouth estuaries in activities and dock or drained (Dorff Lake Superior and many in Lake Michigan access. Higher 1990). Heath (1992, summer water level are located in Wisconsin. These unique 1993b, 1993c) may increase littoral reported an inverse features occur when the mouths of the habitat for fish relationship between tributary rivers have formed estuaries (Johnson 1971) or the degree of late fall enclosed within sand spits formed by waterfowl, but may and winter draw- along-shore currents, coming and going increase spawning down and turtle with changing water levels and storms. habitat for carp or population densities These estuaries provide critical habitat for affect aquatic macro- in several Wisconsin phytes such as wild several bird species such as the least tern reservoirs. A species rice (Fannucchi et al. that matures very and piping plover, as well as dune thistle, 1986). Artificially slowly, like the dwarf lake iris, beach pea, and grass high water levels in Blanding’s turtle, can of parnassus. large, shallow lakes be significantly can also increase impacted by winter wave damage to drawdowns, especially since they are littoral areas, increase turbidity, and elimi- already threatened by fragmentation and nate macrophytes (e.g., Engel and Nichols the loss of habitat (Dorff 1990). Impacts of 1994). Water levels in large reservoirs are winter drawdowns on turtles could be often systematically drawn down to aug- minimized by conducting drawdowns prior ment summer river flows. Unstable water to October 1st in Wisconsin waters which levels typically result in poor macrophyte will allow turtles to seek alternate hiberna- development with associated loss of fish tion sites (Dorff 1990, Heath 1992, 1993b, and macroinvertebrate habitat, and in 1993c). extreme cases can impair fish spawning In addition, when ice is lowered onto activities. Drawdowns conducted for the the bottom substrate during a winter purpose of establishing emergent vegetation drawdown, substrate freezes to the under- can increase the opportunity for seeds of side of the ice, resulting in scouring and exotic nuisance species such as purple resuspension of sediments (Glenn Miller, loosestrife to germinate and become Great Lakes Indian Fish and Wildl. Comm., established (Merendino et al. 1990). pers. comm.). Organic substrates are both compacted and removed by ice settling on ESTUARY HABITAT MANAGEMENT it. During turtle surveys conducted for FERC relicensing on the Chippewa and All of the drowned bay mouth estuar- Peshtigo rivers, it was noted that organic ies in Lake Superior and many in Lake substrates were almost non-existent or Michigan are located in Wisconsin. These compacted in the shallower bays of reser- unique features occur when the mouths of voirs where winter drawdowns were the tributary rivers have formed estuaries routinely done. Macrophyte plant densities enclosed within sand spits formed by along-shore currents, coming and going

184 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE with changing water levels and storms. waters, was conducted in the 1960s and These estuaries provide critical habitat for 1970s (e.g., Carlson and Andrews 1977). A several bird species such as the least tern randomized survey of limnological charac- and piping plover, as well as dune thistle, teristics of 1,140 lakes was also conducted dwarf lake iris, beach pea, and grass of (Lillie and Mason 1983). The Bureau of parnassus. Endangered Resources maintains a database Many of these estuaries have been of rare aquatic species occurrences, but this degraded by filling and dredging for is not the result of a systematic inventory. disposal of industrial wastes, fly ash, The Bureau of Water Resources Manage- taconite tailings, bark and sawdust, and for ment has also recently initiated a program construction of roads, tank farms, coal to collect limnological and storage areas, docks, grain elevators, and macroinvertebrate information as part of small-boat harbors. Developments and their Basin Plan monitoring program. The alterations are isolating the remaining Department has also conducted many pockets of estuary (e.g., the separation of surveys of state waters that were not part of St. Louis River estuary from the Allouez Bay a statewide or regional program. estuary), and are causing habitat simplifica- Numerous surveys of other aquatic tion. Most of Green Bay’s Atkinson’s Marsh organisms have been conducted. is affected by dredging and filling with fly Wisconsin’s snail populations were sur- and bottom ash, dredge spoil, and other veyed in the 1920s (Baker 1928a). State- wastes behind a bulkhead line. Other wide surveys of mussels were conducted in estuaries such as at Port Wing, Marinette, the 1920s and 1970s (Baker 1928b, and Peshtigo are affected by recreational Mathiak 1979), and mussels in the Missis- development, but undisturbed parts also sippi River were surveyed in the late 1970s remain. Only a few, such as Flag River and (Ecological Analysts 1981, Theil 1981, the Kakagon Sloughs, are relatively undis- Duncan and Thiel 1983). Statewide surveys turbed. The Mink River estuary in Door of crayfish and shrimp were published by County and some parts of the Kakagon Bundy (1882), Creaser (1932), and Hobbs Sloughs have been the focus of protection and Jass (1988). Vogt (1981) provided a efforts of The Nature Conservancy. Backpack ACK OF ONITORING electroshocking L M equipment allows managers to survey The Department has collected a and monitor remote substantial amount of information on the waters. Photo by state’s aquatic ecosystems over the years. Robert Queen. Several statewide surveys of fish distribution have been done (Greene 1935, Becker 1983, Fago 1992). However, these surveys did not provide information on relative abundance of species, and the last survey (in the 1980s) covered only 45% of the state’s waters. Other Department or university fish sampling programs cover few waters or a short period of time. The Department’s ambient lakes monitoring program collects water quality and limnological information from 50 selected waters but has been in existence for only a decade. The Waters Classification Program, a major statewide survey of physical, limnological, and fishery characteristics of the state’s

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summary of collections and descriptions of to grow throughout their lifetimes and Wisconsin’s amphibians and reptiles, which attain a weight of several hundred pounds. have been examined in varying degrees of Past experience with exotic species and completeness since 1883. The Milwaukee predictive management are not accurate or Public Museum maintains a Herpetological reliable indicators of likely outcomes from Atlas Project which is the current repository adaptive management and biological for information from these and later engineering. However, to the extent that an surveys. A volunteer frog and toad monitor- emphasis and reliance on management ing survey has been conducted by the diverges from an emphasis on less manage- Department since 1984 (Mossman and ment with reliance on natural reproduction Hine 1984, Mossman and Hine 1985, and a sustainable resource base, conflicts Mossman and Huff 1990). and controversy will result. Judgments on Unfortunately, this apparent wealth of whether the outcomes of these divergent survey information falls far short of meeting approaches are “good” or “bad” will also the critical need for long-term monitoring depend on the observer’s values about of Wisconsin’s aquatic ecosystems and resource management. statewide systematic inventories of aquatic Beyond these important concerns, organisms. Data from these surveys are there is very little evidence that genetic usually not collected in a standardized bioengineering is a viable management manner and rarely contain relative abun- option. We barely understand the impor- dance information, and surveys are not tance of genetic diversity in native species systematically repeated to track distribution which have adapted to Wisconsin’s waters and abundance trends. Information from over millennia. Thus, costly genetic experi- these surveys is often dispersed among mentation poses the potential for major different programs in a variety of computer disruption of existing aquatic ecosystems. and paper file storage formats making accessibility difficult. There is a critical RECREATION need for long-term trend information on the status of aquatic communities using Recreational use of aquatic ecosystems either bioindicator species or community continues to increase. Activities such as samples. Such trend information will only boating, canoeing, swimming, camping, be obtained by institutionalizing a stan- and hiking along riparian areas are among dardized, statistically valid aquatic ecosys- Wisconsinites’ favored activities (Penaloza tem monitoring program as an integral part 1989, 1991, Wis. Dep. Nat. Resour. 1991). of the Department’s aquatic management Boats and motors are becoming more programs. An important basis for such a numerous and larger (Penaloza 1991). program would be a systematic inventory Along with increased demand for these of important aquatic organisms across the activities comes increasing demand for boat state’s waters. launch ramp, canoe access, beach, marina, campsite, and trail development. Both the recreational use and associated develop- BIOENGINEERING ment impact the aquatic ecosystem. Biological engineering to produce Boating can result in direct impacts to faster growing, larger, and more prolific fish habitat. Outboard motors discharge raw species has the potential to alter manage- fuel, oils, and other combustion byproducts ment goals and objectives from reliance on directly into the water (Jackivicz and existing species and strains to designing Kuzminski 1973, Wall and Wright 1977). new species and strains to meet specific Extensive fish kills in the Fox River have management goals and objectives. For been attributed to carbon monoxide instance, it is theoretically possible to discharge from outboard motors (Jim engineer more appealing bait fish, new Kempinger, Wis. Dep. Nat. Resour., pers. predators, or freshwater fish that continue comm.), though this was an area of excep-

186 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE tionally high use. Heavy boat traffic is known to disturb vegetation and macroinvertebrate production and cause shoreline erosion and sediment resuspension (Lagler et al. 1950, Liddle and Scorgie 1980, Smart et al. 1985). Increased sedimentation is known to have detrimental impacts on fish and other aquatic organisms (e.g., Berkman and Rabeni 1987, Ritchie 1972, Ellis 1936). Heavy boat use is often blamed for turbidity which can also adversely affect aquatic organisms (e.g., Van Oosten 1945, Gardner 1981, Breitburg 1988, Robel 1961, Newcombe and MacDonald 1991, Lloyd et al. 1987, ), but there is mixed evidence for this relationship Recreational boating is (Lagler et al. 1950, Moss 1977, Liddle and PRESENT STATUS a favorite activity in Scorgie 1980, Yousef et al. 1980). Boat Wisconsin. Boats and traffic also disturbs waterfowl and other Quantitative analyses of the present motors are becoming more numerous and aquatic wildlife. (See review by York 1994.) status of aquatic ecosystems in Wisconsin larger, placing Movement of boats among waters can require a statewide database of systemati- increased pressure on transport propagules of exotic species such cally collected biological samples. Such a aquatic systems. as Eurasian water milfoil and zebra mus- Photo by Robert database exists only for fish (Greene 1935, Queen. sels. Becker 1983, Fago 1992). Other taxa Heavy use of riparian areas can result including aquatic macrophytes, phy- in bank erosion, vegetative destruction, toplankton, zooplankton, benthic inverte- littering, and demands for increased brates, crayfish, amphibians, and some camping and access site development (e.g., reptile, bird, and mammal species are Manning 1979). Recent interest in marina clearly dependent on aquatic systems, but construction in the Minnesota-Wisconsin there is far less systematic data on their boundary waters of the Mississippi and St. distribution and abundance. This section Croix rivers will add to congestion on what will use fish distribution data to quantita- is already one of the most crowded and tively assess present aquatic ecosystem congested areas in the country for boating health across the state. (Tom Watkins, Wis. Dep. Nat. Resour., The status of the aquatic communities pers. comm.). There is some evidence that can be successfully indexed by fish com- pollution levels are higher near marinas munities (Karr 1981, Fausch et al. 1990, (Mack and D’Itri 1973). Habitat will also be Lyons 1992). Extensive fish surveys of lost due to dredging and riprapping for Wisconsin waters by Greene (1935), Becker development of facilities to support recreational activities such as marinas, condo- Aquatic macrophytes miniums, boat launches, and parks. are an important While perhaps not of great historical component of aquatic biodiversity. Systematic impact on Wisconsin’s aquatic ecosystems, data on their distribu- the current effects of concentrated boating tion and abundance is and other water recreational activities are needed. Photo by documented. Given the dramatic increase Dorothy Cassoday. in the levels of these activities in recent years, resulting changes in the aquatic communities must be monitored carefully.

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(1983) and Fago (1992) provide a quantita- tissues, indicating contamination may still tive basis for discussion of the current be occurring due to atmospheric deposi- status of Wisconsin’s aquatic communities. tion. Monitoring indicates that contaminant Use of indicators such as presence of fish levels have been declining in Lake Michi- species intolerant to environmental degra- gan fish since the 1970s (e.g., Staggs 1987) dation (Lyons 1992), species richness, though rates of decline have slowed be- history of extirpations, current status of cause of internal cycling and continued threatened species, and status of natural atmospheric deposition (Arimoto 1989). reproduction of top-level predators show Monitoring of Lake Superior fish shows trends in aquatic ecosystem health. lower contaminant levels. The Great Lakes aquatic community GREAT LAKES has also been affected by humans. The Great Lakes fish community is a good The shoreline of parts of the Great example. The effects of lamprey predation, Lakes has been modified by urban, indus- water quality degradation, invasion of trial, and second-home development, exotics, and overfishing have combined to especially in urban areas such as Duluth- radically alter the fishery. Some species of Superior, Green Bay, Milwaukee and along cisco have been extirpated in Lake Michi- the Lake Michigan shore near the Illinois gan. Strains of other species, such as lake state line. Shoreline protection efforts such trout and brook trout, have been eliminated as groins, jetties, cribs, dredging, and or greatly reduced. Introductions and navigation channel entries have interfered invasions of non-native fish species, such as with long shore movement of littoral drift. rainbow and brown trout, Pacific salmon, Erosion and scouring of the down-drift side smelt, and alewife have changed the species of these structures is occurring. Other composition of the fishery. However, the shoreline changes such as riprap, sheet pile introduction of the salmon resulted in the walls, gabions, or concrete retaining walls decline of alewives and a comeback in are used in an attempt to stabilize the native perch, sculpins, and coregonids shoreline. They retard the natural process (Stewart et al. 1981, Eck and Brown 1985, of beach formation and destroy unique Jude and Tesar 1985, Wells and Hatch beach and bank plant communities. 1985). Other species, such as perch, Alteration of water levels and natural undergo periods of intensive harvest. fluctuations has affected dune and coastal Undoubtedly predator/prey relationships marsh systems by interrupting nutrient and have been affected by these changes in the organic matter flushing (The Nature fish community. Conservancy 1994). Changing water level The status of the fish communities in fluctuation cycles is leading to the simplifi- Lake Michigan is best described as dis- cation of coastal marshes by eliminating turbed and unstable (Wells and McLain species that require drawdowns at certain 1973). Reproduction of trout and salmon is times to allow germination (Keddy and negligible, and populations are primarily Reznicek 1986). supported by stocking. Sea lamprey preda- Contaminants, particularly PCBs, are tion, angler harvest (Clark and Huang commonly found in many Lake Michigan 1985), and overstocking (Stewart et al. and Green Bay fish and waterfowl at levels 1981) all affect fish populations. Reproduc- which require consumption advisories. tion and populations of native deepwater Contaminants in the Great Lakes sediments sculpins and bloaters are at recent highs, and waters have been passed along through while non-native alewive populations are the food chain, resulting in contamination extremely variable but are at recent lows of invertebrates, fish, wildlife, and humans. (Jude and Tesar 1985, Wells and Hatch Contaminants remain in the bottom 1985). Populations of lake and round sediments, and fish in remote parts of Lake whitefish seem abundant and relatively Superior have mercury and PCB in their stable (Wells and McLain 1973). Of the

188 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE seven cisco species once found in Lake indicators of the status of the entire aquatic Michigan, only the cisco and bloater remain community in the Great Lakes (Ryder and in numbers sufficient to preserve the Edwards 1985, Marshall et al. 1987). Based population. Three cisco species— on this assumption, Lake Michigan has shortnose, blackfin and deepwater—are been dramatically affected by habitat likely extinct (Becker 1983). Two other simplification— primarily dredging, cisco species—shortjaw and kiyi—while wetland filling and water quality declines in likely extirpated in Lake Michigan are still estuarine areas, introduction of exotic relatively abundant in Lake Superior. species, excessive harvest of commercially Excessive commercial harvest and competi- desirable top predators, and pollution. tion from alewives are cited as primary The uncertain status of lake sturgeon causes of cisco population declines. reproduction indicates fragmentation of the Among Lake Michigan warm-water lake ecosystem, as lake populations were species, reproduction and populations of cut off from historical spawning areas by northern pike are currently limited. Wall- dam construction. Lake Michigan, however, eye and yellow perch reproduction is now shows some signs of recovery: declining significant in Green Bay, but populations numbers of exotic species, improving are still experiencing large fluctuations. reproduction of some native species, and Yellow perch are well-established in other declining contaminant burdens (e.g., Wells areas such as Milwaukee harbor, but and McLain 1973). walleye populations are negligible. Lake Conversely, Lake Superior shows few sturgeon populations are probably low but signs of either habitat simplification or the current level of reproduction of this fragmentation. The aquatic community has long-lived species is unknown. Reproduc- primarily been affected by human manage- tion of other warm-water species appears ment activities, including excessive harvest adequate to maintain the stocks. of commercially desirable species, stocking In Lake Superior, fish communities, of domesticated strains of lake trout, and although heavily exploited, are more stable introduction of exotic species (Lawrie and (e.g., Lawrie and Rahrer 1973). There is Rahrer 1973). Localized habitat degrada- significant natural reproduction of most tion in the urban areas of Duluth-Superior trout and salmon species, but angler and may also be occurring but does not appear commercial harvest and sea lamprey to be affecting Lake Superior biodiversity predation have kept adult populations at a on a lake-wide scale. relatively modest level. Reproduction and populations of other cold-water species is INLAND LAKES adequate, but overall productivity in Lake Superior is low so populations are often Fish species intolerant of poor water modest by Lake Michigan standards and quality and environmental degradation cannot support as large a predator popula- (Lyons 1992) such as smallmouth bass, tion. Only four cisco species—cisco, rock bass, Iowa darter, blacknose shiner, bloater, shortjaw, and kiyi—were originally and spottail shiner were found at 56% of present in Lake Superior and all are still the 1,644 sampled lake stations (Table 19). present in sufficient numbers to maintain Species richness averaged 7.2 species per populations. Lake Superior is also home to station with a range of one to 23 species the only known population of pygmy (Table 20). whitefish east of the Rockies. It is abundant There are currently no federally and relatively stable (Becker 1983). With threatened or endangered fish species in some local exceptions, reproduction and Wisconsin lakes, although six species are populations of Lake Superior warm-water under consideration for federal listing. No species are adequate to maintain the stocks. known extinct species were endemic to Fish communities—specifically the Wisconsin lakes (Becker 1983). Several abundance of lake trout—can be used as species are thought to have been extirpated

