Florida panther recovery plan. Technical/agency draft.

              TECHNICAL/AGENCY DRAFT

FLORIDA PANTHER RECOVERY PLAN

(Puma concolor coryi)

THIRD REVISION

Original Approval: December 17, 1981

First Revision Approved: June 22, 1987

Second Revision Approved: March 13, 1995

Prepared by

The Florida Panther Recovery Team

and

South Florida Ecological Services Office

U.S. Fish and Wildlife Service

for

U.S. Fish and Wildlife Service

Southeast Region

Atlanta, Georgia

January 31, 2006 01/31/06 DRAFT Florida Panther Recovery Plan

DISCLAIMER

Recovery plans delineate actions which the best available science indicates are required to recover and protect listed species. Plans are published by the U.S. Fish and Wildlife Service (FWS), sometimes prepared with the assistance of recovery teams, contractors, State agencies, and others. Objectives will be attained and any necessary funds made available subject to budgetary and other constraints affecting the parties involved, as well as the need to address other priorities. Nothing in this plan should be construed as a commitment or requirement that any Federal agency obligate or pay funds in contravention of the Anti-Deficiency Act, 31 U.S.C. 1341, or any other law or regulation. Recovery plans do not necessarily represent the views or the official positions or approval of any individuals or agencies involved in the plan formulation, other than the FWS. They represent the official position of the FWS only after they have been signed by the Regional Director. Approved recovery plans are subject to modification as dictated by new findings, changes in species status, and the completion of recovery actions.

LITERATURE CITATION SHOULD READ AS FOLLOWS:

U.S. Fish and Wildlife Service. 2006. Technical/Agency Draft, Florida Panther Recovery Plan (Puma concolor coryi), Third Revision. U.S. Fish and Wildlife Service. Atlanta, Georgia. XXXpp.

ADDITIONAL COPIES MAY BE OBTAINED FROM:

U.S. Fish and Wildlife Service

1339 20th Street

Vero Beach, FL 32960

772-562-3909

Recovery plans can be downloaded from http://www.fws.gov/endangered or http://www.fws.gov/verobeach ii

01/31/06 DRAFT Florida Panther Recovery Plan

RECOVERY TEAM MEMBERS

Buddy Baker South Carolina Department of Natural Resources

Sonny Bass National Park Service/Everglades National Park

Chris Belden∗ U.S. Fish and Wildlife Service

Skip Bergmann U.S. Army Corps of Engineers

Debbie Blanco* Sarasota County Natural Resources

Dana Bryan* Florida Department of Environmental Protection

Mary Bryant The Nature Conservancy

Jimmy Bullock International Paper Company

Barbara Cintron U.S. Army Corps of Engineers

Joe Clark* U.S. Geological Survey, Biological Resources Division

Mark Cunningham* Florida Fish and Wildlife Conservation Commission

Don Cuozzo National Association of Home Builders

Kipp Frohlich* Florida Fish and Wildlife Conservation Commission

Skip Griep* U.S. Forest Service

Karen Gustin National Park Service/Big Cypress National Preserve

Dennis Hardin* Florida Division of Forestry

Deborah Jansen* National Park Service/Big Cypress National Preserve

Tom Jones Barron Collier Partnership

F. K. Jones Miccosukee Tribe of Indians of Florida

Nick Kapustin* Jacksonville Zoo

∗ Current members, alternates, and U.S. Fish and Wildlife Service participants who actively contributed by attending meetings.

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Robert Lacy Chicago Zoological Society

Darrell Land* Florida Fish and Wildlife Conservation Commission

Dwight LeBlanc U.S. Department of Agriculture, APHIS, Wildlife Services

Gary Lester Louisiana Department of Wildlife and Fisheries

Laurie Macdonald* Defenders of Wildlife

Dave Maehr University of Kentucky

Frank Mazzotti University of Florida

Roy McBride* Livestock Protection Company

Brian Murphy Quality Deer Management Association

Erin Myers* Natural Resources Conservation Service

Stephen O’Brien National Cancer Institute

Tim O’Meara* Florida Fish and Wildlife Conservation Commission

Jim Ozier Georgia Wildlife Resources Division

Pedro Ramos National Park Service/Big Cypress National Preserve

Richard Rummel Mississippi Department of Wildlife, Fisheries & Parks

Mark Sasser Alabama Division of Wildlife and Freshwater Fisheries

David Shindle* Conservancy of Southwest Florida

Mel Sunquist University of Florida

David Thompson White Oak Conservation Center

Steve Williams* Florida Panther Society

Ed Woods* Seminole Tribe of Florida

Wesley Woolf* National Wildlife Federation

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Recovery Team Member Alternates:

Phillip Brouse* Sarasota County Natural Resources

Monika Dey* U.S. Army Corps of Engineers

Elizabeth Fleming* Defenders of Wildlife

Margaret Griep* U.S. Forest Service

Sarah Grubs* Seminole Tribe of Florida

Laura Hartt* National Wildlife Federation

Karen Hill* Florida Panther Society

Jon Moulding* U.S. Army Corps of Engineers

Cynthia Ovdenk* U.S. Army Corps of Engineers

Mike Owen Florida Department of Environmental Protection

Nancy Payton Florida Wildlife Federation

U.S. Fish and Wildlife Service Participants:

Paula Halupa* South Florida Ecological Services Field Office

Layne Hamilton Florida Panther National Wildlife Refuge

Larry Richardson* Florida Panther National Wildlife Refuge

Cindy Schulz* South Florida Ecological Services Field Office

Paul Souza* South Florida Ecological Services Field Office

Meeting Facilitators – Florida Conflict Resolution Consortium:

Chris Pedersen Orlando

Tom Taylor Tallahassee

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Previous Recovery Team members that attended meetings were Lincoln Bormann (The Nature Conservancy), Pete David (South Florida Water Management District), Thomas Eason (Florida Fish and Wildlife Conservation Commission), John Kasbohm (U.S. Fish and Wildlife Service), Jeff Norment (Natural Resources Conservation Service), and Jora Young (The Nature Conservancy).

ACKNOWLEDGMENTS

The initial work (2001 - 2004) on this third revision of the Florida Panther Recovery Plan was led by John Kasbohm with the assistance of Dawn Jennings (U.S. Fish and Wildlife Service). Jora Young guided the Team through the threats analysis process and produced the Threats Analysis tables. Building upon that early work, Chris Belden and Cindy Schulz led the team through to completion of this revision.

Many people contributed to this revision, and some spent countless hours working on specific sections. The Overview and much of the Background Sections were initially written by John Kasbohm. Parts of the Background Section were updated and added to by Chris Belden, Mark Cunningham, Elizabeth Fleming, Paula Halupa, Laura Hartt, Karen Hill, Nick Kapustin, Darrell Land, Laurie Macdonald, Roy McBride, Tim O’Meara, Cindy Schulz, and Wes Woolf. The Recovery Strategy was drafted by Laura Hartt and Karen Hill with assistance from Larry Richardson, Wes Woolf, and Steve Williams. The Recovery Action Outline and Narrative Section and Implementation Schedule were a Team effort, but specific parts were provided by

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Kipp Frohlich, Margaret Griep, Tim O’Meara, and Karen Hill. Karen Hill provided the majority of the Public Awareness and Education parts of these sections.

The major editing for this revision was done by Cindy Schulz, Chris Belden, and Paula Halupa. Editorial suggestions were also provided by Laura Hartt, Deborah Jansen, Elizabeth Fleming, Karen Hill, Tim O’Meara, Joe Clark, Dana Bryan, Laurie Macdonald, and Mark Cunningham. We want to thank Chris Pederson and Tom Taylor for keeping us focused by facilitating our meetings.

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EXECUTIVE SUMMARY

Current Species Status

The Florida panther is the last subspecies of Puma still surviving in the eastern United States. Historically occurring throughout the southeastern United States, today the panther is restricted to less than 5% of its historic range in one breeding population of less than 100 animals, located in south Florida. The panther is threatened with extinction and human development in panther habitat negatively impacts recovery. The panther is federally listed as endangered under the Endangered Species Act of 1973, as amended (16 U.S.C. 1531 et seq.) and is on the State endangered lists for Florida, Georgia, Louisiana, and Mississippi.

Habitat Requirements and Limiting Factors

Panthers are wide ranging, secretive, and occur at low densities. They require large contiguous areas to meet their social, reproductive, and energetic needs. Panther habitat selection is related to prey availability (i.e., habitats that make prey vulnerable to stalking and capturing are selected). Dense understory vegetation provides some of the most important feeding, resting, and denning cover for panthers. Although daytime habitat use has been well documented for the panther, the species is predominantly active at night, and therefore a comprehensive understanding of panther habitat relationships has not yet been determined. Telemetry monitoring and ground tracking, however, indicate that panthers use the majority of habitats available to them.

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Limiting factors for the Florida panther are habitat availability, prey availability, and lack of human tolerance. Habitat loss, degradation, and fragmentation are among the greatest threats to panther survival, while lack of human tolerance is one of the greatest threats to panther recovery. Problems associated with being a single, small, isolated population and vehicle strikes have continued to keep the panther population at its current low numbers. Potential panther habitat throughout the Southeast continues to be affected by urbanization, residential development, conversion to agriculture and silviculture, mining and mineral exploration, and lack of land use planning that recognizes panther needs. Public opinion is critical to attainment of recovery goals and reintroduction efforts. Political and social issues will be the most difficult aspects of panther recovery and must be resolved before reintroduction efforts are initiated. A lack of public support and tolerance could prevent the reintroduction of panthers anywhere outside of south Florida.

Recovery Strategy

The strategy for Florida panther recovery sets an intermediate goal of downlisting from endangered to threatened with the ultimate goal of delisting. To achieve both the intermediate and ultimate goals, the recovery plan identifies three objectives which, collectively, describe the conditions necessary to achieve recovery. This plan presents objective, measurable criteria that when met would result in a determination that delisting is warranted. These criteria are based on population numbers and number of populations that provide for demographically and genetically viable populations as determined by several population viability analyses to ensure resilience to catastrophic events. The threats to the Florida panther will need to be addressed to attain these criteria.