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Table 19 Stations Percent Stations Percent Percent of stations with tolerant and with with with with intolerant fish species Total Intolerant Intolerant Tolerant Tolerant in Wisconsin lakes and Ecoregion Stations Species* Species* Species* Species* rivers, classified by ecoregion as defined Lakes by Omernik and Gallant (1988), based Driftless Area (DRFT)2211501777 on nearest county boundary. (Data N. C. Hardwood Forest (NCHF) 355 203 57 266 75 source: Wisconsin N. Lakes and Forest (NOLF) 660 424 64 511 77 Fish Distribution Study Master Fish File.) S.E. Wis. Till Plains (SETP) 607 276 45 471 78 Statewide 1,644 914 56 1,265 77 Streams Driftless Area (DRFT) 1,586 886 56 1,466 92 N. C. Hardwood Forest (NCHF) 1,079 850 79 977 91 N. Lakes and Forest (NOLF) 1,317 1,029 78 1,149 87 S.E. Wis. Till Plains (SETP) 1,433 662 46 1,376 96 Statewide 5,415 3,427 63 4,968 92 Rivers Driftless Area (DRFT) 331 289 87 245 74 N. C. Hardwood Forest (NCHF)7546615573 N. Lakes and Forest (NOLF)6859873957 S.E. Wis. Till Plains (SETP)2619732077 Table 20 Statewide 500 413 83 359 72 Analysis of fish *Tolerance and intolerance to environmental degradation as defined by Lyons (1992). species richness in Wisconsin lakes and rivers, classified by from Wisconsin waters, including ghost in Wisconsin and were never common in ecoregion as defined shiner, ironcolor shiner, and creek lakes. by Omernik and Gallant (1988), based chubsucker, but these were probably not Differences in fish communities on nearest county common in lakes. Wisconsin lists nine among ecoregions suggest that biodiversity boundary. (Data species of endangered fish and 11 fish in SETP lakes has been more heavily source: Wisconsin Fish Distribution Study species as threatened, although most of influenced by human activities compared Master Fish File.) these species are on the edge of their range with NOLF and NCHF lakes. Comparisons

No. Fish Species, by Water Type

Lake River Stream All

Ecoregion Mean STD Mean STD Mean STD Mean STD

Driftless Area (DRFT) 8.00 4.15 15.19 7.36 9.08 5.43 10.11 6.23 N. C. Hardwood Forest (NCHF) 7.41 4.02 11.53 7.99 10.35 6.30 9.72 6.08 Lakes and Forest (NOLF) 7.02 3.52 11.99 5.92 8.10 4.59 7.88 4.42 S. E. Wis. Till Plains (SETP) 7.19 4.29 11.42 8.76 9.88 5.76 9.11 5.56 All 7.18 3.94 14.01 7.52 9.31 5.58 9.16 5.64

190 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE with DRFT lakes are difficult because few Piscivorous birds have been impacted lakes exist or were sampled in that by water quality problems in NOLF lakes. ecoregion. Species regarded as intolerant of Common loons rarely nest on lakes with environmental degradation were present at low pH and elevated mercury levels and only 45% of sampled SETP stations (Table suffer higher reproductive mortality in 19). Five state-threatened and one state- these locations (Meyer 1994). endangered species were found in SETP With regard to aquatic vegetation, lakes including pugnose shiner, redfin wild rice areas in northern Wisconsin have shiner, river redhorse, starhead topminnow, been lost to flooding caused by dams striped shiner, and longear sunfish. (Vennum 1988). In contrast, fish communities in NOLF and NCHF lakes showed less STREAMS evidence of biodiversity impacts. Species intolerant of environmental degradation No federally threatened or endangered were present at 64% of sampled NOLF fish species are found in Wisconsin stations and 57% of NCHF stations. streams, but six species are currently under Tolerant species such as carp and green consideration for federal listing. At least sunfish are generally uncommon (see Table three species, ghost shiner, ironcolor shiner 13). Intolerant species such as the small- and creek chubsucker, have been extirpated mouth bass, rock bass, and Iowa darter are from state streams. The nine state-endan- more common. Only the state threatened gered and 11 state-threatened species are pugnose shiner, redfin shiner, Ozark found in state streams. Most are on the minnow, and longear sunfish were found in edge of their distribution, but species such any NOLF or NCHF lakes, and it is un- as the river redhorse, pallid shiner, crystal NOLF Northern likely that any species were extirpated from darter, and gilt darter are declining across Lakes and Forest lakes in these regions. One species thought their ranges. Wisconsin has some of the best populations of greater redhorse and to be extirpated from SETP waters (Becker NCHF North 1983), the black redhorse, was recently pugnose shiners across their ranges (Lee et Central found in a NCHF reservoir (Fago and al. 1980) even though they are listed as Hardwood Forest Hauber 1993). threatened in this state’s streams. Species intolerant of environmental The current status of NOLF and DRFT Driftless NCHF lakes as indexed by fish communi- degradation, such as brook trout, small- Area ties is healthy and stable. Reproduction and mouth bass, and rock bass (Lyons 1992), abundance of top level predators is gener- were found at 63% of the 5,415 stations SETP Southeast sampled statewide (Table 19). Species Wisconsin ally adequate (Staggs et al. 1990, U.S. Dep. Till Plains Int. 1991), and a large proportion of richness is higher in streams than lakes, sampled stations have species that are averaging 9.3 statewide and with a range of intolerant to environmental degradation. one to 40 species per station (Table 20). Species richness is less than that of south- In comparison with NOLF and NLHF ern Wisconsin lakes, but waters in more streams, patterns of fish distribution northerly latitudes are often species poor to suggest that some streams in the SETP and begin with (Lyons 1992). DRFT ecoregions have diminished biologi- Localized impacts such as dam cal integrity. Intolerant species were found construction in NCHF lakes are not readily at only 46% and 56%, respectively, of apparent in this regional analysis but are sampled stations. While many species shifts known to be important in specific waters. are due to underlying habitat differences Water level fluctuations and high nutrient such as larger streams and warmer tem- loading are thought to have resulted in peratures, the increased abundance of poor water quality and high carp popula- tolerant species such as carp, green sunfish, tions in the region’s two largest reservoirs, bluntnose and fathead minnows, and Castle Rock and Petenwell (Jim Kreitlow, yellow bullhead provide evidence for Wis. Dep. Nat. Resour., pers. comm.). environmental perturbation (Table 15).

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Other studies suggest that smallmouth bass damming and paper mill pollution on the populations in DRFT streams have experi- Wisconsin and Chippewa rivers. enced major declines during the last three Several state-threatened fish species decades (Forbes 1985, Mason et al. 1991). are common in NOLF, NCHF, and DRFT Two state-endangered and eight state- rivers. Only three state-threatened spe- threatened fish species were found at SETP cies—gilt darter, river redhorse, and blue stations; one state-endangered and nine sucker—have been found in NOLF rivers, state-threatened fish species were found at but the gilt darter and river redhorse were DFRT stations; no state-threatened fish found at one third of the sampled stations species were found at NCHF stations; and suggesting they are not threatened in the three state-threatened fish species were NOLF ecoregion. Seven state-threatened found at NOLF stations. fish species were found in NCHF rivers. DRFT rivers have nine state-threatened and LARGE RIVERS one state-endangered fish species. The four state-threatened and one state-endangered Species regarded as intolerant to fish species found in SETP rivers but are environmental degradation (Lyons 1992) not common in that region’s rivers. occurred at 83% of sampled stations. Rivers NOLF rivers contain the lowest exhibit the highest species richness of all percentage of intolerant species among the the aquatic communities, averaging 14 four ecoregions suggesting there has been species with a range of one to 40 species environmental degradation in the per station. ecoregion. However, there is evidence that No federally threatened or endangered NOLF fish communities are still relatively fish species are found in Wisconsin rivers, healthy. River redhorse, a state-threatened but six species are currently under consid- species known to be affected by dam eration for federal listing. At least three construction, was present at 30% of species, ghost shiner, ironcolor shiner and sampled stations. Also, the gilt darter, a creek chubsucker, have been extirpated state-threatened species intolerant of from state streams and rivers. The nine environmental degradation, was found at state-endangered fish species and many of 32% of sampled stations. Most top predator the 11 state-threatened fish species cur- species exhibit self-sustaining populations. rently inhabit state rivers. Most are on the These analyses show that some edge of their distribution, but species such stations have degraded fish communities, as the paddlefish, blue sucker, river red- while fish communities at many stations horse, pallid shiner, crystal darter, and gilt remain intact. This finding is consistent darter are declining across their ranges. with the observation that dam construction Wisconsin has some of the best populations is a major environmental impact on of greater redhorse across their ranges (Lee Wisconsin’s rivers since the most significant et al. 1980) even though they are listed as effects of a dam would be localized in areas threatened in this state’s rivers. near the dam. Differences in river fish communities between ecoregions were not pronounced. Average species richness is higher in the PROJECTED STATUS DRFT ecoregion, but this is probably because the largest rivers, the Mississippi Some taxa dependent on aquatic River and the lower Wisconsin River, are systems in Wisconsin face a difficult future. located here. Species intolerant to environ- The abundance and zoogeographical mental degradation are found at a large distribution of Wisconsin’s river mussels percentage of sampling sites in all have been dramatically altered. Three ecoregions (Table 17). Intolerant species species have been extirpated (scaleshell, fat were found at only 61% of NCHF river pocketbook and pyramid pigtoe), two have stations perhaps reflecting impacts of only remnant populations (ebony shell and

192 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE elephant ear), and two are on the federal endangered species list (winged mapleleaf, higgins eye). If the invading zebra mussel affects native mussel species as severely as predicted, up to half of the states stream and river mussel species will be threatened or endangered. Other taxa are in better condition. Although the abundance of many fish species has been greatly altered, very few fish species have been extirpated in Wis- consin. Of the fish species locally extirpated, some have later been found again, although in very low numbers (e.g., black redhorse and skipjack herring), and most extirpations involved species on the edge of their range. There are no federally endan- develop pollution prevention technology. The abundance and gered or threatened fish species in Wiscon- distribution of Many priority watershed plans already Wisconsin’s river sin, although the lake sturgeon, blue sucker recommend removal of dams to improve mussels have been and paddlefish are among the candidates water quality. Dam removal may be an dramatically altered. for listing. It appears most extirpations effective option for restoration of riverine The winged mapleleaf, have been caused by habitat destruction, shown here, is on the ecosystems. state and federal primarily associated with agriculture and The most cost-effective management endangered species dams. In general, most native fish species strategy is protection of existing healthy, lists. Photo by William Smith. are self-sustaining, especially in Lake self-sustaining aquatic ecosystems. How- Superior and the northern ecoregions. Even ever, pressure remains to develop and the heavily developed agricultural areas and destroy riparian habitat, modify land use densely populated areas of southeastern patterns in watersheds, intensify agricul- Wisconsin still support self-sustaining fish ture, divert water for irrigation and indus- populations and good species diversity in trial uses, and provide more harvest oppor- some waters. tunities for sport The relative anglers and subsis- abundance of The most cost-effective management tence users. Govern- currently healthy strategy is protection of existing healthy, ment agencies and aquatic communities self-sustaining aquatic ecosystems. users will have to in many inland establish and main- waters is an excellent tain strong partner- indicator for the future. Attention can be ships if long-term sustainability of aquatic focused on identification and restoration of ecosystems is to be maintained. specific degraded habitats and on protect- The accidental or intentional intro- ing and restoring species whose numbers duction of exotics has already been impli- are in local decline. Existing aquatic cated in major changes in native communities provide a source for biodiversity. Past invaders, including carp, recolonization of native species and a Eurasian milfoil, and purple loosestrife are model for restoration efforts. Past restora- notoriously hard to control and eradicate. tion successes, such as those on the Wis- The state has invested heavily in control, consin and lower Fox Rivers, provide clear education, and monitoring, but efforts to evidence that such efforts will work. eliminate targeted exotics have shown poor Further progress and additional successes results. Undoubtedly, new exotic species can be expected as the Department works will continue to be introduced, and the cooperatively with industry to install the potential threats to the aquatic community latest pollution abatement equipment and will increase.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 193 AQUATIC COMMUNITIES

recreational fisheries are now thought to be approaching overexploitation, although this concern is not well documented in Wiscon- sin. The popularity of fishing has come at a price. The state’s fishing public, which is among the five largest in the nation (U.S. Dep. Int./U.S. Dep. Comm. 1993), demands an intensive fishery management effort. Resort owners want more large predators such as walleye, muskellunge, and northern pike, while trout anglers want more trout. More access is required to meet the demands of more people who own more boats with bigger motors and who purchase more licenses and pay higher fees. Healthy, diverse Two other possible threats to the More demands require stretching the aquatic communities resources of the aquatic community to are present in many aquatic community present unknown inland waters. Photo by dangers—the potential effects of climate provide for more return, much as a farmer Dean Tvedt. change and the effects of acid deposition may try to increase yield from a corn field. (Bergquist 1991). Assessing the likelihood The short-term expectations of resort of either of these threats is dependent on owners and other interests lead to increased the development of accurate predictive pressure for management actions, such as models. Global climate change could fish-stocking, habitat manipulation, and change the numbers and distribution of single species management, rather than aquatic species in Wisconsin. However, the practiced restraint and reliance on natural climate change threat is only in the early recruitment to replenish a fishery. As the stages of monitoring and model develop- pressure for providing short-term solutions ment (U.S. Dep. Energy 1990). increases, the interest in the long-term health and diversity of the aquatic community could diminish. SOCIO-ECONOMIC ISSUES There is heavy economic pressure to continue developing shoreline property. The aquatic community has a faithful Lake homes continue to be in high de- and dedicated following of avid anglers, mand. Some counties, such as Vilas and recreationists, and other user groups. Oneida, are growing rapidly due to the Conflicts can occur as more people try to demand of retirees for lake and waterfront use a limited resource for purposes that are homes. Former resorts are being converted not easily compatible, such as water-skiing to condominiums. This demand now and fishing. In some cases, there is a strong threatens the smaller, more isolated, feeling of “my lake,” “my river,” “my trout shallow lakes that were not developed stream,” and “my spot,” particularly among earlier because they were “less desirable.” local residents and riparian owners, which The agriculture industry is an ex- may place them in conflict with other tremely important component of statewide users. Wisconsin’s economy, and its impacts on Recreational fishing has a significant aquatic ecosystems in Wisconsin are well impact on aquatic communities. In Wiscon- documented. The Priority Watershed sin, fishing is the sixth most popular program along with stream bank protection outdoor activity among all adults (26% of acquisitions and easements under the all adults) (Wis. Dep. Nat. Resour. 1991). Stewardship Program are the main manage- In some areas of the country, fishing is now ment activities targeted at reducing this so intense that previously underexploited

194 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE large source of pollutants and erosion, but both are largely voluntary programs. POTENTIAL FOR COMMUNITY An evaluation of the socio-economic RESTORATION implications of managing Wisconsin’s aquatic ecosystems reveals many of the Aquatic systems can probably recover same problems that mark similar analyses more quickly than terrestrial systems from of natural resource issues. Documentation the impacts of fragmentation or simplifica- of the value of development and industry tion if the causes are corrected. In rivers needs is straightforward and readily avail- and streams, sediments and contaminants able. Hydropower will generate electricity are flushed downstream, hydrologic worth a certain amount. Lakeshore devel- processes will restore channel morphology, opment yields a certain amount of property riparian areas will revegetate, and aquatic taxes for local government. Industries using organisms will return provided there are no aquatic resources employ a certain number barriers to recolonization from unaffected of people and contribute a certain amount areas (Detenbeck et al. 1992). of tax revenues to local governments. Macroinvertebrates, for example, generally Family farms employ a certain number of recover very quickly (months to just a few people and generate a certain level of years) from most kinds of disturbances expenditures in the rural communities. (Niemi et al. 1990) and colonize new However, documentation of the value of the habitats very rapidly (e.g., Williams and aquatic resource Hynes 1977, Doeg et degraded or lost is al. 1989). Narf less certain and often A major challenge to maintaining the long- (1985) found aquatic involves hard-to- term sustainability of Wisconsin’s aquatic insect colonization of quantify intrinsic ecosystems will be to develop adequate available habitat in a values. What is the valuations of the importance and uses of relocated stream segment in northern value of the nongame these ecosystems, and to ensure that Wisconsin was species killed by a society and future generations are not pollution discharge? complete after 5.5 paying for short-term benefits that limit What is the value to years. Many other the local economy future options. aquatic organisms when tourism de- are mobile and clines because of poor fecund, allowing fishing, increased pollution, or loss of rapid recolonization and repopulation of scenic beauty? What is it worth to be sure affected areas. future generations will be able to enjoy Restoration of lakes may take longer clean water, good fishing, and scenic or require more directed management recreational areas? A major challenge to actions. Lakes that have been exposed to maintaining the long-term sustainability of contamination or excessive nutrient loading Wisconsin’s aquatic ecosystems will be to usually take longer to recover than rivers develop adequate valuations of the impor- and streams because flushing rates are tance and uses of these ecosystems, and to much longer and nutrients and contami- ensure that society and future generations nants are continually recycled from sedi- are not paying for short-term benefits that ments. Morphology in lakes is not shaped limit future options. However, according to by strong currents, so restoration of altered Clark (1991), “the political realities are that habitat may take extremely long periods. exploiters of large resource stocks have every Revegetation of riparian areas would be incentive to impose major external costs on similar to moving water systems, but the public at large, and these externalized replacement of woody debris habitat would costs add up to nonsustainability.” be slower since it is not being actively transported from upstream areas. Lakes also tend to be more isolated from other