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The recovery strategy for the Florida panther is to maintain, restore, and expand the panther population and its habitat in south Florida, expand this population into south-central Florida if sufficient habitat exists, reintroduce at least two additional viable populations within the historic range outside of south and south-central Florida, and facilitate panther recovery through public awareness and education. The panther depends upon habitat of sufficient quantity, quality, and spatial configuration for long-term persistence, therefore the plan is built upon habitat conservation and reducing habitat-related threats. Range expansion and reintroduction of additional populations are recognized as essential for panther recovery. Similarly, fostering greater public understanding and support is necessary to achieve panther conservation and recovery.

Recovery Goal

The goal of this revised recovery plan is to achieve long-term viability of the Florida panther to a point where it can be reclassified from endangered to threatened, and then removed from the list of endangered / threatened species.

Recovery Objectives

1. To maintain, restore, and expand the Florida panther population and its habitat in south Florida and, if feasible, expand the known occurrence of Florida panthers north of the Caloosahatchee River to maximize the probability of the long-term persistence of this metapopulation.

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2. To identify, secure, maintain, and restore habitat in potential reintroduction areas within the panther’s historic range, and to establish viable populations of the panther outside south and south-central Florida.

3. To facilitate panther conservation and recovery through public awareness and education.

Recovery Criteria

Downlisting should be considered when:

1. Two viable populations of at least 240 individuals (adults and subadults) each have been established and subsequently maintained for a minimum of fourteen years.

2. Sufficient habitat quality, quantity, and spatial configuration to support these populations is retained / protected or secured in the long-term.

A viable population, for purposes of Florida panther recovery, has been defined as one in which there is a 95% probability of persistence for 100 years. This population may be distributed in a metapopulation structure composed of subpopulations that total 240 individuals. There must be exchange of individuals and gene flow among subpopulations. For downlisting, exchange of individuals and gene flow can be either natural or through management. If managed, a commitment to such management must be formally documented and funded. Habitat should be in relatively unfragmented blocks that provide for food, shelter, and characteristic movements (e.g., hunting, breeding, dispersal, and territorial behavior) and support each metapopulation at a

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density of 2 to 3 animals per 100 square miles (259 square kilometers), resulting in a minimum of 8,000 – 12,000 square miles (20,720 – 31,080 square kilometers) per metapopulation of 240 panthers.

Delisting should be considered when:

1. Three viable, self-sustaining populations of at least 240 individuals (adults and subadults) each have been established and subsequently maintained for a minimum of fourteen years.

2. Sufficient habitat quality, quantity, and spatial configuration to support these populations is retained / protected or secured in the long-term.

For delisting, exchange of individuals and gene flow among subpopulations must be natural (i.e., not manipulated or managed).

Actions Needed

1. Maintain, restore, and expand the panther population and its habitat in south Florida.

2. Expand the known occurrence of panthers north of the Caloosahatchee River, if feasible.

3. Identify potential reintroduction areas within the historic range of the panther.

4. Reestablish viable panther populations outside of south and south-central Florida within the historic range.

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5. Secure, maintain, and restore habitat in reintroduction areas.

6. Facilitate panther conservation and recovery through public awareness and education.

Total Estimated Cost of Recovery

Cost estimates reflect costs for specific actions needed to achieve Florida panther recovery. Estimates do not include costs that agencies or other entities normally incur as part of their mission or normal operating expenses. The following table provides cost estimates for five years for recovery actions listed in the Implementation Schedule of this document. Costs for some recovery actions were not determinable; therefore, the total cost for recovery during this period is higher than this estimate.

Estimated Cost of Recovery for Five Years by Recovery Action Priority (Dollars x 1,000):

Year

Priority 1 Action

Priority 2 Actions

Priority 3 Actions

Total

1

875

1,981

1,713.5

4,569.5

2

875

1,696

1,506.5

4,077.5

3

835

1,561

1,231.5

3,627.5

4

835

921

981.5

2,737.5

5

835

921

981.5

2,737.5

Total

4,255

7,080

6,414.5

17,750

Date of Recovery

If all actions are fully funded and implemented as outlined, including full cooperation of all partners needed to achieve recovery, criteria for downlisting from endangered to threatened

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could be accomplished within 30 years; criteria for delisting could be accomplished within 45 years following reclassification.

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TABLE OF CONTENTS

DISCLAIMER..............................................................................................................................ii

RECOVERY TEAM MEMBERS.................................................................................................iii

ACKNOWLEDGMENTS..............................................................................................................vi

EXECUTIVE SUMMARY..........................................................................................................viii

Current Species Status........................................................................................................................viii

Habitat Requirements and Limiting Factors....................................................................................viii

Recovery Strategy.................................................................................................................................ix

Recovery Goal........................................................................................................................................x

Recovery Objectives..............................................................................................................................x

Recovery Criteria.................................................................................................................................xi

Actions Needed.....................................................................................................................................xii

Total Estimated Cost of Recovery......................................................................................................xiii

Date of Recovery.................................................................................................................................xiii

I. BACKGROUND.......................................................................................................................1

A. Overview...........................................................................................................................................4

B. Description........................................................................................................................................5

C. Taxonomy.........................................................................................................................................7

D. Population Trends and Distribution...............................................................................................8

E. Life History / Ecology.....................................................................................................................11

F. Habitat Characteristics / Ecosystem..............................................................................................21

G. Habitat and Prey Management.....................................................................................................27

H. Response to Management Activities.............................................................................................29

I. Reasons for Listing / Threats Assessment......................................................................................31

J. Conservation Efforts.......................................................................................................................56

K. Population Viability Analysis........................................................................................................74

II. RECOVERY STRATEGY......................................................................................................84

III. RECOVERY GOAL, OBJECTIVES, AND CRITERIA.....................................................91

Recovery Goal......................................................................................................................................91

Recovery Objectives............................................................................................................................91

Recovery Criteria................................................................................................................................92

A. Reclassification to Threatened.....................................................................................................................94

B. Delisting......................................................................................................................................................94 xv

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Existing Population..............................................................................................................................96

South Florida...................................................................................................................................................96

Expansion into South-Central Florida.........................................................................................................109

Reintroduction...................................................................................................................................111

Select Reintroduction Sites...........................................................................................................................111

Reintroduce Panthers into Suitable Sites.....................................................................................................112

Actions Once Populations Are Established..................................................................................................114

Public Awareness and Education......................................................................................................114

Design and Develop Materials and Programs.............................................................................................114

Provide Materials and Programs..................................................................................................................118

Evaluation......................................................................................................................................................120

V. IMPLEMENTATION SCHEDULE....................................................................................122

VI. LITERATURE CITED.......................................................................................................151

FIGURES..................................................................................................................................170

APPENDIX A. DEFINITIONS...............................................................................................176

APPENDIX B. THREATS ANALYSIS USING THE FIVE LISTING FACTORS..............178

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I. BACKGROUND

The Florida panther (Puma concolor coryi) was listed as endangered throughout its range in 1967 (32 FR 4001), and received Federal protection under the passage of the Endangered Species Act of 1973, as amended (16 U.S.C. 1531 et seq.) (ESA). Since the panther was designated as an endangered species prior to enactment of the ESA, there was no formal listing package identifying threats to the species as required by section 4(a)(1) of the ESA. No critical habitat has been designated for the panther.

The ESA establishes policies and procedures for identifying, listing, and protecting species of plants, fish, and wildlife that are endangered or threatened with extinction. The purposes of the ESA are “to provide a means whereby the ecosystems upon which endangered species and threatened species depend may be conserved, [and] to provide a program for the conservation of such endangered species and threatened species….” The ESA defines an “endangered species” as “any species which is in danger of extinction throughout all or a significant portion of its range.” A “threatened species” is defined as any species which is likely to become an endangered species within the foreseeable future throughout all or a significant portion of its range.” Under the definition of “species” in the ESA, the U.S. Fish and Wildlife Service (FWS) can apply the protections of the ESA to any species or subspecies of fish, wildlife, or plants, or any distinct population segment of any species of vertebrate fish or wildlife that meets the definition of endangered or threatened. The ESA does not attempt to define “species” in biological terms, and thus allows the term to be applied according to the best current biological knowledge and understanding of evolution, speciation, and genetics. So although the Florida

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panther is a subspecies, the protection it receives under the ESA is the same as for all other federally listed taxa whether they are species, subspecies, or distinct population segments.

The Secretary of the Department of the Interior is responsible for administering the ESA’s provisions as they apply to the Florida panther. Day-to-day management authority for endangered and threatened species under the Department’s jurisdiction has been delegated to the FWS. To help identify and guide species recovery needs, section 4(f) of the ESA directs the Secretary to develop and implement recovery plans for listed species or populations. Such plans are to include: (1) a description of site-specific management actions necessary to conserve the species; (2) objective, measurable criteria which, when met, will allow the species or populations to be removed from the endangered and threatened species list; and (3) estimates of the time and funding required to achieve the plan’s goals and intermediate steps. Section 4 of the ESA and regulations (50 CFR Part 424) promulgated to implement its listing provisions also set forth the procedures for reclassifying and delisting species on the Federal lists. A species can be delisted if the Secretary of the Interior determines that the species no longer meets the endangered or threatened status based upon these five factors listed in section 4(a)(1) of the ESA: (1) the present or threatened destruction, modification, or curtailment of its habitat or range; (2) overutilization for commercial, recreational, scientific, or educational purposes; (3) disease or predation; (4) the inadequacy of existing regulatory mechanisms; and (5) other natural or manmade factors affecting its continued existence.

Further, a species may be delisted, according to 50 CFR Part 424.11(d), if the best scientific and commercial data available substantiate that the species or population is neither endangered nor

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threatened for one of the following reasons: (1) extinction, (2) recovery, or (3) original data for classification of the species were in error.