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A basic barrier to both restoration and maintenance of sustainable aquatic ecosystems is the lack of meaningful ability to regulate habitat destruction. The Depart- ment has only minimal authority to regulate environmental impacts from agriculture. On paper the Department should be able to minimize destruction of riparian habitat areas, but in practice such destruction continues. Land use practices in nonriparian areas of a watershed can have major impacts on downstream aquatic systems, but zoning and other land use regulations rarely consider aquatic impacts. The successes of point-source water pollution regulation should be a model for Remote lakes such as lakes, which would slow recolonization by regulation of nonpoint pollution and Gobler Lake in Oneida County offer valuable aquatic organisms. riparian development. The substantial insight into restoring There are few if any undisturbed investments made by both governmental aquatic communities. aquatic ecosystems in Wisconsin to use as agencies and state industries in water This lake is located templates for restoration efforts. However, within a State Natural quality improvements are showing results Area. Photo by William there are many systems that, while dis- and should serve as a model for other Tans. turbed, still maintain a healthy complement restoration efforts. For example, water of native species. quality in the Wis- These systems must consin and Fox rivers serve as models for There are few if any undisturbed aquatic has been considerably restoration and as ecosystems in Wisconsin to use as improved and the sources of genetic templates for restoration efforts. However, aquatic resource, stock for there are many systems that, while most notably fish recolonization efforts. species, has improved disturbed, still maintain a healthy When restoration of as well. pre-settlement complement of native species. These However, many aquatic ecosystems is systems must serve as models for waters in Wisconsin a desirable goal, restoration and as sources of genetic still need attention. studies of undis- stock for recolonization efforts. The atmospheric turbed systems in transport of pollut- other regions or use ants across the of paleolimnology techniques may be continent and throughout the region results needed to establish realistic goals. Deter- in the deposition of combustion mining restoration objectives will not byproducts, sulfur and nitrous oxides, always be straightforward. Historically and mercury, and PCB’s from industries in other currently, recreational and commercial states or nations to Wisconsin’s waters. The fishing demands guide management efforts results, such as acid deposition, have been and some components of the aquatic implicated in raising the level of mercury community may consequently be consid- contamination in fish from waters that are ered less important in restoration projects. not exposed to other sources of pollution Careful consideration of the costs of (Lathrop et al. 1990). Contaminant adviso- different management activities and a ries still remain for some species of fish, balancing of management objectives across such as carp and white bass. In addition, various scales must be part of any restora- the allowable residue in fish flesh continues tion plan. to be lowered, reflecting the improving technology of contaminant detection and

196 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE the interests of Americans in maintaining a Concerns about the effect of climate healthful lifestyle. Despite some declines in change on the aquatic community are the levels of PCB’s in Great Lakes fish starting to surface. Regier et al. (1990) (Staggs 1987), the problem of toxics from describe three levels of connections be- some materials, such as dioxins and PCB’s, tween climate change and fish. First, there will continue for a long time due to their is a direct connection between local climate persistence and cycling in the environment, and local assemblages of fish. Second, there which may be on the order of several are indirect linkages between climate, hundred years. Some in-place pollutants hydrology, and the biotic system. Finally, have been covered by recent deposition of there is the human and cultural response to cleaner sediments, but these overlaying these changes. layers can be removed during dredging or Weather extremes of the past decade scouring during floods. Additional sources and the possibility of global climate change of pollutants may occur as a result of make it difficult to predict any resulting groundwater inputs from the thousands of changes in the aquatic community. The past abandoned landfills and leaking under- decade has been marked by unprecedented ground storage tanks. droughts preceded by periods of extreme State industries have already made wetness. The cold-water resources of the substantial investments in pollution abate- northeastern and southwestern parts of the ment equipment, which has reduced the state have been affected by past droughts, level of pollutants being added to aquatic and the Great Lakes littoral areas and systems. However, remedial actions to clean shorelines have been affected by high water up past pollution will be technically and resulting shoreline erosion and wave difficult and very costly. There is increasing damage. The recent variations in climate recognition among government agencies fall within the bounds of predictions from and state industries that development of the atmospheric general circulation model pollution prevention technology is a more that predicts climate change. cost-effective option than paying for later Effects of climate change could clean-up of pollutants. include decreases in winter ice cover on the There are few selective controls for Great Lakes (Sanderson 1987); the devel- exotic species and none are cost-effective opment of a permanent thermocline in the on a large scale. One method to control deeper parts of Lake Michigan overlain by a exotic fish in small waters is to eradicate seasonal thermocline such as occurs in the entire fish community and start over. most of the world’s oceans (McCormick More often than not, the inconvenience of 1990); hypolimnetic anoxia (Schertzer and an exotic fish species is tolerated because of Sawchuk 1990); increased bacterial activity the costs, controversy, and difficulty of in the hypolimnion and sediments completely removing it along with most of (Blumberg and Di Toro 1990); changes in the rest of the aquatic community. Main- habitat for Great Lakes cold-, cool-, and taining a large predator biomass may be warm-water fish (Magnuson et al. 1990); helpful over a long period (Stewart et al. changes in fish growth rates (Hill and 1981), but extirpation of local forage fish Magnuson 1990); reduced stream habitat species or shifts in species composition of for brook trout (Meisner 1990); expansion zooplankton and phytoplankton may of the range of the exotic white perch in the occur. Herbicides can be applied to small Great Lakes (Johnson and Evans 1990); infestations of certain plants, but large scale extension of the northernmost ranges of control is expensive and damaging to yellow perch and smallmouth bass (Shuter related native species. No control methods and Post 1990); possible local extinctions exist for exotic bivalves and zooplankton. of southern fish populations and northward There is evidence that healthy native aquatic invasion of southern fish populations (Tonn ecosystems are more resistant to invasions of 1990); and the invasion of species adapted exotics (Baltz and Moyle 1993). to warm conditions concurrent with local

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extinctions of some cold-water species 2. Use a landscape scale approach to set (Mandrak 1989). watershed or ecoregion-based goals for Most of the consequences of today’s the protection and management of lake aquatic habitat problems are the result of systems. This will involve work with changes brought about by agriculture, many partners, public and private, to forestry, and urban development practices apply the ecological, socio-economic, several decades ago. Some positive trends and institutional aspects of ecosystem are now on the horizon, such as the Con- management to a comprehensive view of servation Reserve Program provisions of the our lake resources and the conservation 1985 Food Security Act. In 1988, Wiscon- of statewide biological diversity. sin adopted Water Quality Standards for a broad range of contaminants. The 1990 3. Recognize the importance of protecting Farm Bill strengthened the requirements for certain unique types of aquatic commu- environmental protection and attempted to nities (e.g., historic wild rice stands), lessen the water quality impacts of agricul- undisturbed aquatic communities (e.g., ture (Pajak 1991). However, later versions wilderness lakes), and long-lived native could change these gains. Additionally, the species (e.g., 100 year-old-ebony shell Department’s Stewardship and Forestry Best mussels, 75-year-old snapping turtles, Management Practice Programs have a and 120-year-old lake sturgeons). These substantial component devoted to water are often are economically valuable, add resources protection. Increased interest in stability to ecosystems, are a reservoir wetland protection and sustainable agricul- for genetic diversity, and have tremen- ture may lead to lower chemical inputs and dous scientific value for understanding less erosion, while changes in manufactur- the processes that affect managed and ing methods and waste treatment portend harvested systems. The concept of old possible decreases in wastewater dis- growth, usually applied to forest com- charges. munities, may help us manage and value populations of long-lived aquatic species and species assemblages. POSSIBLE ACTIONS 4. Manage rivers as ecological continuums The following possible actions are from headwater to mouth, taking into consistent with ecosystem management, account adjacent floodplain and terres- but require more analysis and discussion. trial habitats. Recognize the role of How priorities are set within this list will be floods in maintaining the integrity of based on ecoregion goals, staff workload, river ecosystems. To do this, we will fiscal resources, public input and support, work with many public and private and legal authority. We will work with our partners to develop ecoregion or water- customers and clients to set priorities and shed goals and objectives based on bring recommendations to the Natural ecosystem management principles. This Resources Board for consideration begin- will include reaching a consensus on the ning in the 1995-97 biennium. desired outcomes after considering a full 1. Apply the principles of ecosystem range of management opportunities management to the many kinds of (e.g., nonpoint source control, recre- aquatic communities and their associ- ational use, industrial activity, aquatic ated species. Put less emphasis on single community restoration, and enhance- species management and more on ment of fisheries and aquatic life). communities and ecosystems. One benefit will be the cost-effectiveness of a. Emphasize the protection of the last managing for native, naturally reproduc- large river systems without dams. ing species assemblages and up-front These include the Lower Chippewa protection of habitat.

198 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE River, Lower Black River, and the Namekagon/St. Croix Rivers.

b. Where appropriate, identify opportunities for upstream and downstream ﬁsh passage at existing and proposed dams.

c. Where flows are adversely affected by dams, seek to establish adequate minimum flows to protect recreation, water quality, and fish and aquatic life. Prepare drought contingency plans for rivers where there are consumptive uses or conflicting uses.

d. Document the cost and beneﬁt of dam removal for selected restoration projects, and use the analysis within the ecosystem management decision model to recommend appropriate action.

e. Encourage the preparation of hydroelectric ﬂow models based on entire river systems.

f. Examine the practice of removing natural woody debris from stream beds for channel maintenance. If needed, prepare guidelines to protect instream habitat structure. 6. Develop programs, regulations, and Aerial photographs and satellite imagery 5. Manage riparian and shoreline forests guidelines that effectively protect are among the tools using ecosystem management principles. riparian and shoreline habitats. This will that help managers Allow ﬂoodplains to develop mature include work with local governments take a landscape scale and private groups to develop a com- view of aquatic forests to minimize the impacts of systems. mon understanding of the impacts and ﬂooding and to maintain channel [Top] Photo by geomorphology. Use Wisconsin’s For- long-term costs of poorly planned National Aerial estry Best Management Practices guide- riparian and shoreline development and Photography Program. lines, which require a buffer area of at to provide the support needed to design [Bottom] Photo by Univ. of Wisconsin- least 100 feet along shorelines, to plan and implement long-range plans that Madison Environmen- timber harvest. These buffers are sources provide adequate protection. We will tal Remote Sensing of fallen woody debris to maintain need to promote a combination of Center instream habitat structure, and they approaches that include: provide shade to control stream temperature. They also protect banks and zoning practices, such as those that ground vegetation from damage caused protect sensitive areas from overuse, by heavy equipment. disturbance, or destruction (e.g., special designations for spawning areas or undisturbed natural communities);

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alternative methods of protection 8. Continue to develop a long-term inven- (e.g., new technologies for erosion tory and monitoring program that control and shoreline protection, includes the state’s aquatic communities incentives for property owners to and the species dependent on aquatic protect habitats); and systems. No agency can hope to sample all aquatic taxa in all state waters; an traditional policies and regulations effective approach is to sample a statisti- (e.g., enforcement, legislation, and cally valid subset of waters using cost- grants). effective biocriteria (e.g., indices of biotic integrity or abundance of environ- 7. Identify and restore degraded aquatic mentally sensitive species) and apply communities, working in partnership results to a well-developed waters with other public and private groups to classification system. Such a program ensure success of these projects. Re- will develop meaningful trend informa- moval of the Woolen Mills dam and tion and help identify problem areas in restoration of river and riparian habitat need of special protection or restoration on the Milwaukee River provides an efforts. excellent template for similar projects and demonstrates that there is wide 9. Consider the long-term, cumulative support for such activities, particularly impacts of Department actions. Indi- in urban areas. Some northern rivers vidual regulations or decisions may affected by historical log drives are seem independent of one another, but in among the candidates for restoration. In combination some may be inconsistent some projects, dredging and disposal of or have unintended impacts on the contaminated sediments may be neces- efforts of other programs or on aquatic sary, and the lack of an approved resources. For example, any single hazardous waste site in Wisconsin may macrophyte removal permit may seem present problems. There is great poten- minor, but the cumulative impact of tial for additional joint restoration many such permits may have significant projects involving state, federal, county, effects on the ecosystem. municipal, industrial, and citizen partners. For example, the Great Lakes a. Develop policy to establish an Indian Fish and Wildlife Commission is integrated, comprehensive approach interested in restoring wild rice beds. to aquatic plant management. Three Other organizations, such as Ducks different programs are now involved Unlimited, are interested in shallow lake in aquatic plant management, and restoration. their decisions are often based on different considerations. The Water Regulation program evaluates permits Dragonflies, such as for structures such as sand blankets this nymph of the and mechanical weed control devices, extra-striped snaketail, are important indicator the Lake Management program is species for ecosystem responsible for permitting chemical health. Photo by and mechanical aquatic plant man- William Smith. agement proposals, and Fisheries Management is involved in the habitat issues related to both programs.

b. Consider a pilot program for applying ecosystem management principles to selected aquatic regulatory

200 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE and management programs. This 13. Take action at the state and federal would explore ways to correct levels to prevent the invasions of problems arising when one program exotic species. Contingency plans makes decisions that impact parts of would prepare the Department to be the ecosystem managed by another proactive when small infestations program. This kind of approach occur. Public education and aware- might be easier to pilot for aquatic ness programs can help minimize the than terrestrial systems, because risk of importation and introduction. aquatic community boundaries are more easily defined, and they are 14. Exercise extreme caution in imple- typically already under Department menting biological engineering to management authority. intensively manage the aquatic community. It is doubtful that such 10. Work with the agricultural commu- technology will be cost-effective or nity and other public and private desirable when compared with the interests to address the effects of benefits of protecting naturally agriculture on aquatic ecosystems. reproducing populations within self- Much is known about the impacts of sustaining ecosystems. erosion, nutrients, pesticides, and land use changes. Voluntary pro- 15. Support and conduct additional grams have not always been success- research to apply the principles of ful in mitigating these impacts but conservation biology to the manage- mandatory programs have not been ment of aquatic communities and popular. Incentives to alleviate the ecosystems. Many important ques- environmental effects of agricultural tions remain unanswered, for in- practices need greater attention in stance: do rivers that have been federal farm legislation and pro- fragmented by dams follow the grams. principles of island biogeography? How does a lake’s size affect its 11. Emphasize critical aquatic habitat susceptibility to various kinds of protection and restoration priorities disturbance? Is fish stocking of in land acquisition and easement smaller waters more detrimental to programs. Undeveloped shoreline biodiversity than stocking of larger areas deserve special consideration waters? because these opportunities are rapidly declining.

12. Study the genetic composition of selected native ﬁsh species and modify ﬁsh stocking, transfer, and bait collecting policies if they appear detrimental to genetic diversity.

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Case Study

HABITAT RESTORATION FOLLOWING DAM REMOVAL ON THE MILWAUKEE RIVER AT WEST BEND Contributed by Mike Staggs, John Lyons, and Kris Visser

There are over 50 dams in the Mil- or replace the dam in conjunction with the waukee River Basin, most holding back construction of a new bridge. small impoundments of 50 acres or less. A DNR team studied the 67-acre Because most of these impoundments impoundment and found siltation; poor originated as mill dams, they are located in water quality; high turbidity; low recre- the heart of urban areas and are valued by ational values due to shallow depth; a fish local residents for ice skating, waterfowl population heavily dominated by carp, viewing, and their aesthetic qualities. suckers, and bluntnose minnows; and a Ecologically, however, these impoundments lack of aquatic vegetation throughout the fragment fish habitat, create barriers to fish entire impoundment. Both upstream and movement, may create thermal pollution downstream from the impoundment, where problems, and typically have poor water the river still flowed freely within its banks, quality as a result of sedimentation and the fish population was dominated by a related eutrophication. In some cases, the variety of minnows, darters, crappie, dams are more than a century old, creating bluegills and other panfish, and small- safety concerns for their public and private mouth and largemouth bass. In the river owners. Thus, management of these dams itself, carp, suckers, and bluntnose min- and their associated impoundments poses nows were much less abundant than in the an ecologically and socially complex impoundment; carp made up 83% of the problem in the Milwaukee River Basin. catch in the impoundment but only 23% In West Bend, a wooden dam was above and below it. built across the river as early as 1870 to After considerable public discussion, operate a woolen mill. In 1919 it was the City decided to accept the Department’s replaced by a concrete dam, which was recommendation to remove the dam, The impoundment in operated privately for nearly 40 years to rehabilitate the riverbed, and stock the Milwaukee River at produce hydropower. The City took gamefish. The goal was to restore self- West Bend was drawn ownership of the dam in 1959. By 1987, sustaining habitat for smallmouth bass and down in February 1988 the dam was in obvious need of removal or other native fish species, to eliminate to prepare for dam removal. Photo by Paul replacement. The City either had to remove barriers to fish migration, to improve water Kanehl. the dam and restore the associated riverbed quality, to create an urban park along the shoreline to provide recreational opportunity, and to use cost-effective methods to achieve these goals. The dam was removed in May 1988. After dam removal, Department managers did some habitat improvement work throughout 1989 and 1990, including removing material from a portion of the channel area; placing logs, tree root masses, boulders, and similar materials underwater to create “instant” habitat; and rip-rapping some areas to prevent erosion. However, most of the formerly impounded area was allowed to recover naturally, without management.

202 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE The restoration produced 1.5 miles of free flowing river. Fish access to one mile of river upstream from the former impoundment was also regained. Floodplain areas of the former impoundment were developed as parkland, and oaks and maples were planted along the banks. Aquatic habitat quality improved dramatically. Aquatic vegetation quickly returned. Carp populations declined, and smallmouth bass and panfish populations increased. One threatened species, the greater redhorse, is now found in this restored area of the river.

This section of the Milwaukee River was restored following dam removal in May 1988. The formerly fragmented habitat is once again a free- flowing river with abundant aquatic vegetation and a diverse fish community. This photo was taken in June 1991. Photo by Paul Kanehl.

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206 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Ecological Analysts. 1981. Survey of Forney, J.L. 1974. Interactions between freshwater mussels (Pelecypoda: yellow perch abundance, walleye Unionacea) at selected sites in Pools 11 predation and survival of alternate prey through 24 of the Mississippi River. in Oneida Lake, New York. Trans. Am. U.S. Army Corps of Engineers, Rock Fish. Soc. 103(1):15-24. Island District, Rock Island, Ill. 188 pp. Fredricksen, R.L. 1970. Erosion and Eddy, S. and J.C. Underhill. 1974. Northern sedimentation following road fishes. Univ. Minn. Press, Minneapolis. construction and timber harvest on 414 pp. unstable soils in three small western Ellis, M.M. 1936. Erosion silt as a factor in Oregon watersheds. U.S. Dep. Ag., For. aquatic environments. Ecology Ser. Res. Pap. PNW-104. 17(1):29-42. Fremling, C.R., J.L. Rasmussen, R.E. Engel, S. and S.A. Nichols. 1994. Restoring Sparks, S.P. Cobb, C.F. Bryan, and T.O. Rice Lake at Milltown, Wisconsin. Wis. Claflin. 1989. Mississippi River Dep. Nat. Resour. Tech. Bull. No. 186. fisheries: a case history. pp. 309-351 in 42 pp. D.P. Dodge, ed. Proceedings of the international large river symposium. Extence, C.A. 1981. The effect of drought Can. Spec. Pub. Fish. Aquat. Sci. No. on benthic invertebrate communities in 106. 628 pp. a lowland river. Hydrobiologia 83:217- 224. Frey, D.G. 1963. Wisconsin: the Birge-Juday era. in D.G. Frey, ed. Limnology of Fago, D. 1992. Distribution and relative North America. Univ. Wis. Press. abundance of fishes in Wisconsin. VIII. Final Report. Wis. Dep. Nat. Resour. Fuller, S.L.H. 1974. Clams and mussels Tech. Bull. No. 175. (Mollusca:Bivalvia) in C.W. Hart, Jr., and S.L.H. Fuller, eds. Pollution Fago, D. and A.B. Hauber. 1993. Black ecology of freshwater invertebrates. redhorse, Moxostoma duquesnei Academic Press, Inc., New York. 389 rediscovered in Wisconsin. Can. Field- pp. Naturalist 107(3):351-352. Gardner, M.B. 1981. Effects of turbidity on Fannucchi, G.T., W.A. Fannucchi, and S. feeding rates and selectivity of bluegills. Craven. 1986. Wild rice in Wisconsin: Trans. Am. Fish. Soc. 110:446-450. its ecology and cultivation. Univ. Wis. Extension Pub. No. G3372. 8 pp. Garton, D.W. and D.J. Berg. 1990. Occurrence of Bythotrephes cederstroemi Fausch, K.D., J. Lyons, J.R. Karr, and P.L. (Schoedler 1877) in Lake Superior, Angermeier. 1990. Fish communities as with evidence of demographic variation indicators of environmental within the Great Lakes. J. Great Lakes degradation. Am. Fish. Soc. Symp. Res. 16:148-152. 8:123-44. Gersich, F. M. and M. A. Brusven. 1981. Fisher, S.G. and A. LaVoy. 1972. Differences Insect colonization rates in near-shore in littoral fauna due to fluctuating water regions subjected to hydroelectric levels below a hydroelectric dam. J. power peaking flows. J. Freshw. Ecol. Fish. Res. Bd. Can. 29:1472-1476. 1:231-236. Forbes, A.M. 1985. Summary of survey Giesy, J.P., J.P. Ludwig, and D.E. Tillit. 1994. data for smallmouth bass in Wisconsin Deformities in birds of the Great Lakes streams, 1952-80. Wis. Dep. Nat. region: assigning causality. Environ. Sci. Resour. Bur. Res. Rep. No. 133. Technol. 28(3):128-135.

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Gray, J.R.A. and J.M. Edington. 1969. Effect Heath, D.J. 1992. A preliminary survey for of woodland clearance on stream rare reptiles at selected Wisconsin temperature. Can. J. Fish. Aquat. Sci. Valley Improvement Company 26:399-403. impoundment projects located in Greene, C.W. 1935. The distribution of Wisconsin. Wis. Dep. Nat. Resour., Wisconsin fishes. State of Wis. Conserv. North Central District Headquarters, Comm., Madison. 235 pp. Rhinelander. Gresswell, R.E. and J.D. Varley. 1988. Heath, D.J. 1993a. Justification and Effects of a century of human influence feasibility analysis for freshwater mussel on the cutthroat of Yellowstone Lake. reestablishment upstream of the Prairie Am. Fish. Soc. Symp. 4:45-52. du Sac dam (FERC project #11162), Wisconsin River, Wisconsin. Wis. Dep. Hall, R.J. and E. Kolbe. 1980. Nat. Resour., North Central District Bioconcentrations of organophosphorus Headquarters, Rhinelander. pesticides to hazardous levels by amphibians. J. Toxicol. Environ. Health Heath, D.J. 1993b. A preliminary survey for 6:853-60. rare reptiles at the Wausau hydroelectric project (FERC # 1999) Handford, P., G. Bell, and T. Reimchen. located in Marathon county, Wisconsin. 1977. A gillnet fishery considered as an Wis. Dep. Nat. Resour., North Central experiment in artificial selection. Can. J. District Headquarters, Rhinelander. Fish. and Aquat. Sci. 34:954-61. Heath, D.J. 1993c. A preliminary survey for Hansen, M.J. 1989. A walleye population the Wisconsin state threatened model for setting harvest quotas. Wis. Blanding’s turtle (Emydoidea blandingii) Dep. Nat. Resour. Bur. Fish. Manage. at the Kings (FERC #2239) (Lake Alice) Rep. No. 143. 12 pp. hydroelectric project located near Hansen, M.J., M.D. Staggs, and M.H. Hoff. Tomahawk, Wisconsin. Wis. Dep. Nat. 1991. Derivation of safety factors for Resour., North Central District setting harvest quotas on adult walleyes Headquarters, Rhinelander. from past estimates of abundance. Hebert, P.N.D., C.C. Wilson, M.H. Trans. Am. Fish. Soc. 120:620-628. Murdoch, and R. Lazar. 1991. Hansmann, E.W. and H.K. Phinney. 1973. Demography and ecological impacts of Effects of logging on periphyton in the invading mollusc Dreissena coastal streams of Oregon. Ecology polymorpha. Can. J. Zool. 69:405-409. 54:194-198. Helms, D.R. 1974. Shovelnose sturgeon in Hauber, A. 1989. Impact of continuous the Mississippi River, Iowa. Iowa fishing on the walleye population of Conserv. Comm. Comm. Fish. Res. Lake DuBay, Marathon County, Tech. Ser. 74-3. 68 pp. Wisconsin, 1983-84. Wis. Dep. Nat. Hewlett, J.D. and J.C. Fortson. 1982. Resour. Bur. Fish. Manage. Rep. No. Stream temperature under an 142. inadequate buffer strip in the southeast Hazelwood, E. 1970. Frog pond Piedmont. Water Resour. Bull. 18:983- contaminated. Brit. J. Herpe. 4:177-85. 988. Healey, M.C. 1980. Growth and recruitment Hibbert, A.R. 1969. Water yield changes in experimentally exploited lake after converting a forested catchment to whitefish (Coregonus clupeaformis) grass. Water Resour. Res. 5:634-640. populations. Can. J. Fish. and Aquat. Sci. 37:255-67.

208 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Hicks, B.J., R.L. Beschta, and R.D. Harr. Johnson, C.E. 1971. Factors affecting fish 1991. Long-term changes in streamflow spawning. Wis. Cons. Bull. 36(4):16- following logging in western Oregon 17. and associated fisheries implications. Johnson, F.H. 1961. Walleye egg survival Water Resour. Bull. 27:217-226. during incubation on several types of Hill, D.K. and J.J. Magnuson. 1990. bottom in Lake Winnibigoshish, Potential effects of global warming on Minnesota and connecting waters. Tran. the growth and prey consumption of Am. Fish. Soc. 90:312-322. Great Lakes fish. Trans. Am. Fish. Soc. Johnson, T.B. and D.O. Evans. 1990. Size- 119:(2):265-75. dependent winter mortality of young- Hilsenhoff, W.L. 1971. Changes in the of-the-year white perch: climate downstream insect and amphipod warming and invasion of the Laurentian fauna caused by an impoundment with Great Lakes. Trans. Am. Fish. Soc. a hypolimnion drain. Ann. Am. 119(2):301-13. Entomol. Soc. 64:743-746. Jude, J.D. and F.J. Tesar. 1985. Recent Hilsenhoff, W.L. 1987. An improved biotic changes in the inshore forage fish of index of organic stream pollution. Great Lake Michigan. Can. J. Fish. Aquat. Sci. Lakes Entomol. 20:31-39. 42:1154-57. Hindar, K., R. Ryman, and F. Utter. 1991. Junk, W.J., P.B. Bayley, and R.E. Sparks. Genetic effects of cultured fish on 1989. The flood pulse concept in river- natural fish populations. Can. J. Fish. floodplain systems. pp. 110-127 in D.P. and Aquat. Sci. 48:945-57. Dodge, ed. Proceedings of the Hobbs, H.H., III and J.P. Jass. 1988. The international large river symposium. crayfishes and shrimp of Wisconsin. Can. Spec. Publ. Fish. Aquat. Sci. No. Milwaukee Public Museum, Milwaukee, 106. Wis. 177 pp. Kallemeyn, L.W. 1987. Correlations of Holland, L.E. 1987. Effect of brief regulated lake levels and climatic navigation-related dewaterings on fish factors with abundance of young-of- eggs and larvae. N. Am. J. Fish. the-year walleye and yellow perch in Manage. 7:145-147. four lakes in Voyageurs National Park. N. Am. J. Fish. Manage. 7:513-521. Holland, L.E. and M.L. Huston. 1984. Relationship of young-of-the-year Kanehl, P. and J. Lyons. 1992. Impacts of northern pike to aquatic vegetation in-stream sand and gravel mining on types in backwaters of the Upper stream habitat and fish communities, Mississippi River. N. Am. J. Fish. including a survey on the Big Rib River, Manage. 4:514-522. Marathon county, Wisconsin. Wis. Dep. Nat. Resour. Bur. Res. Rep. No. 155. 32 Hornbeck, J.W., R.S. Pierce, and C.A. pp. Federer. 1970. Streamflow changes after forest clearing in New England. Water Kapuscinski, A.R. and L.D. Jacobson. 1987. Resour. Res. 6:1124-1132. Genetic guidelines for fisheries management. Univ. Minn. Sea Grant Jackivicz, T.P., Jr., and L.N. Kuzminski. Res. Rep. No. 17. 1973. A review of outboard motor effects on the aquatic environment. J. Kapuscinski, A.R.D. and J.E. Lannan. 1986. Water Poll. Con. Fed. 45(8):1759- A conceptual genetic fitness model for 1770. fisheries management. Can. J. Fish. Aquat. Sci. 43:1606-16.

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Kapuscinski, A.R. and D.P. Philipp. 1988. Korth, R. 1993. About jewels, magic and Fisheries genetics: issues and priorities lakes. Lake Tides 18(2):4-5. for research and policy development. Kroger, R.L. 1973. Biological effects of Fisheries 13:4-10. fluctuating water levels in the Snake Karr, J.R. 1981. Assessment of biotic River, Grand Teton National Park, integrity using fish communities. Wyoming. Am. Midl. Nat. 89:478-481. Fisheries 6(6):21-27. Kuhm, H.W. 1928. Wisconsin Indian Karr, J.R., L.A. Toth, and D.R. Dudley. fishing - primitive and modern. Wis. 1985. Fish communities of midwestern Archeol. 7:61-115. rivers: a history of degradation. Lagler, K.F., A.S. Hazzard, W.E. Hazen, and BioScience 35(2):90-95. W.A. Tompkins. 1950. Outboard Keddy, P.A. 1983. Shoreline vegetation in motors in relation to fish behavior, fish Axe Lake, Ontario: effects of exposure production, and angling success. Proc. on zonation patterns. Ecology 64:331- 15th N. Am. Wild. Conf., March 6-9, 344. 1950: 280-303. Keddy, P.A. and A.A. Reznicek. 1986. Great Larson, G.L. and S.E. Moore. 1985. Lakes vegetation dynamics: the role of Encroachment of exotic rainbow trout fluctuating water levels and buried into stream populations of native brook seeds. J. Great Lakes Res. 12:25-36. trout in the southern Appalachian Kempinger, J.J., W.S. Churchill, G.R. mountains. Trans. Am. Fish. Soc. Priegel, and L.M. Christensen. 1975. 114:195-203. Estimates of abundance, harvests, and Lathrop, R.C., P.W. Rasmussen, and D.R. exploitation of the fish population of Knauer. 1990. Walleye mercury Escanaba Lake, Wisconsin, 1946-69. concentrations in Wisconsin lakes. Pap. Wis. Dep. Nat. Resour. Tech. Bull. No. presented at International Conference 84. 30 pp. on Mercury, Gavle, Sweden, 11 June Kitchell, J.F. 1992. Introduction: the 1990. rationale and goals for food web Lawrie, A.H. and J.F. Rahrer. 1973. Lake management in Lake Mendota. in J.F. Superior - a case history of the lake and Kitchell, ed. Food web management: a its fisheries. Great Lakes Fish. Comm. case study of Lake Mendota. Springer- Tech. Rep. No. 19. Verlag, New York. 553 pp. Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Kitchell, J.F. and P.R. Sanford. 1992. Jenkins, D.E. McAllister, and J.R. Paleolimnological evidence of food web Stauffer, Jr. 1980. Atlas of North dynamics in Lake Mendota. in J.F. American Fishes. North Carolina State Kitchell, ed. Food web management: a Museum of Nat. Hist., Raleigh. case study of Lake Mendota. Springer- Lemly, A.D. 1982. Modification of benthic Verlag, New York. 553 pp. insect communities in polluted streams: Kleinert, S.J., P.E. Degurse, and J. Ruhland. combined effects of sedimentation and 1974. Concentration of metals in fish. nutrient enrichment. Hydrobiologia Wis. Dep. Nat. Resour. Tech. Bull. No. 87:229-245. 74. Leonard, P.M. and D.J. Orth. 1988. Use of Konrad, J.G. and S.J. Kleinert. 1974. habitat guilds of fishes to determine Removal of metals from waste waters by instream flow requirements. N. Am. J. municipal sewage treatment plants. Fish. Manage. 8(4):399-409. Wis. Dep. Nat. Resour. Tech. Bull. No. 74.