The FWS has lead responsibility for conservation and recovery of the Florida panther, but all Federal agencies are responsible for contributing to panther conservation pursuant to section 7(a)(1) of the ESA. In 1981, FWS issued the initial recovery plan, and the plan was revisited in the mid-1980s culminating in the first major revision in 1987. A minor revision to incorporate a task to address genetic restoration and management was approved in 1995. In 1999, the FWS approved the South Florida Multi-species Recovery Plan (MSRP) (FWS 1999) that identified recovery needs of 68 threatened and endangered species in south Florida. The MSRP included recovery actions for the panther, but only for the portion of its range in south Florida. The FWS acknowledges that portions of the MSRP are now outdated and the habitat descriptions need to be clarified to more accurately describe panther habitat.

In 2001, the FWS initiated the process to revise the overall recovery plan for a third time. A new Florida Panther Recovery Team, consisting of representatives of the public, agencies, and groups that have an interest in panther recovery and / or could be affected by proposed actions, was established to assist with this revision.

Since approval of the original recovery plan in 1981 (FWS 1981), significant research has been conducted and important conservation and recovery activities have been accomplished primarily by the Florida Game and Freshwater Fish Commission (now the Florida Fish and Wildlife Conservation Commission [FWC]). This third revision of the recovery plan reflects many of

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those accomplishments, addresses current threats and needs, specifically addresses the planning requirements of the ESA, and supersedes previous recovery plans including the Florida panther component of the MSRP.

A. Overview

The Florida panther, a subspecies of Puma (also known as mountain lion, cougar, painter, or catamount) is the last subspecies still surviving in the eastern U.S. Historically occurring throughout the southeastern U.S., today the panther is restricted to less than 5% of its historic range in one breeding population of less than 100 animals, located in south Florida (Figure 1).

Persecution and prey decline resulted in a panther population threatened with extinction. Prior to 1949, panthers could be killed in Florida at any time of the year. In 1950, FWC declared the panther a regulated game species due to concerns over declining numbers. The FWC removed panthers from the game animal list in 1958 and gave them complete legal protection. On March 11, 1967, the FWS listed the panther as endangered (32 FR 4001) throughout its historic range. The Florida Panther Act (State Statute 372.671), a 1978 Florida State law, made killing a panther a felony. The Florida panther is listed as endangered by the States of Florida, Georgia, Louisiana, and Mississippi.

FWS uses recovery priority numbers, ranging from a high of 1C to a low of 18, to assign recovery priorities to listed species. The criteria on which the recovery priority number is based are degree of threat, recovery potential, taxonomic distinctiveness, and presence of an actual or imminent conflict between the species and development activities. The FWS has assigned the

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panther a recovery priority number of 6C. This priority number identifies the panther as a subspecies with a high degree of threat of extinction, but low recovery potential because recovery is in conflict with construction, other development projects, or other forms of economic activity.

Presently, habitat loss and fragmentation, problems associated with small isolated populations including inbreeding depression and reduced genetic health, and vehicle strikes have continued to keep the panther population at its currently low numbers. Today, survival and recovery of the Florida panther are dependent upon maintaining, restoring, and expanding the panther population and its habitat in south Florida and facilitating panther conservation and recovery through public awareness and education. In addition, recovery also requires expanding the known occurrence of panthers north of the Caloosahatchee River, if feasible, and identifying potential reintroduction areas within the historic range and establishing and maintaining at least two additional viable populations with associated habitats outside of south and south-central Florida.

B. Description

An adult Florida panther is unspotted and typically rusty reddish-brown on the back, tawny on the sides, and pale gray underneath. There has never been a melanistic (black) puma documented in North America (Tinsley 1970, 1987). Adult males can reach a length of seven feet (ft) (2.1 meters [m]) from their nose to the tip of their tail and may exceed 161 pounds (lbs) (73 kilograms [kg]) in weight; but, typically adult males average around 116 lbs (52.6 kg) and stand approximately 24 - 28 inches (in) (60 - 70 centimeters [cm]) at the shoulder (Roelke 1990). Female panthers are smaller with an average weight of 75 lbs (34 kg) and length of 6 ft (1.8 m) 5

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(Roelke 1990). The skull of the Florida panther is unique in that it has a broad, flat, frontal region, and broad, high-arched or upward-expanded nasal bones (Young and Goldman 1946).

Florida panther kittens are gray with dark brown or blackish spots and five bands around the tail. The spots gradually fade as the kittens grow older and are almost unnoticeable by the time they are six months old. At this age, their bright blue eyes slowly turn to the light-brown straw color of the adult (Belden 1988).