210 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Liddle, M.J. and H.R.A. Scorgie. 1980. The Lyons, J. 1989b. Changes in the abundance effects of recreation on freshwater of small littoral-zone fishes in Lake plants and animals: a review. Biol. Mendota, Wisconsin. Can. J. Zool. Cons. 17:183-206. 67:2910-2916. Lillie, R.A. and J.W. Mason. 1983. Lyons, J. 1992. Using the index of biotic Limnological characteristics of integrity (IBI) to measure Wisconsin lakes. Wis. Dep. Nat. Resour. environmental quality in warmwater Tech. Bull. No. 138. 116 pp. streams of Wisconsin. U.S. Dep. Ag., Linder, G., J. Barbitta, and T. Kwaiser. 1990. For. Serv., N. Cent. For. Exp. Stn., Gen. Short-term amphibian toxicity tests and Tech. Rep. NC-149. 51 pp. paraquat toxicity assessment. pp. 189- Lyons, J. 1993. Status and biology of 98 in W.G. Landis and W.H. van der paddlefish (Polyodon spathula) in the Schalie, eds. Aquatics toxicology and lower Wisconsin river. Trans. Wis. risk assessment. Vol. 13. Am. Soc. for Acad. Sci. Arts Let. 81:123-135. Testing and Materials, Philadelphia. Lyons, J., A.M. Forbes, and M.D. Staggs. Lloyd, D.S., J.P. Koenings, and J.D. 1988. Fish species assemblages in LaPerriere. 1987. Effects of turbidity in southwestern Wisconsin streams with fresh waters of Alaska. N. Am. J. Fish. implications for smallmouth bass Manage. 7:18-33. management. Wis. Dep. Nat. Resour. Lubinski, K.S. and S. Gutreuter. 1993. Tech. Bull. No. 161. Ecological information and habitat Mack, W.N. and F.M. D’Itri. 1973. Pollution rehabilitation on the Upper Mississippi of a marina area by watercraft use. J. River. pp. 87-100 in L.W. Hesse, C.B. Water Poll. Cont. Fed. 45(1):97-104. Stalnaker, N.G. Benson, and J.R. Zuboy, Mackie, G.L. 1991. Biology of the exotic eds. Proceedings of the symposium on zebra mussel, Dreissena polymorpha, in restoration planning for the rivers of the relation to native bivalves and its Mississippi River ecosystem. U.S. Dep. potential impact in Lake St. Clair. Int., Nat. Biol. Surv., Biol. Rep. No. 19. Hydrobiologia 219:251-268. 502 pp. Magnuson, J.J. 1976. Managing with Ludwig, D., R. Hilborn, and C. Walters. exotics—a game of chance. Trans. Am. 1993. Uncertainty, resource Fish. Soc. 105(1):1-9. exploitation, and conservation: lessons from history. Science 260:17. Magnuson, J.J., J.D. Meisner, and D.K. Hill. 1990. Potential changes in thermal Lynch, J.A., G.B. Reshez, and E.S. Corbett. habitat of Great Lakes fish after global 1984. Thermal alteration of streams climate warming. Trans. Am. Fish. Soc. draining clear-cut watersheds: 119(2):254-64. quantification and biological implications. Hydrobiologia 111:161- Mandrak, N.E. 1989. Probable invasions of 169. the Great Lakes by new species due to climate warming. J. Great Lakes Res. Lyons, J. 1989a. Correspondence between 15:306-16. the distribution of fish assemblages in Wisconsin streams and Omernik’s Manning, R.E. 1979. Impacts of recreation ecoregions. Am. Midl. Nat. 122:163-82. on riparian soils and vegetation. Wat. Res. Bull. 15(1):30-43.

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212 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Morizot, D.C., S.W. Calhoun, L.L. Clepper, Napoli, J. 1975. The coasts of Wisconsin. M.E. Schmidt, J.H. Williamson, and Univ. Wis. Sea Grant WIS-SG-75-122, G.J. Carmichael. 1991. Multispecies March 1975. hybridization among native and Narf, R.P. 1985. Aquatic insect colonization introduced centrarchid basses in central and substrate changes in a relocated Texas. Trans. Am. Fish. Soc. 120:283- stream segment. Great Lakes Entomol. 289. 18(2):83-92. Morton, J.W. 1977. Ecological effect of Neel, J.K. 1963. Impact of reservoirs. pp. dredging and dredge spoil disposal: a 575-593 in D.G. Frey, ed. Limnology in literature review. U.S. Fish and Wild. North America. Univ. Wis. Press, Serv., Tech. Pap. 94, 33 pp. Madison. Mosindy, T.A., W.T. Momot, and P.J. Colby. Neves, R.J. and J.C. Widlak. 1987. Habitat 1987. Impact of angling on the ecology of juvenile freshwater mussels production and yield of mature (Bivalvia:Unionidae) in a headwater walleyes and northern pike in a small stream in Virginia. Amer. Malacol. Bull. boreal lake in Ontario. N. Am. J. Fish. 5:1-7. Manage. 7:493-501. Newbold, J.D., D.C. Erman, and K.B. Roby. Moss, B. 1977. Conservation problems in 1980. Effects of logging on the Norfolk broads and rivers of East macroinvertebrates in streams with and Anglia, England - phytoplankton, boats without buffer strips. Can. J. Fish. and the causes of turbidity. Biol. Cons. Aquat. Sci.37:1076-1085. 12:95-114. Newcombe, C.P. and D.D. MacDonald. Mossman, M.J. and R. Hine. 1984. The 1991. Effects of suspended sediments Wisconsin frog and toad survey: on aquatic ecosystems. N. Am. J. Fish. establishing a long-term monitoring Manage. 11:72-82. program. Wis. Dep. Nat. Resour. Bur. Endangered Resour. Rep. No. 9. 24 pp. Ney, J.J. and M. Mauney. 1981. Impact of a small impoundment on benthic Mossman, M.J. and R. Hine. 1985. macroinvertebrate and ﬁsh Wisconsin’s frog and toad survey. Wis. communities of a headwater stream in Dep. Nat. Resour. Bur. Endangered the Virginia Piedmont. Am. Fish. Soc. Resour. Rep. No. 16. 16 pp. Warmwater Streams Symposium, pp. Mossman, M.J. and J. Huff. 1990. Results of 102-112. Wisconsin’s frog and toad survey 1984- Niemi, G.J. et al. 1990. Overview of case 1989. [unpubl. rep.] studies on recovery of aquatic systems Moyle, P.B. 1991. Ballast water from disturbance. Environ. Manage. introductions. Fisheries 16(1):4-6. 14(5):571-587. Murphy, M.L. and K.V. Koski. 1989. Input Nilles, E. 1993. Emeralds, pygmies, and depletion of woody debris in skimmers, and snaketails: rare Alaska streams and implications for dragonﬂies in Wisconsin. The Niche streamside management. N. Am. J. 6(1):1-4. Fish. Manage. 9:427-36. Noel, D.S., C.W. Martin, and C.A. Federer. Nalepa, T.F. 1989. Estimates of 1986. Effects of forest clearcutting in macroinvertebrate biomass in Lake New England on stream Michigan. J. Great Lakes Res. 15:437- macroinvertebrates and periphyton. 443. Environ. Manage. 10:661-670.

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Nuhfer, A.J. and G.R. Alexander. 1991. Pflieger, W.L. 1971. A distributional study Growth, survival and vulnerability of of Missouri River fishes. Univ. Kans., angling of three wild brook trout strains Lawrence. Publ. Mus. Nat. Hist. exposed to different levels of angler 20(3):225-570. exploitation over time. Mich. Dep. Nat. Philipp, D.P. 1991. Genetic implications of Resour. Fish. Res. Rep. No. 1973. introducing Florida largemouth bass. Olsen, T.M., D.M. Lodge, G.M. Capelli, and Can. J. of Fish. and Aquat. Sci. R.J. Houlihan. 1991. Mechanisms of 48(Suppl. 1):58-65. impact of an introduced crayfish Philipp, D.P., W.F. Childers, and G.S. Whitt. (Orconectes rusticus) on littoral cogeners, 1983. A biochemical genetic evaluation snails and macrophytes. Can. J. Fish. of the northern and Florida subspecies Aquat. Sci. 48:1853-1861. of largemouth bass. Trans. Am. Fish. Omernik, J.M. and A.L. Gallant. 1988. Soc. 112:1-20. Ecoregions of the upper midwest states. Philipp, D.P., J.M. Epifanio, and M.J. EPA/600/3-88/037. Jennings. 1991. AFS position statement Pajak, P. 1991. Fisheries implications of the - Protecting native fish stocks: the 1990 Farm Bill. Fisheries 16(4):4. elimination of stock transfers (draft). Ill. Panek, F.M. 1979. Cumulative effects of Nat. Hist. Surv., Cent. for Aquat. Ecol. small modifications to habitat. Fisheries [memo 26 Aug 1991] 4(2):54-57. Phillip, D.P. and G.S. Whitt. 1991. Survival Pearce, A.J. and L.K. Rowe. 1979. Forest and growth of northern, Florida and management effects on interception, reciprocal F1 hybrid largemouth bass in evaporation and water yield. J. Hydrol. central Illinois. Trans. Am. Fish. Soc. 18:73-87. 120:58-64. Pechmann, J.H.K., D.E. Scott, R.D. Policansky, D. 1993. Fishing as a cause of Semlitsch, J.P. Caldwell, L.J. Vitt, and evolution in fishes. pp. 2-18 in T.K. J.W. Gibbons. 1991. Declining Stokes, J.M. McGlade and R. Law, eds. amphibian populations: the problem of The exploitation of evolving resources. separating human impacts from natural Lecture Notes in Bio-Mathematics fluctuations. Science 253:892-95. Volume 99. Springer-Verlag, Berlin. Penaloza, L.J. 1989. Wisconsin recreation Radford, D.S. and R. Hartland-Rowe. 1971. survey - 1986. Wis. Dep. Nat. Resour. A preliminary investigation of bottom Tech. Bull. No. 167. 24 pp. fauna and invertebrate drift in an unregulated and a regulated stream in Penaloza, L.J. 1991. Boating pressure on Alberta. J. Appl. Ecol. 8:883-903. Wisconsin’s lake and rivers. Wis. Dep. Nat. Resour. Tech. Bull. No. 174. 52 Regier, H.A., J.J. Magnuson, and C.C. pp. Coutant. 1990. Introduction to proceedings: symposium on effects of Peterman, R.M. 1990. Statistical power climate change on fish. Trans. Am. Fish. analysis can improve fisheries research Soc. 119 (2)173-75. and management. Can. J. Fish. Aquat. Sci. 47:2-15. Reid, D.M. and W.T. Momot. 1985. Evaluation of pulse fishing for the Peterman, R.M. and M.J. Bradford. 1987. walleye, Stizostedion vitreum vitreum, in Statistical power of trends in fish Henderson Lake, Ontario. J. Fish Biol. abundance. Can. J. Fish. Aquat. Sci. 27(A):235-51. 44:1879-1889.

214 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Ritchie, J.C. 1972. Sediment, fish and fish Schertzer, W. M. and A.M. Sawchuk. 1990. habitat. J. Soil and Water Cons. 27:124- Thermal structure of the lower Great 125. Lakes in a warm year: implications for Robel, R.J. 1961. Water depth and turbidity the occurrence of hypolimnion anoxia. in relation to growth of Sago Trans. Am. Fish. Soc. 119(2):195-209. Pondweed. J. Wild. Manage. 25(4):436- Schlosser, I.J. 1982. Trophic structure, 438. reproductive success, and growth rate Robins, C.R., R.M. Bailey, C.E. Bond, J.R. of fishes in a natural and modified Brooker, E.A. Lachner, R.N. Lea, and headwater stream. Can. J. Fish. and W.B. Scott. 1991. Common and Aquat. Sci. 39:968-78. scientific names of fishes from the Schneberger, E. and J.L. Funk, eds. 1971. United States and Canada. Am. Fish. Stream channelization: a symposium. Soc. Spec. Publ. No. 20. 183 pp. Am. Fish. Soc. N. Cent. Div. Spec. Publ. Rosenberg, D.M. and A.P. Wiens. 1978. No. 2. Effects of sediment addition on Sedell, J.R. and J.L. Frogatt. 1984. macrobenthic invertebrates in a Importance of streamside forests to northern Canadian river. Water Res. large rivers: the isolation of the 12:753-763. Willamette River, Oregon, USA, from its Ryder, R.A. and C.J. Edwards. 1985. A floodplain by snagging and streamside conceptual approach for the application forest removal. Verh. Int. Ver. Limnol. of biological indicators of ecosystem 22:1828-34. quality in the Great Lakes basin. Rep. to Sedell, J.R. and F.J. Swanson. 1984. Great Lakes Sci. Adv. Bd., Int. Joint Ecological characteristics of streams in Comm., Windsor, Ontario. March, old-growth forests of the Pacific 1985. Northwest. pp. 9-16 in Fish and Sagar, A.D. 1991. Pest control strategies: wildlife relationships in old-growth concerns, issues and options. Environ. forests. Am. Inst. of Fish. Res. Biol. Impact Assess. Rev. 11(3):257-79. Morehead City, N.C. Salmon, A. and R.H. Green. 1983. Seeb, J.E., L.W. Seeb, D.W. Oates, and F.M. Environmental determinants of unionid Utter. 1987. Genetic variation and clam distribution in the middle Thames postglacial dispersal of populations of River, Ontario. Can. J. Zool. 61:832- northern pike (Esox lucius) in North 838. America. Can. J. Fish. and Aquat. Sci. 44:556-61. Sanders, H.O. 1970. Pesticide toxicity to tadpoles of the western chorus frog Selgeby, J.H. 1974. Immature insects (Pseudacris triseriata) and Fowler’s toad (Plecoptera, Trichoptera, and (Bufo woodhousii fowleri). Copeia Ephemeroptera) collected from deep 1970:246-51. water in western Lake Superior. J. Fish. Res. Bd. Can. 31:109-111. Sanderson, M. 1987. Implications of climatic change for navigation and Shapiro, J., V. Lamarra, and M. Lynch. power generation in the Great Lakes. 1975. Biomanipulation: an ecosystem Environ. Can., Clim. Change Digest approach to lake restoration. in P. L CCD 87-03, Toronto. Brezonik and J.L Fox, eds. Proceedings of a symposium on water quality Schaeffer, J.S. and F.J. Margraf. 1987. management through biological control. Predation of fish eggs by white perch, Univ. Flor., Gainesville. Morone americana, in western Lake Erie. Environ. Biol. Fish. 18(1):77-80.

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Shuter, B.J. and J.R. Post. 1990. Climate, Staggs, M.D., R.C. Moody, M.J. Hansen, and population viability, and the M.H. Hoff. 1990. Spearing and sport zoogeography of temperate fishes. angling for walleye in Wisconsin’s Trans. Am. Fish. Soc. 119(2):314-36. ceded territory. Wis. Dep. Nat. Resour. Simmons, G.M. and J.R. Voshell. 1978. Pre- Bur. Fish. Manage. Admin. Rep. No. 31. and post-impoundment benthic Stansbery, D.H. 1970. American macroinvertebrate communities of the malacological union symposium: rare North Anna River. pp. 46-51 in J. and endangered mollusks. 2. Eastern Cairns, E.F. Benfield, and J.R. Webster, freshwater mollusks. 1. The Mississippi eds. Current perspectives on river- and St. Lawrence river systems. reservoir ecosystems. N. Am. Benth. Malacologia 10(1):9-22. Soc., Blacksburg, Va. Stark, W.F. 1988. Wisconsin River of Simonson, T. and J. Lyons. 1992. history. [privately printed] 354 pp. Evaluation and monitoring of fish and Stewart, D.J., J.F. Kitchell, and L.B. habitat during priority watershed Crowder. 1981. Forage fishes and their projects. Wis. Dep. Nat. Resour., Bur. salmonid predators in Lake Michigan. Res., Fish Res. Section Progress Rep. 30 Trans. Am. Fish. Soc. 110:751-63. pp. Stewart, K.W. and C.C. Lindsey. 1983. Smart, M.M., R.G. Rada, D.N. Nielsen, and Postglacial dispersal of lower T.O. Claflin. 1985. The effect of vertebrates in the Lake Agassiz region. commercial and recreational traffic on in J.T. Teller and L. Clayton, eds. Glacial the resuspension of sediment in Lake Agassiz. Geol. Assoc. Can. Spec. navigation pool 9 of the Upper Pap. 26:392-419. Mississippi River. Hydrobiologia 126:263-274. Stone, R. 1994. Dioxins dominate Denver gathering of toxicologists. Science Smith, P. W. 1968. An assessment of 266:1162-1163. changes in fish fauna of two Illinois rivers and its bearing on their future. Strahler, A.N. 1957. Quantitative analysis of Trans. Ill. Acad. Sci. 61(1):31-45. watershed geomorphology. Trans. Am. Geophys. Union 38:913-20. Spangler, G.R., N.R. Payne, J.E. Thorpe, J.M. Byrne, H.A. Regier, and W.J. Strayer, D.L. and J. Ralley. 1993. Christie. 1977. Responses of percids to Microhabitat use by an assemblage of exploitation. Can. J. Fish. Aquat. Sci. stream-dwelling unionaceans (Bivalvia), 34:1983-1988. including two rare species of Alasimidonta. J. N. Am. Benthol. Soc. Spence, J.A. and H.B.N. Hynes. 1971. 12:247-258. Differences in benthos upstream and downstream of an impoundment. J. Sweeney, B.W. 1993. Effects of streamside Fish. Res. Bd. Can. 28:35-43. vegetation on macroinvertebrate communities of White Clay Creek in SPOF. 1983. Strategic planning for Ontario eastern North America. Proceedings of fisheries: the identification of the Academy of Natural Sciences of overexploitation. Report of SPOF Philadelphia 144:291-340. Working Group No. 15, Ontario Ministry of Nat. Resour. 85 pp. The Nature Conservancy. 1994. The conservation of biological diversity in Staggs, M.D. 1987. Sampling design for fish the Great Lakes ecosystem: issues and contaminant monitoring program in opportunities. TNC Great Lakes Lake Michigan. Wis. Dep. Nat. Resour. Program, Chicago Ill. Bur. Res. Rep. No. 140.