Three external characters—a right angle crook at the terminal end of the tail, a whorl of hair or cowlick in the middle of the back, and irregular, white flecking on the head, nape, and shoulders—not found in combination in other subspecies of Puma (Belden 1986), were commonly observed in Florida panthers through the mid-1990s. The kinked tail and cowlicks were considered manifestations of inbreeding (Seal 1994a), whereas the white flecking was thought to be a result of scarring from tick bites (Maehr 1992, Wilkins et al. 1997). Four other abnormalities prevalent in the panther population prior to the mid-1990s included cryptorchidism (one or two undescended testicles), low sperm quality, atrial septal (the opening between two atria fails to close normally during fetal development) defects, and immune deficiencies and were also suspected to be the result of low genetic variability (Roelke et al. 1993b).

A plan for genetic restoration and management of the Florida panther was developed in September 1994 (Seal 1994a) and eight non-pregnant adult female Texas panthers (Puma concolor stanleyana) were released in five areas of south Florida from March to July 1995. Since this introgression, rates of genetic defects, including crooked tails and cowlicks, have

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dramatically decreased (Land et al. 2004). In addition, to date neither atrial septal defects nor cryptorchidism have been found in introgressed panthers (M. Cunningham, FWC, pers. comm. 2005).

C. Taxonomy

The Florida panther was first described by Charles B. Cory in 1896 as Felis concolor floridana (Cory 1896). The type specimen was collected in Sebastian, Florida. Bangs (1899), however, believed that the Florida panther was restricted to peninsular Florida and could not intergrade with other Felis spp. Therefore, he assigned it full specific status and named it Felis coryi since Felis floridana had been used previously for a bobcat (Lynx rufus).

The taxonomic classification of the Felis concolor group was revised and described by Nelson and Goldman (1929) and Young and Goldman (1946). These authors differentiated 30 subspecies using geographic and morphometric (measurement of forms) criteria and reassigned the Florida panther to subspecific status as Felis concolor coryi. This designation also incorporated F. arundivaga which had been classified by Hollister (1911) from specimens collected in Louisiana into F. c. coryi. Nowell and Jackson (1996) reviewed the genus Felis and placed mountain lions, including the Florida panther, in the genus Puma.

Culver et al. (2000) examined genetic diversity within and among the described subspecies of Puma concolor using three groups of genetic markers and proposed a revision of the genus to include only six subspecies, one of which encompassed all Puma in North America including the Florida panther. However, Culver et al. (2000) determined that the Florida panther was one of

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several smaller populations that had unique features, the number of polymorphic microsatellite loci and amount of variation were lower, and it was highly inbred (eight fixed loci). The degree to which the scientific community has accepted the results of Culver et al. (2000) and the proposed change in taxonomy is not resolved at this time. The Florida panther remains listed as a subspecies and continues to receive protection pursuant to the ESA.

D. Population Trends and Distribution

The Florida panther once ranged throughout the southeastern U.S. from Arkansas and Louisiana eastward across Mississippi, Alabama, Georgia, Florida, and parts of South Carolina and Tennessee (Young and Goldman 1946) (Figure 1). Historically, the panther intergraded to the north with P. c. cougar, to the west with P. c. stanleyana, and to the northwest with P. c. hippolestes (Young and Goldman 1946).

Although generally considered unreliable, sightings of panthers regularly occur throughout the Southeast. However, no reproducing populations of panthers have been found outside of south Florida for at least 30 years despite intensive searches to document them (Belden et al. 1991, McBride et al. 1993, Clark et al. 2002). Survey reports and more than 70,000 locations of radio-collared panthers recorded between 1981 and 2004 clearly define the panther’s current breeding range (Figure 1). Reproduction is known only in the Big Cypress Swamp / Everglades physiographic region in Collier, Lee, Hendry, Dade, and Monroe Counties south of the Caloosahatchee River (Belden et al. 1991). Although confirmed panther sign, male radio-collared panthers, and uncollared males killed by vehicles have been recorded outside of south Florida in recent years, no female panthers have been documented north of the Caloosahatchee

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River since 1973 (Nowak and McBride 1974, Belden et al. 1991, Land and Taylor 1998, Land et al. 1999, Shindle et al. 2000, McBride 2002, Belden and McBride 2005).

Puma are wide ranging, secretive, and occur at low densities. However, their tracks, urine markers, and scats are readily found by trained observers, and resident populations are easily located. Van Dyke (1986b) determined that all resident puma, 78% of transient puma, and 57% of kittens could be detected by track searches in Utah. In south Florida, the Florida panther’s limited range and low densities may make the population count derived from track searches more accurate than in Utah. During two month-long investigations – one late in 1972 and early 1973 and another in 1974 – funded by the World Wildlife Fund to determine if panthers still existed in Florida, McBride searched for signs of panthers in portions of south Florida. In 1972, McBride authenticated a road-killed male panther in Glades County and a female captured and released from a bobcat trap in Collier County (R. McBride, Livestock Protection Company, pers. comm. 2005). In 1973, McBride captured one female in Glades County (Nowak and McBride 1974). Based on this preliminary evidence, Nowak and McBride (1974) estimated the “population from the Lake Okeechobee area southward to be about 20 or 30 individuals.” In 1974, McBride found evidence of only two additional panthers in the Fakahatchee Strand and suggested that “there could be not more than ten individual panthers in the area around Lake Okeechobee and southward in the state” (Nowak and McBride 1975). This initial survey, while brief in nature, proved that panthers still existed in Florida and delineated areas where a more exhaustive search was warranted. After this initial investigation, more comprehensive surveys on both public and private lands were completed (Reeves 1978; Belden and McBride 1983a, b; Belden et al. 1991).

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Thirty individual panthers were identified during a wide-ranging survey in 1985 in south Florida (McBride 1985).

Maehr et al. (1991a) provides the only published population estimate based on a substantial body of field data (Beier et al. 2003). Maehr et al. (1991a) estimated a density of 1 panther / 43 mi2 [110 square kilometers (km2)] based on 17 concurrently radiocollared and four uncollared panthers. They extrapolated this density to the area occupied (1,946 mi2 [5,040 km2]) by radio-collared panthers during the period 1985 - 1990 to achieve a population estimate of 46 adult panthers for southwest Florida (excluding Everglades National Park [ENP], eastern Big Cypress National Preserve [BCNP], and Glades and Highlands Counties). Beier et al. (2003), however, argued that this estimate of density, although “reasonably rigorous,” could not be extrapolated to other areas because it was not known whether densities were comparable in those areas.

More recently, McBride (2000, 2001, 2002, 2003) obtained minimum population counts (i.e., number known alive) based on panthers treed with hounds, physical evidence (e.g., tracks where radio-collared panthers were not known to occur), documentation by trail-camera photos, and sightings of uncollared panthers by a biologist or pilot from a monitoring plane or via ground telemetry. He counted 62, 78, 80, and 87 panthers (which includes adult and subadult panthers but not kittens at the den) in 2000, 2001, 2002, and 2003, respectively. Population counts were not conducted in 2004 and are not yet available for 2005.

McBride (2002) documented an increase in the number of uncollared panthers captured each year between 2000 and 2002 relative to 1981 through 1999, while Shindle et al. (2001) reported

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data showing an apparent increase in the number of panthers killed by vehicles since 1999. In neither case, however, was supporting evidence provided that would be necessary to conclude that these increases reflected population trends.

Although the breeding segment of the panther population occurs only in south Florida, panthers have been documented north of the Caloosahatchee River over 125 times since February 1972. This has been confirmed through field sign (e.g., tracks, scrapes, scats), camera-trap photographs, seven highway mortalities, four radio-collared animals, two captured animals (one of which was radiocollared), and one skeleton. From 1972 through 2004, panthers have been confirmed in 11 counties (Flagler, Glades, Highlands, Hillsborough, Indian River, Okeechobee, Orange, Osceola, Polk, Sarasota, Volusia) north of the river (Belden et al. 1991, Belden and McBride 2005). However, no evidence of a female or reproduction has been documented north of the Caloosahatchee River in over 30 years (Belden and McBride 2005).

E. Life History / Ecology

Reproduction--Male Florida panthers are polygynous, maintaining large, overlapping home ranges containing several adult females and their dependent offspring. The first sexual encounters for males normally occur at about three years based on 26 radio-collared panthers of both sexes (Maehr et al. 1991a). Based on genetics work, some males may become breeders as early as 17 months (W. Johnson, National Cancer Institute, pers. comm. 2005). Breeding activity peaks from December to March (Shindle et al. 2003). Litters (n = 82) are produced throughout the year, with 56 - 60% of births occurring between March and June (Jansen et al. 2005, Lotz et al. 2005). The greatest number of births occur in May and June (Jansen et al. 2005,

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Lotz et al. 2005). Female panthers have bred as young as 18 months (Maehr et al. 1989a) and successful reproduction has occurred up to 11 years old. Mean age of denning females is 4.6 ± 2.1 (standard deviation [sd]) years (Lotz et al. 2005). Age at first reproduction for 19 known-aged female panthers averaged 2.2 ± 0.246 (sd) years and ranged from 1.8 - 3.2 years. Average litter size is 2.4 ± 0.91 (sd) kittens. Seventy percent of litters are comprised of either two or three kittens. Mean birth intervals (elapsed time between successive litters) are 19.8 ± 9.0 (sd) months for female panthers (n = 56) (range 4.1 - 36.5 months) (Lotz et al. 2005). Females that lose their litters generally produce another more quickly; five of seven females whose kittens were brought into captivity (see Captive Breeding section of F. Conservation Efforts) successfully produced another litter an average of 10.4 months after the removal of the initial litter (Land 1994).

Den sites are usually located in dense, understory vegetation, typically saw palmetto (Serenoa repens) (Maehr 1990a, Shindle et al. 2003). Den sites are used for up to two months by female panthers and their litters from birth to weaning. Independence and dispersal of young typically occurs at 18 months, but may occur as early as one year (Maehr 1992).