216 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Thiel, P.A. 1981. A survey of Unionid U.S. Department of the Interior. 1991. mussels in the Upper Mississippi River Casting light upon the waters: a joint (pools 3 through 11). Wis. Dep. Nat. fishery assessment of the Wisconsin Resour., Tech. Bull. 124, Madison. 24 ceded territory. U.S. Dep. Int., Bur. pp. Indian Aff., Minneapolis, Minn. 102 pp. Threinen, C.W. 1982. The nature of the bait U.S. Department of the Interior/U.S. business in Wisconsin: an informational Department of Commerce. 1993. 1991 report to the Natural Resources Board. National survey of fishing, hunting and Wis. Dep. Nat. Resour. Adm. Rep. No. wildlife-associated recreation. U.S. Dep. 13. 10 pp. Int., Fish and Wildl. Serv. and U.S. Thuemler, R., A. Hauber, R. Theis, F. Pratt, Dep. Commerce, Bur. Census. P. Pajak, M. Staggs, L. Kernen, and D. Washington. 125 pp. Gebken. 1989. FERC Project Review Van Oosten, J. 1945. Turbidity as a factor in Procedures, Vol. 1 Wis. Dep. Nat. the decline of Great Lakes fishes with Resour. Bur. Fish. Manage., Madison special reference to Lake Erie. Trans. [unpubl. rep.] Am. Fish. Soc. 75:281-322. Todd, T.N. 1990. Genetic differentiation of Vannote, R.L., G.W. Minshall, K.W. walleye stocks in Lake St. Clair and Cummins, J.R. Sedell, and C.E. western Lake Erie. U.S. Fish and Wildl. Cushing. 1980. The river continuum Serv. Fish and Wildl. Tech. Rep. 28. concept. Can. J. Fish. and Aquat. Sci. Tonn, W.M. 1990. Climate change and fish 37:130-37. communities: a conceptual framework. Vennum, T. 1988. Wild rice and the Trans. Am. Fish. Soc. 119(2):337-52. Ojibway people. St. Paul Minn Tonn, W.M. and J.J. Magnuson. 1982. Historical Society Press. 358 pp. Patterns in the species composition and Verry, E.S. 1986. Forest harvesting and richness of fish assemblages in northern water: the lake states experience. Water Wisconsin lakes. Ecology 63(4):1149- Resour. Bull. Rep. No. 84184:1039- 66. 1047. Treloar, D. and T. Ehlinger. 1991. Walleye Vetrano, D.M. 1988. Unit construction of stock differentiation and analysis of trout habitat improvement structures recruit contributions in the Lake for Wisconsin coulee streams. Wis. Winnebago system. Wis. Dep. Nat. Dep. Nat. Resour. Bur. Fish. Manage. Resour. Sport Fish Restor. Grant F-83-R Admin. Rep. No. 27. Inter. Rep. Vogt, R.C. 1981. Natural history of Trotzky, H.M. and R.W. Gregory. 1974. The amphibians and reptiles of Wisconsin. effects of water flow manipulation Milwaukee Pub. Mus., Milwaukee, Wis. below a hydroelectric power dam on 205 pp. the bottom fauna of the Upper Wall, G. and C. Wright. 1977. The Kennebec River, Maine. Trans. Am. environmental impact of outdoor Fish. Soc. 103:318-324. recreation. Univ. Waterloo, Ontario, Tyus, H.M. 1990. Effects of altered stream Dep. Geography Publication Series 11. flows on fishery resources. 69 pp. Fisheries 15(3):18. Ward, J.V. 1976. Effects of flow patterns U.S. Department of Energy. 1990. Energy below large dams on stream benthos: a and climate change. Rep. DOE Multi- review. pp. 235-253 in J.F. Orsborn and Lab. Climate Change Comm. Lewis C.H. Allman, eds. Instream flow needs Publishers, Chelsea, Mich. 160 pp. symposium, Vol. 2. Am. Fish. Soc., Bethesda.

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218 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Yousef, Y.A., W.M. McLellon, and H.H. Zillioux, E.J., D.P. Porcella, and J.M. Benoit. Zebuth. 1980. Changes in phosphorus 1993. Mercury cycling and effects in concentrations due to mixing by freshwater wetland ecosystems. motorboats in shallow lakes. Water Res. Environ. Toxicol. Chem. 12:2245-2264. 14:841-852.

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based on ecological associations of plants and animals. Also known as biomes and major life zones. Examples are tundra, northern coniferous forest, desert, grassland, and tropical forest. APPENDIX ? Bracken-Grassland. The northern version of prairie, similar in structure but floristically very different, with bracken fern being the dominant species. Brush Prairie. Prairie with young trees and Glossary shrubs less than 6 ft tall, maintained in this condition by repetitive fire. Capability (Ecological). The long-term ability of an ecosystem to sustain diverse assemblages of microbial, plant, biotic Environment. The animal, and human communities and to nonliving components of the retain the integrity of ecological environment. composition, structure, and function under different management Adaptive Management. A alternatives. formal, structured approach to dealing with uncertainty Cedar Glade. Savannas occurring on dry in natural resource management, using limestone bluffs, with red cedar more the experience of management as an prevalent than oaks. ongoing, continually improving Climax Community. In theory, the final process. community in a successional series, a Barrens. Areas of sandy soil dominated by community that is self-perpetuating, in grasses, low shrubs, and small trees and equilibrium with the physical habitat, subject to frequent wildfire. In general, and controlled by the regional climate. the barrens community takes the form Disturbance and topographic and soil of pine barrens in northern and central conditions strongly influence the Wisconsin and oak barrens in southern species we associate with climax and west-central Wisconsin. Bracken conditions at a particular sites. grasslands are also a part of the barrens Community. An assemblage of species living community. together in a particular area, at a Biological Diversity. The spectrum of life particular time, in a prescribed habitat. forms and the ecological processes that Communities usually bear the name of support and sustain them. Biological their dominant plant species but diversity occurs at four interacting include all of the microbes, plants, and levels: genetic, species, community, and animals living in association with the ecosystem. In shortened form, dominant plant species at a given time. biological diversity is known as Community Diversity. The variety and type biodiversity. of species present in a community, the Biosphere. The parts of the earth’s surface complexity of their interactions, and the layers, waters, and atmosphere in which age and stability of the community. The living things exist. community diversity of a region is influenced by the number of Biotic Environment. The living components communities present, the degree of of the environment. difference among the communities, and Biotic Province. Subdivisions of the earth’s how the communities are distributed surface into geographical land units across the region.

220 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Composition. The makeup of an ecological Ecosystem Management. A system to assess, unit in terms of the organisms or conserve, protect, and restore the groups of organisms present in a unit composition, structure, and function of area or geographical area contains. ecosystems, to ensure their Critical Thinking. A process of reflection and sustainability across a range of temporal analysis that involves the identification and spatial scales and to provide of assumptions, the identification of desired ecological conditions, economic existing knowledge, the exploration of products, and social benefits. alternatives, and the integration of new Edge. The zone where two different habitat understandings into thought and types meet. It can range from an abrupt behavior patterns. As applied to natural change from one to the other (hard resource or environmental decision- edge) to a gradual integration of the making, it results in the integration of two (soft edge). current scientific knowledge and Edge Effects. The ecological impact of clarified values. interfacing two or more habitat types. Dystrophic Lakes. Nutrient-limited lakes, Edge is inherent or natural in nature with low species diversity, simplified but can have negative impacts if its systems, and extremely low levels of creation alters ecological processes. In energy transfer (i.e., short food chains) general, edge effects increase in relation (also known as bog lakes). to the dissimilarity between adjoining cological Processes. Actions or habitats. events that link organisms and Environmental Pollution. The human- their environment. Examples are induced movement of many types of nutrient cycling, carbon cycling, substances within and between air, predation, and primary land, and water components of productivity. ecosystems in quantities and at rates Ecoregion. Areas of relatively homogenous that adversely impact organisms, ecological systems. Ecoregions are habitats, communities, ecosystems, or usually based on patterns of land use, public health. topography, present and potential Eutrophic Lakes. Lakes that are high in natural vegetation, and soils. Ecoregion nutrients but continue to show normal designations are used by resource species diversity and ecological managers to develop logical, regional processes. strategies for land acquisition and Fen. A highly restricted type of wet prairie management. that supports an unusually specialized Ecosystem. A biotic community and its flora. It forms on wet to moist and often abiotic environment, considered peaty, calcareous soils that have together as a unit. Ecosystems are developed over a diffuse groundwater characterized by flow of energy that discharge area that is often under leads to trophic structure and material artesian pressure. cycling (i.e., exchange of materials Fragmentation. The breaking up of large and between living and nonliving parts). continuous ecosystems, communities, Ecosystem is a shortened form of the and habitats into smaller areas term ecological system. surrounded by altered or disturbed land Ecosystem Diversity. The diversity in or aquatic substrate. structure and function within an Function. The roles played by the biotic and ecosystem. It is determined by the abiotic components of ecosystems in amount and complexity of linkages driving the processes (e.g., carbon among the plants and animals of an cycle, water cycle, nutrient cycle) that ecosystem and their abiotic sustain the ecosystem. environment.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 221 GLOSSARY

Genes. The functional unit of heredity—the Landscape Scale. The appropriate spatial components of DNA molecules that and temporal scale for planning, control heredity characteristics in analysis, and improvement of organisms. management activities to sustain Genetic Diversity. The spectrum of genetic ecosystem capability and achieve material carried by different organisms. ecosystem management objectives. Genetic diversity has the potential to Mesotrophic Lakes. Midway between increase or decrease over time due to oligotrophic and eutrophic lakes in recombination. nutrients. Geographic Information System (GIS). A Northern Forest. A Wisconsin community system of computer hardware and characterized by mixed deciduous and software that can input, manipulate, coniferous tree species. In broader and analyze large amounts of terms, it may be characterized as a geographically referenced data to region representing the area north of support the decision-making processes the tension zone that divides Wisconsin of an organization. into two distinct climatic zones. Geographically Referenced Data. Oak Opening. Savanna on rich, mesic soils Information that is spatially keyed to a with mostly bur or white oak. coordinate system for the earth so that Old Growth. A community with dominant different data layers (or maps) can be trees at or near biological maturity. The overlayed or integrated. This type of age and structure of an old-growth data is the foundation of Geographic community varies with species and site. Information Systems (GIS). Old-growth stands are sometimes Grassland. Refers collectively to several characterized by a multi-layered, native Wisconsin plant communities, uneven age and size class structure; a including prairie, brush prairie, sand high degree of compositional and barrens, bracken-grassland, fen, and structural patchiness and heterogeneity; sedge meadow. and significant amounts of woody abitat. The place where an debris and tip-up mounds. organism lives and its Oligotrophic Lakes. Lakes that are low in surrounding environment, nutrients, with low levels of energy including its biotic and abiotic transfer and simplified systems, but not components. Habitat includes to the extent of dystrophic lakes. everything an organism needs to Oligotrophic lakes, such as Lake survive. Superior, are often considered to be the Hypereutrophic Lakes. Lakes that have epitome of desirable water quality nutrient levels so high that the conditions. functions of the system may be affected. Postglacial. The period after the melting of Interior Habitat. That portion of a the last glacier (the Wisconsin Glacier), community not influenced by edge from approximately 10,500 years ago effects because it is far enough removed up to the present. from its outside boundaries. Prairie. A plant community dominated by Isolation Effects. The impact of isolating grasses and forbs, although woody habitat patches from similar habitat shrubs and occasional tree seedlings through fragmentation. may also be present. Prairies frequently grade imperceptibly into other Landscape. An area composed of adjacent communities such as oak savanna and and interacting ecosystems that are sedge meadow. related because of geology, land forms, soils, climate, biota, and human influences.

222 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE Presettlement. The period before the arrival Size Effects. The ecological impact of and extended presence of non-Native decreasing or increasing the size of land American people in Wisconsin. units. egion. Region has no specific Southern Forest. A community spatial or fixed-area definition. characterized by several species of oak Like scale, the appropriate and by the presence of several tree definition of region will vary species normally not found north of the according to the scope of the tension zone (i.e., shagbark hickory, problem or project being hackberry, boxelder, and black walnut). considered. The geographic area Spatial Scale. The geographic size of a included in any particular definition of community or ecosystem. Spatial scale region will be determined by our can range from a microsite such as the knowledge of the breadth of the underside of a leaf on the forest floor, to interconnections among the biotic a forest, to the larger landscape. The communities involved. biosphere (i.e., the planet earth) can be Sand Barrens. Similar to dry sand prairie, thought of as the maximum spatial but with far sparser vegetation and scale. generally including exposed sand or Species. A group of individuals that can sandblows. Most sand barrens today are interbreed successfully with one artifacts of post-settlement activity, another, but not with members of other primarily failed attempts at agriculture. groups. Plants and animals are Savanna. A community that was historically identified as belonging to a given part of a larger ecotone complex species based on similar morphological, bordered by the prairies of the west and genetic, and biochemical the deciduous forests of the east. This characteristics. ecotone was a mosaic of plant Species Diversity. The variety of species in community types that represented a an area. It includes not only the continuum from prairie to forest. number of species in the area but also Savannas were the communities in the their relative abundance and spatial middle of this continuum. distribution. Species richness is one Characteristically, savannas have less component of species diversity, but not than 50% crown cover. the only determinant. Scale. The relative amount or degree of Species Richness. The number of species in something. In relation to ecosystems, an area. scale has both spatial and temporal meanings (see spatial scale and Structure. The pattern or physical temporal scale). organization of an area. It has both vertical and horizontal components. Sedge Meadow. Distinguished from wet Succession. prairie by having (1) more sedge than Progressive temporal changes grass vegetation, (2) more organic than in species composition, organic mineral soil, and (3) seasonally structure, and energy flow in a standing water. It also supports a less community. diverse flora than wet prairie. Sustainability. Long-term management of ecosystems Simplification. A reduction in the diversity to meet the needs of of genetic, species, or community present human populations without resources, and a reduction in the interruption, weakening, or loss of the complexity of the interrelationships resource base for future generations. within them. Simplification affects the composition, structure, and/or function of ecosystems.

WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 223 GLOSSARY

allgrass Prairie. The eastern particular discipline (e.g.,ﬁsheries, portion of the grassland biome forestry, wildlife management) to (including Wisconsin), consider the widest range of options for characterized by productive soils, managing a particular landscape unit. ample precipitation, and tall Trophic structure. The inter-relationship of grasses as the dominant all organisms within a community food vegetation.. web as determined by the speciﬁc role Temporal Scale. The time required to each plays as a producer and/or complete a life history event or consumer of energy or food resources. ecological process. Temporal scale can Values. Principles, standards, or qualities vary from a few seconds for considered worthwhile or desirable biochemical reactions to thousands of from a particular viewpoint. years for ecosystem development. For geologic changes, temporal scale Wetland. An area where water is at, near, or reaches millions of years. above the land surface long enough to be capable of supporting aquatic or Transdisciplinary Perspective. The ability to hydrophytic vegetation and which has transcend the focus and tenets of a soils indicative of wet conditions.

224 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 225 APPENDIXES

ﬁr, balsam ...... Abies balsamea hackberry...... Celtis occidentalis hemlock ...... Tsuga canadensis APPENDIX A hickory, bitternut ...... Carya cordiformis hickory, shagbark...... Carya ovata ironwood ...... Ostrya virginiana maple, red ...... Acer rubrum Common maple, sugar ...... Acer saccharum oak, black ...... Quercus velutina oak, black-jack ...... Quercus marilandica and oak, bur ...... Quercus macrocarpa oak, Hill’s = northern pin oak ...... Quercus ellipsoidalis Scientiﬁc oak, red...... Quercus rubra oak, pin ...... Quercus palustris oak, swamp white ...... Quercus bicolor Names of oak, white ...... Quercus alba pine, jack ...... Pinus banksiana pine, Norway = Organisms red pine ...... Pinus resinosa pine, red ...... Pinus resinosa pine, scrub ...... n.f. (jack pine?) Cited pine, white ...... Pinus strobus spruce ...... Picea spp. spruce, white ...... Picea glauca spuce, black ...... Picea mariana TREES tamarack ...... Larix laricina walnut, black ...... Juglans nigra ash, black ...... Fraxinus nigra ash, white ...... Fraxinus americana aspen ...... Populus tremuloides SHRUBS basswood ...... Tilia americana alder, tag ...... Alnus spp. beech, American ...... Fagus grandifolia bearberry ...... Arctostaphylos birch, white...... Betula papyrifera uva-ursi birch, yellow ...... Betula alleghaniensis bilberry, dwarf...... Vaccinium boxelder...... Acer negundo cespitosum butternut ...... Juglans cinerea blueberry ...... Vaccinium cedar, red ...... Juniperus virginiana angustifolium cedar, white ...... Thuja occidentalis also elm ...... Ulmus spp. blueberry, low sweet..... Vaccinium angustifolium elm, American...... Ulmus americana

226 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE buckthorn, common .... Rhamnus cathartica broomrape, Louisiana .. Orobanche cherry, sand...... Prunus pumila ludoviciana cranberry ...... Vaccinium broomrape, macrocarpon one-flowered ...... Orobanche uniflora gooseberry, bulrush, tussock ...... Scirpus cespitosus hawthorn-leaved ...... Ribes oxyacanthoides bush-clover, prairie ...... Lespedeza hazel, beaked ...... Corylus cornuta leptostachya honeysuckle, bush ...... Diervilla lonicera bush-clover, slender ..... Lespedeza virginica honeysuckle, Japanese . Lonicera japonica clover, villous prairie .... Petalostemum villosum huckleberry...... Gaylussacia baccata coneflower, pale purple Echinacea pallida tea, early New Jersey .... Ceanothus herbaceus cow-wheat ...... Melampyrum lineare tea, New Jersey ...... Ceanothus americanus cress, European water .. Rorippa nasturtium- aquaticum willow, sand = sandbar willow or ...... Salix exigua dandelion, prairie...... Agoseris cuspidata = sand-dune willow ..... S. cordata eupatorium, sessile-leaved ...... Eupatorium yew, Canada...... Taxus canadensis sessilifolium fame-flower, prairie ...... Talinum rugospermum FORBS, GRASSES, AND LOWER PLANTS fern, bracken...... Pteridium aquilinum alfalfa ...... Medicago sativa fern, Braun’s holly ...... Polystichum braunii anemone, Carolina ...... Anemone fern, goblin ...... Botrychium mormo caroliniana fern, sweet ...... Myrica asplenifolia anemone, Hudson Bay . Anemone multifida fern, ternate grape ...... Botrychium arbutus, trailing ...... Epigaea repens ternatum asphodel, false ...... Tofieldia glutinosa fescue, western...... Festuca occidentalis aster family ...... Asteraceae foamflower ...... Tiarella cordifolia [Compositae] foxglove, eared false ..... Tomanthera aster, forked ...... Aster fureatus ariculata beardtongue, hairy ...... Penstemon hirsutus foxglove, mullein ...... Dasistoma beardtongue, pale ...... Penstemon pallidus macrophylla bellwort, sessile ...... Uvularia sessilifolia foxglove, pale false ...... Agalinis skinneriana bladderpod ...... Lesquerella foxglove, round- ludoviciana stemmed false ...... Agalinis gattingeri blazing star, marsh ...... Liatris spicata gentian, cream ...... Gentiana sp. n.f. blazing star, dotted...... Liatris punctata gentian, horse ...... Triosteum spp. bluegrass, bog ...... Poa paludigena gentian, prairie ...... Gentiana puberula bluegrass, Kentucky ..... Poa praetensis gentian, small fringed... Gentianopsis procera bluets ...... Houstonia caerulea gentian, yellowish ...... Gentiana alba boneset, woodland ...... Eupatorium giant hyssop, yellow ..... Agastache nepetoides sessilifolium goldenrod, elm-leaved.. Solidago ulmifolia goldenrod, Ohio ...... Solidago ohioensis

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grass, beak = mallow, poppy ...... Callirhoe triangulata beak-rush or ...... Rhynchospora spp. marsh-marigold, = beaked spike-rush ..... Eleocharis rostellata floating ...... Caltha natans grass, brome...... Bromus spp. milfoil, Eurasian grass family ...... Gramineae water...... Myriophyllum spicatum grass, poverty ...... Aristida dichotoma milkweed, Mead’s...... Asclepias meadii grass, quack ...... Elytrigia repens milkweed, prairie ...... Asclepias sullivantia grass, reed canary...... Phalaris arundinacea milkweed, purple ...... Asclipias purpurascens grass, Smith melic ...... Melica smithii milkweed, tall ...... Asclipias exaltata grass, Wilcox’s panic .... Panicum Wilcoxianum milkweed, wooly...... Ascplepias lanuginosa grass-of-parnassus ...... Parnassia spp. milkwort, cross ...... Polygala cruciata grass-of-parnassus, marsh...... Parnassia palustris milkwort, pink ...... Polygala incarnata grass-of-parnassus, moonwort ...... Botrychium lunaria small-flowered ...... Parnassia parviflora moss, reindeer...... Cladonia spp. groundcherry, white ..... Physalis grandiflora nutrush, netted ...... Scleria reticularis hairgrass, common ...... Deschampsia orchid, calypso...... Calypso bulbosa ?flexuosa? orchid, Hooker’s...... Platanthera hookeri heath family ...... Ericaeae [Ericaceae] orchid, prairie hyacinth, wild ...... Camassia scilloides white-fringed ...... Platanthera indigo, false wild ...... Baptisia spp. leucophaea iris, dwarf lake ...... Iris lacustris orchid, small round-leaved...... Platanthera kitten tails ...... Besseya bullii orbiculata lady-slipper, orchid, tubercled...... Platanthera flava ram’s head ...... Cypripedium arietinum orchid, white-fringed ... Platanthera blephariglottis lady-slipper, white ...... Cypripedium candidum parsley, prairie...... Polytaenia nuttallii lead plant ...... Amorpha canescens parsnip ...... ?Pastinaca sativa? legume family ...... Fabaceae pea, beach ...... Lathyrus maritimus lespedeza, Virginia ...... Lespedeza virginica pea, silvery scurfy ...... Psoralea argophylla lettuce, great white ...... Prenanthes petunia, wild ...... Ruellia humilis crepidinea phlox, smooth...... Phlox glaberrima lettuce, rough white ..... Prenanthes aspera pimpernel, yellow ...... Taenidia integerrima lily family ...... Liliaceae pine-drops ...... Pterospora loco weed, Fassett’s ...... Oxytropis campestris andromedea v: chartacea plantain, pale Indian .... Cacalia atriplicifolia loosestrife, purple ...... Lythrum salicaria plantain, prairie Indian Cacalia tuberosa lupine ...... Lupinus perennis plum, prairie ...... Astragalus mallow, glade ...... Napaea dioica crassicarpus

228 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE pomme-de-prairie ...... Psoralea esculenta sundew, English ...... Drosera anglica prickly pear, brittle ...... Opuntia fragilis sundew, linear leaved ... Drosera linearis primrose, toothed sweetfern ...... Comptonia peregrina evening ...... Oenothera switchgrass ...... Panicum virgatum serrulatus thistle, Canada ...... Cirsium arvense puccoon, hoary ...... Lithospermum canescens thistle, dune ...... Cirsium pitcheri quinine, wild ...... Parthenium thistle, prairie...... Cirsium hillii integrifolium thistle, tall ...... Cirsium altissimum redroot ...... Ceanothus ovatus thistle, woodland ...... Cirsium sp. n.f. reed, sand ...... Calamovilfa twayblade, auricled ...... Listera auriculata longifolia twayblade, rice, wild ...... Zizania aquatica broad-leaved ...... Listera rose family ...... Rosaceae convallarioides rush, bald ...... Juncus sp. n.f. valerian, marsh ...... Valeriana sitchensis uliginosa rush, bog ...... Juncus sp. n.f. vetch, Cooper’s milk..... Astragalus neglectus rye, downy wild ...... Elymus villosus violet, sand ...... Viola adunca sagewort, prairie ...... Artemisia frigida wintergreen ...... Gaultheria sedge, chestnut ...... Fimbristylis procumbens puberula sedge, coast ...... Carex sp. n.f. sedge, crow-spur ...... Carex sp. n.f. MAMMALS sedge, drooping ...... Carex prasina badger...... Taxidea taxus sedge, hop-like...... Carex sp. n.f. bear, black ...... Euarctos americanus sedge, lenticular ...... Carex lenticularis beaver ...... Castor canadensis sedge, Michaux’s ...... Carex michauxiana michiganensis sedge, prairie straw ...... Carex suberecta bison ...... Bison bison sedge, Richardson’s ...... Carex richardsonii bobcat ...... Lynx rufus sedge, Torrey’s ...... Carex torreyi caribou, woodland ...... Rangifer caribou senna, Maryland...... Cassia marilandica cougar ...... Felis concolor shinleaf, small ...... Pyrola minor coyote ...... Canis latrans skullcap, small ...... Scutellaria parvula deer, white-tailed ...... Odocoileus spike-rush, angle virginianus stemmed = angled s-r ... Eleocharis elk ...... Cervus canadensis quadrangulata ﬁsher...... Martes pennanti spike-rush, beaked ...... Eleocharis rostellata fox, red ...... Vulpes fulva spike-rush, Robbin’s..... Eleocharis robbinsii ground squirrel, spike-rush, wolf ...... Eleocharis wolﬁi Franklin’s ...... Citellus franklinii spurge, leafy spurge ..... Euphorbia esula ground squirrel, St. John’s wort, thirteen-lined ...... Citellus round-fruited ...... Hypericum tridecemlineatus sphaerocarpum hare, snowshoe ...... Lepus americanus

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lynx ...... Lynx canadensis BIRDS marten, pine ...... Martes americana mice ...... Peromyscus spp. blackbird, Brewer’s ...... Euphagus cyanocephalus mink ...... Mustela vison blackbird, red-winged .. Agelaius phoeniceus mole, prairie ...... Scalopus aquaticus bluebird, eastern ...... Sialia sialis moose, northwestern ... Alces alces bobolink ...... Dolichonyx mouse, harvest ...... Reithrodontomys oryzivorus megalotis bunting, indigo ...... Passerina cyanea mouse, prairie deer ...... Peromyscus maniculatus catbird, gray ...... Dumetella carolinensis mouse, white-footed .... Peromyscus leucopus noveboracensis chickadee, black-capped ...... Parus atricapillus muskrat ...... Ondatra zibethicus chickadee, boreal ...... Parus hudsonicus otter ...... Lutra canadensis cormorant, pocket gopher, plains ... Geomys bursarius double-crested ...... Phalacrocorax porcupine ...... Erethizon dorsatum auritus rabbit, cottontail ...... Sylvilagus floridanus cowbird, rabbit, white-tailed brown-headed ...... Molothrus ater jack ...... Lepus townsendii crane, sandhill ...... Grus canadensis raccoon ...... Procyon lotor crane, whooping ...... Grus americana shrew, least...... Cryptotis parva creeper, brown ...... Certhia americana shrew, masked...... Sorex cinereus crossbill, red ...... Loxia curvirostra shrew, short-tailed...... Blarina spp. crossbill, shrews...... Blarina spp. and white-winged ...... Loxia leucoptera Sorex spp. crow, American ...... Corvus skunk, striped ...... Mephitis mephitis brachyrhynchos squirrel, flying...... Glaucomys sabrinus cuckoo, black-billed..... Coccyzus macrotis erythropthalmus squirrel, fox ...... Sciurus niger cuckoo, yellow-billed ... Coccyzus squirrel, gray ...... Sciurus carolinensis americanus squirrel, red ...... Tamiasciurus curlew, long-billed ...... Numenius hudsonicus americanus vole, prairie ...... Microtus ochrogaster dickcissel ...... Spiza americana voles ...... Microtus spp. dove, mourning ...... Zenaida macroura weasel, long-tailed ...... Mustela frenata eagle, bald ...... Haliaeetus leucocephalus wolf, timber ...... Canis lupus finch, purple ...... Carpodacus wolverine ...... Gulo luscus luscus purpureus woodchuck ...... Marmota monax flicker, northern ...... Colaptes auratus flycatcher, acadian...... Empidonax virescens flycatcher, great crested ...... Myiarchus crinitus

230 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE flycatcher, least ...... Empidonax minimus nuthatch, flycatcher, olive-sided .. Contopus borealis red-breasted ...... Sitta canadensis goldfinch, American ..... Carduelis tristis nuthatch, white-breasted ...... Sitta carolinensis goshawk, northern ...... Accipiter gentilis oriole, orchard ...... Ictreus spurius grackle, common ...... Quiscalus quiscula osprey ...... Pandion haliaetus grosbeak, evening ...... Coccothraustes vespertinus ovenbird ...... Seiurus aurocapillus grosbeak, pine...... Pinicola enucleator owl, barn ...... Tyto alba grosbeak, owl, barred ...... Strix varia rose-breasted ...... Pheucticus owl, great horned ...... Bubo virginianus ludovicianus owl, long-eared ...... Asio otus grouse, ruffed ...... Bonasa umbellus owl, northern grouse, sharp-tailed...... Tympanuchus saw-whet ...... Aegolius acadicus phasianellus owl, short-eared ...... Asio flammeus grouse, spruce ...... Dendragapus parakeet, Carolina ...... Conuropsis canadensis carolinensis gulls ...... Larus spp. parula, northern...... Parula americana harrier, northern ...... Circus cyaneus phalarope, Wilson’s ...... Phalaropus tricolor hawk, broad-winged .... Buteo platypterus pheasant, ring-necked .. Phasianus colchicus hawk, Cooper’s...... Accipiter cooperii phoebe, eastern ...... Sayornis phoebe hawk, red-shouldered .. Buteo lineatus pigeon, passenger...... Ecopistes hawk, sharp-shinned ... Accipiter striatus migratorius jay, blue...... Cyanocitta cristata pintail, northern...... Anas acuta jay, gray...... Perisoreus plover, piping...... Charadrius melodus canadensis prairie chicken, junco, dark-eyed ...... Junco hyemalis greater...... Tympanuchus cupido kestrel, American ...... Falco sparverius quail, bobwhite ...... Colinus virginianus killdeer ...... Charadrius vociferus rail, black ...... Laterallus kingbird, eastern ...... Thrannus tyrannus jamaicensis kinglet, rail, yellow ...... Coturnicops golden-crowned ...... Regulus satrapa noveboracensis kinglet, ruby-crowned.. Regulus calendula raven, common ...... Corvus corax kite, swallow-tailed ...... Elanoides forficatus redstart, American...... Setophaga ruticilla lark, horned ...... Eremophila alpestris robin, American ...... Turdus migratorius loon, common ...... Gavia immer sandpiper, upland ...... Bartramia longicauda meadowlark, eastern .... Sturnella magna sapsucker, meadowlark, western ... Sturnella neglecta yellow-bellied ...... Sphyrapicus varius merlin ...... Falco columbarius shrike, loggerhead ...... Lanius ludovicianus night-heron, sparrow, chipping ...... Spizella passerina yellow-crowned ...... Nyctanassa violacea sparrow, clay-colored ... Spizella pallida nighthawk, common .... Chordeiles minor sparrow, field ...... Spizella pusilla

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sparrow, grasshopper ... Ammodramus warbler, savannarum chestnut-sided ...... Dendroica sparrow, Henslow’s ...... Ammodramus pensyvanica henslowii warbler, Connecticut .... Oporornis agilis sparrow, lark ...... Chondestes warbler, grammacus golden-winged ...... Vermivora sparrow, Le Conte’s ...... Ammodramus chrysoptera leconteii warbler, Kirtland’s ...... Dendroica kirtlandii sparrow, savannah...... Passerculus warbler, magnolia ...... Dendroica magnolia sandwichensis warbler, mourning ...... Oporornis sparrow, sharp-tailed.... Ammodramus philadelphia caudacutus warbler, Nashville ...... Vermivora sparrow, song ...... Melospiza melodia ruﬁcapilla sparrow, vesper ...... Pooecetes gramineus warbler, Tennessee ...... Vermivora peregrina tanager, scarlet ...... Piranga olivacea warbler, teal, blue-winged ...... Anas discors yellow-rumped ...... Dendroica coronata tern, least ...... Sterna antillarum warbler, yellow-throated ...... Dendroica dominica thrasher, brown...... Toxostoma rufum waterthrush, northern .. Seiurus thrush, hermit ...... Catharus guttatus noveboracensis thrush, Swainson’s...... Catharus ustulatus waxwing, cedar ...... Bombycilla thrush, wood ...... Hylocichla mustelina cedrorum towhee, rufous-sided.... Pipilo whip-poor-will ...... Caprimulgus erythrophthalmus vociferus turkey ...... Meleagris gallopavo wood-pewee, eastern ... Contopus virens veery ...... Catharus fuscescens woodcock, American ... Scolopax minor vireo, Bell’s ...... Vireo bellii woodpecker, vireo, red-eyed ...... Vireo olivaceus black-backed...... Picoides arcticus vireo, solitary ...... Vireo solitarius woodpecker, downy ..... Picoides pubescens vireo, warbling ...... Vireo gilvus woodpecker, hairy ...... Picoides villosus vireo, yellow-throated .. Vireo ﬂavifrons woodpecker, pileated ... Dryocopus pileatus warbler, woodpecker, black-and-white ...... Mniotilta varia red-headed...... Melanerpes erythrocephalus warbler, black-throated blue ...... Dendroica wren, sedge ...... Cistothorus platensis caerulescens wren, winter ...... Troglodytes warbler, troglodytes black-throated green .... Dendroica virens yellowthroat, warbler, Blackburnian .. Dendroica fusca common ...... Geothlypis trichas warbler, Canada ...... Wilsonia canadensis warbler, Cape May ...... Dendroica tigrina warbler, cerulean...... Dendroica cerulea

232 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE snake, eastern milk ...... Lampropeltis AMPHIBIANS AND REPTILES triangulum bullsnake ...... Pituophis snake, Eastern melanoleucus plains garter ...... Thamnophis radix radix frog, Blanchard’s cricket ...... Acris crepitans snake, northern blanchardi ribbon ...... Thamnophis sauritus septentrionalis frog, chorus...... Pseudacris triseriata triseriata snake, northern red-bellied ...... Storeria frog, green...... Rana clamitans occipitomaculata melanota snake, queen ...... Regina septemvittata frog, mink ...... Rana septentrionalis snake, ringneck frog, wood ...... Rana sylvatica = prairie ...... Diadophis punctatus lizard, western arnyi slender glass...... Ophisaurus = northern ...... Diadophis punctatus attenuatus edwarsi newt, central ...... Notophthalmus snake, smooth green .... Opheodrys vernalis viridescens snake, western fox ...... Elaphe vulpina racer, blue ...... Coluber constrictor foxi toad, American...... Bufo americanus racerunner, six-lined .... Cnemidophorus treefrog, Cope’s gray ..... Hyla chrysoscelis sexlineatus treefrog, grey = rattlesnake, eastern Cope’s gray treefrog or massasauga ...... Sistrurus catenatus Eastern gray treefrog .... Hyla versicolor rattlesnake, timber ...... Crotalus horridus treefrog, wood = wood frog ...... Rana sylvatica salamander, blue-spotted ...... Ambystoma laterale turtle, Blandings...... Emydoidea blandingi salamander, turtle, common eastern tiger ...... Ambystoma tigrinum snapping ...... Chelydra serpentina salamander, four-toed .. Hemidactylium turtle, ornate box ...... Terrapene ornata scutatum turtle, wood ...... Clemmys insculpta salamander, red-backed ...... Plethodon cinereus salamander, spotted ..... Ambystoma FISH maculatum alewife ...... Alosa skink, ﬁve-lined ...... Eumeces fasciatus pseudoharengus skink, northern bass, smallmouth ...... Micropterus prairie ...... Eumeces dolomieui septentrionalis bass, white ...... Morone chrysops snake, black rat ...... Elaphe obsoleta bass, yellow...... Morone snake, Dekay’s mississippiensis (also brown snake) ...... Storeria dekayi bloater ...... Coregonus hoyi snake, eastern hognose ...... Heterodon bowﬁn ...... Amia calva platyrhinos buffalo, bigmouth ...... Ictiobus cyprinellus

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bullhead...... Ictaluridae lamprey, American burbot...... Lota lota brook ...... Lampetra appendix carp, common ...... Cyprinus carpio lamprey, sea ...... Petromyzon marinus carp, grass ...... Ctenopharyngodon minnow, bluntnose ...... Pimephales notatus idella minnow, bullhead ...... Pimephales vigilax catfish, channel ...... Ictalurus punctatus minnow, Ozark ...... Notropis nubilus chub, gravel ...... Erimystax x- minnow, suckermouth . Phenacobius punctatus mirabilis chubs ...... Corgenonus spp. mudminnow, central .... Umbra limi chubsucker, creek ...... Erimyzon oblongus muskellunge ...... Esox masquinongy cisco, blackfin ...... Coregonus northern pike ...... Esox lucius nigripinnis paddlefish ...... Polyodon spathula cisco, deepwater...... Coregonus johannae perch, white ...... Morone americana cisco, shortjaw ...... Coregonus zenithicus perch, yellow ...... Perca flavescens cisco, shortnose ...... Coregonus reighardi redhorse, black ...... Moxostoma dace, southern duquesnei redbelley ...... Phoxinus redhorse, golden ...... Moxostoma erythrogaster erythrurum darter, bluntnose ...... Etheostoma redhorse, river...... Moxostoma chlorosomum carinatum darter, crystal ...... Ammocrypta redhorse, shorthead ..... Moxostoma asprella macrolepidotum darter, fantail...... Etheostoma ruffe ...... Gymnocephalus flabellare cernua darter, gilt ...... Percina evides salmon ...... Oncorhynchus spp. darter, Iowa...... Etheostoma exile salmon, Atlantic ...... Salmo salar darter, Johnny ...... Etheostoma nigrum sauger ...... Stizostedion darter, least ...... Etheostoma canadense microperca sculpin ...... Cottidae darter, mud ...... Etheostoma shad, Ohio = asprigene Alabama shad...... Alosa alabamae eel, American ...... Anguilla rostrata shiner, bigmouth ...... Notropis dorsalis freshwater drum...... Aplodinotus shiner, ghost ...... Notropis buchanani grunniens shiner, ironcolor...... Notropis chalybaeus gar, longnose ...... Lepisosteus osseus shiner, pallid ...... Notropis amnis gar, shortnose...... Lepisosteus platostomus shiner, pugnose ...... Notropis anogenus goldfish ...... Carassius auratus shiner, redfin...... Notropis umbratilis grayling ...... Thymallus arcticus shiner, sand...... Notropis stramineus herring, lake...... Coregonus artedii shiner, spotfin ...... Notropis spilopterus herring, skipjack ...... Alosa chrysochloris shiner, striped ...... Notropis chrysocephalus kiyi ...... Coregonus kiyi smelt, rainbow ...... Osmeridae

234 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE stickleback ...... Gasterosteidae butterfly, Henry’s elfin .. Incisalia henrici stoneroller, central ...... Campostoma butterfly, hoary elfin ..... Incisalia polia anomalum butterfly, Indian sturgeon, lake ...... Acipenser fluvescens skipper...... Hesperia sassacus sturgeon, shovelnose .... Scaphirhynchus butterfly, Juvenal’s platorynchus dusky wing ...... Erynnis juvenalis sucker, blue ...... Cycleptus elongatus butterfly, Karner blue ... Lycaeides melissa sucker, northern hog .... Hypentelium samuelis nigricans butterfly, Laurentian sucker, white...... Catostomus skipper...... Hesperia comma commersoni laurentina sunfish, green...... Lepomis cyanellus butterfly, Leonard’s skipper...... Hesperia leonardus sunfish, longear...... Lepomis megalotis leonardus topminnow, starhead.... Fundulus notti butterfly, mustard trout...... Salmonidae white...... Pieris napi oleracea trout, brook ...... Salvelinus fontinalis butterfly, northern trout, rainbow ...... Oncorhynchus blue ...... Lycaeidesidas mykiss nabokovi walleye ...... Stizostedion vitreum butterfly, Olympian marble ...... Euchloe olympia butterfly, Persius INSECTS dusky wing ...... Erynnis persius butterfly, pine butterfly, American elfin ...... Incisalia niphon copper ...... Lycaena phlaeas clarki americana butterfly, pink-edged butterfly, brown elfin .... Incisalia augustinus sulphur ...... Colias interior butterfly, chryxus butterfly, Powesheik arctic ...... Oeneis chryxus skipper...... Oarisma powesheik strigulosa butterfly, regal fritillary . Speyeria idalia butterfly, cobweb butterfly, roadside skipper...... Hesperia metea skipper...... Amblyscirtes vialis butterfly, coral butterfly, silvery blue .... Glaucopsyche hairstreak ...... Harkenclenus titus lygdamus couperi butterfly, dusted butterfly, sleepy skipper...... Atrytoropsis hianna dusky wing ...... Erynnis brizo butterfly, Edward’s butterfly, swamp hairstreak ...... Satyrium edwardsii metalmark...... Calephelis muticum butterfly, European butterfly, tawny cabbage ...... Pieris rapae crescent...... Phyciodes batesii butterfly, frosted elfin ... Incisalia irus butterfly, western butterfly, Gorgone tailed blue ...... Everes amyntula checkerspot...... Charidryas gorgone dragonfly ...... Anisoptera carlota

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moth, graceful clearwing ...... Hemaris gracilis OTHER INVERTEBRATES moth, Nevada buck...... Hemileuca clam, Asiatic...... Corbicula fluminea nevadensis clam, winged moth, phlox flower ...... Schinia indiana mapleleaf ...... Quadrula fragosa moth, pink prominent . Hyparpax aurora hydra ...... Hydra spp. moth, silphium borer ... Papaipema silphii jellyfish ...... Scyphozoa moth, Sprague’s mussel, ebony shell ...... Fusconaia ebena pygarctia ...... Pygarctia spraguei mussel, zebra ...... Oreissena [moth] ...... Grammia celia polymorpha [moth] ...... Heliothis borealis rusty crayfish ...... Orconectes rusticus sponge ...... Porifera [cladoceran] ...... Bythotrephes cederstroemi

236 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE 237 APPENDIXES

William Moorman, Bruce Moss, Jeffrey Smoller, and Michael Staggs

APPENDIX B The Applications Team created the ecosystem management decision model and developed a number of the case studies presented in the report. Members were:

Acknowledgements Dennis Schenborn and Darrell Zastrow (Leaders), Walter Adams, Neil Ambourn, Gerry Bartelt, Michael Beaufeaux, Ronald Benjamin, Michael Coshun, Alan Crossley, Jane This report is much more than the ideas Cummings-Carlson, Paul expressed on paper. Perhaps more impor- Cunningham, Ronald Eckstein, Tom tant than the content of these pages is the Eggert, Eric Epstein, Tim Grunewald, process that we navigated to reach this ﬁnal David Hammer, Richard Henderson, copy. The following credits bear witness to John Huff, Thomas Isaac, Kenneth the interest and commitment of the many, Johnson, Paul Kooiker, James many people who shared their time, Kreitlow, James Lealos, Charles Ledin, thoughts, and expertise—and to the Mark Martin, Robert Martini, Robert challenge of writing a Department state- Mather, David Mladenoff, Sam Moore, ment on such an emerging and complex William Moorman, Bruce Moss, Carol issue. Nielsen, Timothy Rasman, James Rau, The Biodiversity Working Group coordi- Michael Staggs, Michael Talbot, Craig nated the overall effort and integrated the Thompson, and Tom Watkins work of the various teams on behalf of the Management Team, Division of Resource The Public Involvement Planning Team Management. Members of the Working prepared short and long range recommen- Group were: dations for including various interest groups, other agencies, and the general Anne Forbes (Leader 1994-95), Betty public in the dialogue on biodiversity. Les (Leader 1993-94), Mary Hamel, Members were: Paul Matthiae, Wendy McCown, Thomas Mickelson, William Mary Hamel (Leader), Kristin Belling, VanderZouwen, Kristin Visser, Wendy David Crehore, John Grosman, David Weisensel, Darrell Zastrow, and Laura Hammer, Kirsten Held, Gary Kulibert, Komai (staff assistant) Thomas Mickelson, Michael Ries, and David Weitz The Biodiversity Technical Team prepared much of the material for the ﬁrst draft of The Editorial and Production Team, whose this report and acted as a discussion body members are listed on the title page, guided to resolve differences of opinion to com- this report through the revision, editing, plete that draft. Members were: and publication stages. In addition to the above team members, we Paul Matthiae (Leader), Ronald offer thanks to the many others who Eckstein, DuWayne Gebken, Jonathan assisted in shaping this report. In the space Gilbert, Richard Henderson, Ralph below, we would like to recognize you and Hewett, Roy Jacobson, John Kotar, thank you for your varied contributions.

238 WISCONSIN’S BIODIVERSITY AS A MANAGEMENT ISSUE During the fall of 1993, over 600 Depart- ment employees participated in a dialogue Rusch, UW-Madison; Craig Shafer, on biodiversity at five regional open houses USDI-National Park Service; Forest and discussion sessions held around the Stearns, UW-Madison; Ann Swengel, state. These employees came from through- North American Butterfly Association; out the agency—foresters, wildlife manag- Stanley Temple, UW-Madison; Jack ers, planners, researchers, water quality Troyer, Chequamegon National specialists, environmental engineers, Forest; Donald Waller, UW-Madison; program assistants, education specialists, and Bill Willers, UW-Oshkosh and others. Their ideas broadened our perspective on biodiversity and resulted in From Wisconsin DNR: Charles major revisions to the draft report. We Adams, Lloyd Andrews, Thomas thank David Lentz for computer and Beard, Ronald Benjamin, Jon database support for the regional meetings Bergquist, Cathy Bleser, Lawrence and Michael Koutnick for assistance with Claggett, Paul Cunningham, Robert the traveling exhibits. In addition, District DuBois, Ronald Eckstein, Tom Eggert, personnel were a tremendous help in Eric Epstein, David Evenson, Bureau setting up the meetings and making them a of Forestry staff, Meg Galloway, success. A report, “The Dialogue on DuWayne Gebken, Beth Goodman, Biodiversity: What We Learned from DNR Mary Hamel, Robert Hansis, Robert Employees at the Six Regional Meetings on Hay, David Heath, Randy Hoffman, Biodiversity,” summarized what we learned Steve Holaday, William Ishmael, at the meetings. We thank Elizabeth Ivers David Ives, Roy Jacobson, Martin for her role in preparing that report. Jennings, Kenneth Johnson, Kelly Scientists within the Department, the Kearns, Greg Kessler, Willard Kiefer, university system, industry, and elsewhere James Lealos, Charles Ledin, John shared their expertise and ideas through Lyons, James March, Mark Martin, formal technical review. Their efforts Robert Martini, Wendy McCown, improved the report and contributed to its James McNelly, David Mladenoff, Sam scientific accuracy. Those providing written Moore, Doug Morrissette, Bruce Moss, reviews were: Timothy Mulholland, Kathleen Patnode, Thomas Pellett, Ronald Poff, David Sample, Patrick Savage, Dennis Robert Ahrenhoerster, Prairie Seed Schenborn, Candy Schrank, Duane Source; David Cieslewicz, The Nature Schuettpelz, David Siebert, Kenneth Conservancy; Grant Cottam, UW- Sloan, Theodore Smith, Michael Madison; Thomas Crow, USDA-Forest Talbot, Donald Tills, William Service; Douglas County Forestry VanderZouwen, Karen Voss, Philip Committee; Nan Field, Writer; Wallace, Dreux Watermolen, Shane Thomas Frost, UW-Madison; Earl Weber, Cliff Wiita, Peter Wisdom, and Gustafson, Wisconsin Paper Council; Adrian Wydeven Alan Haney, UW-Stevens Point; Harold Jordahl, UW-Madison; Virginia Kline, UW-Madison Arboretum; We appreciate the time that a number of Limnology Graduate Seminar stu- leaders of industry, environmental, aca- dents, UW-Madison; Michael demic, and conservation groups spent in Luedeke, Burnett County Forest & sharing their ideas with us. These included: Parks; John Magnuson, UW-Madison; Rob Manes, Kansas Wildlife and James Baldock, Wisconsin Wildlife Parks; Jim Olson, Sierra Club; Neil Federation; David Cieslewicz, The Payne, UW-Stevens Point; Donald Nature Conservancy; Sharon Clark-

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Gaskill, Audubon Council; Noel Oneida County Forestry; Robert Cutright, Wisconsin Electric Power; Schmidt, Town of LeRoy; Robert Starr, Robert Ellingson, Wisconsin Conser- Town of LeRoy; John Staszcuk, Eau vation Congress; Earl Gustafson, Claire County Parks and Forest; Wisconsin Paper Council; Alan Haney, Robert Uschan, RUST Env. and UW-Stevens Point; Mark Heyde, Infrastructure; Mike Wharton, UW- Wisconsin County Forest Association, River Falls; and Judy Kircher (re- Marathon County Forestry Dept.; corder), WDNR-Eau Claire William Johnson, Governor’s Council on Forestry, Johnson Timber; Brian The dialogues and forums mentioned above Klatt, RUST Environment & Infra- went smoothly due to the expert help of structure; Walter Kuhlman, Environ- Department facilitators. These included: mental attorney, Boardman, Suhr, Curry & Field; John Magnuson, UW- Madison Limnology Laboratory; Paul Suzan Acre, David Birren, Larry Mikulak, Lake States Lumber Associa- Claggett, Linda Freitag, Carole Lee, tion, Kretz Lumber Company; Keith David Lentz, Tom Mickelson, Lynn Reopelle, Wisconsin Environmental Montgomery, Ed Nelson, Jordan Decade; and Caryl Terrell, Sierra Club Petchenik, Karen Rahmeir, Dennis Schenborn, Pat Sheahan, Al Stenstrup, Eric Thompson and Kristin Members of the general public were invited Visser to discussion sessions at meetings held in Milwaukee, Eau Claire, Rhinelander, and Madison in 1994. These participants A number of staff attended meetings in May helped add to our understanding of the and June 1994 to help develop ideas and issue and helped identify areas of common materials for presentations on biodiversity. agreement. Those who attended were: Those attending these meetings were:

Osh Andersen, Forestry Sciences Lab; Gerald Bever, Alan Crossley, Ronald D. Craig Beardsley, Nicolet National Eckstein, Glen Eveland, John Forest; Geoff Chandler, Nicolet Grosman, Tim Grunewald, Ralph National Forest; Jan Dahl, Izaak Hewett, Mark Heyde, Rebecca Walton League; Jim Dahl, Conserva- Isenring, Mary Judd, Kelly Kearns, tion Congress; Miriam Dahl, Izaak Ruth King, Lisa Kosmond, Robert Walton League; Andy Holschbach, Martini, Wendy McCown, Bruce Ozaukee County LCD; Michael Moss, William Mytton, John Nelson, Kaltenberg, UW-River Falls; Bart Dick Nikolai, John Olson, Mark Kotarba, Northwoods Wildlife Center; Schmidt, Craig Thompson, William Rick Koziel, Beaver Creek Reserve; VanderZouwen, Robert Wallen and Jim Lee, Wausau Daily Herald; Cy Charles Zosel Lyle, Audubon Society; Janette Marsh, USEPA; Dave Newhouse, Nicolet In addition to all of the above, many National Forest; Jim Olson, Sierra individuals shared thoughts at staff meet- Club; Joe Panci, Trees For Tomorrow; ings and public interest group meetings, Reggie Rensink, Wazee Park Comm; through phone calls, E-mail and Internet Tony Rinaldi, Nicolet National Forest; messages, personal conversations, and the Gene Roark, member of sharing of books and articles. Our sincere TNC,WWOA...; Russ Roberts, Geor- thanks to these individuals and all others gia Paciﬁc Corp.; Rick Rollman, who helped with clarifying the issue of biodiversity.

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