Survivorship and Causes of Mortality--Intraspecific aggression accounts for 42% of all mortalities among radio-collared panthers (Jansen et al. 2005, Lotz et al. 2005). Unknown causes and collisions with vehicles account for 24 and 19% of mortalities, respectively. From 1990 - 2004, mean annual survivorship of radio-collared adult panthers was greater for females (0.894 ± 0.099 sd) than males (0.779 ± 0.125 sd) (Lotz et al. 2005). Most intraspecific aggression occurs between male panthers; but, aggressive encounters between males and

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females, resulting in the death of the female, have occurred. Defense of kittens and / or a kill is suspected in half (5 of 10) of the known instances through 2003 (Shindle et al. 2003).

Female panthers are considered adult residents if they are older than 18 months, have established home ranges and bred (Maehr et al. 1991a). Land et al. (2005) reported that all 24 female panthers first captured as kittens survived to become residents and 19 (79.2%) produced litters. Male panthers are considered adult residents if they are older than three years and have established a home range that overlaps with females. Thirty-one male panthers were captured as kittens and 12 (38.7%) of these cats survived to become residents (Jansen et al. 2005, Lotz et al. 2005). “Successful male recruitment appears to depend on the death or home-range shift of a resident adult male” (Maehr et al. 1991a). Turnover in the breeding population is low with documented mortality in radio-collared panthers being greatest in subadults and non-resident males (Maehr et al. 1991a, Shindle et al. 2003).

Den sites of female Florida panthers and Texas puma females have been visited since 1992 and the number of kittens that survived to six months for 38 of these litters has been documented. Florida and introgressed panther kitten survival to six months were estimated to be 52 and 72%, respectively, but were not significantly different (P = 0.2776) (Lotz et al. 2005). Survival of kittens greater than six months old was determined by following the fates of 55 radio-collared dependent-aged kittens, including 17 introgressed panthers from 1985 - 2004. Only one of these 55 kittens died before reaching independence, resulting in a 98.2% survival rate (Lotz et al. 2005). The FWC and NPS are continuing to compile and analyze existing reproductive and kitten data.

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Dispersal--Panther dispersal begins after a juvenile becomes independent from its mother and continues until it establishes a home range. Dispersal distances are greater for males (n = 18) than females (n = 9) (42.5 mi [68.4 km] vs. 12.6 mi [20.3 km], respectively) and the maximum dispersal distance recorded for a young male was 139.2 mi (224.1 km) over a seven-month period followed by a secondary dispersal of 145 mi (233 km) (Maehr et al. 2002a). Males disperse an average distance of 25 mi (40 km); females typically remain in or disperse short distances from their natal ranges (Comiskey et al. 2002). Female dispersers are considered philopatric because they usually establish home ranges less than one average home range width from their natal range (Maehr et al. 2002a). Maehr et al. (2002a) reported that all female dispersers (n = 9) were successful at establishing a home range whereas only 63% of males (n = 18) were successful. Young panthers become independent at 14 months on average for both sexes, but male dispersals are longer in duration than for females (9.6 months and 7.0 months, respectively) (Maehr et al. 2002a). Dispersing males usually go through a period as transient (non-resident) subadults, moving through the fringes of the resident population and often occupying suboptimal habitat until an established range becomes vacant (Maehr 1997a).

Most panther dispersal occurs south of the Caloosahatchee River with only four radio-collared panthers crossing the river and continuing north since 1981 (Land and Taylor 1998, Land et al. 1999, Shindle et al. 2000, Maehr et al. 2002a, Belden and McBride 2005). Western subspecies of Puma have been documented crossing wide, swift-flowing rivers up to a mile in width (Seidensticker et al. 1973, Anderson 1983). The Caloosahatchee River, a narrow (295 - 328 ft [90 - 100 m]), channelized river, probably is not a significant barrier to panther movements, but

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the combination of the river, State Route (SR) 80, and land uses along the river seems to have restricted panther dispersal northward (Maehr et al. 2002a). Documented physical evidence of at least 15 other uncollared male panthers have been confirmed north of the river since 1972, but no female panthers nor reproduction have been documented in this area since 1973 (Belden and McBride 2005).

Home Range Dynamics and Movements--Panthers require large areas to meet their needs. Numerous factors influence panther home range size including habitat quality, prey density, and landscape configuration (Belden 1988, Comiskey et al. 2002). Home range sizes of 26 radio-collared panthers monitored between 1985 and 1990 averaged 200 mi2 (519km2) for resident adult males and 75 mi2 (193 km2) for resident adult females; transient males had a home range of 240 mi2 (623 km2) (Maehr et al. 1991a). Comiskey et al. (2002) examined the home range size for 50 adult panthers (residents greater than 1.5 years old) monitored in south Florida from 1981 - 2000 and found resident males had a mean home range of 251 mi2 (650 km2) and females had a mean home range of 153 mi2 (396 km2). Beier et al. (2003) found home range size estimates for panthers reported by Maehr et al. (1991a) and Comiskey et al. (2002) to be reliable.

Annual minimum convex polygon home range sizes of 52 adult radio-collared panthers monitored between 1998 and 2002 ranged from 24 - 459 mi2 (63 - 1,188 km2), averaging 140 mi2 (362 km2) for 20 resident adult males and 69 mi2 (179 km2) for 32 resident adult females (Land et al. 1999, Shindle et al. 2000, Shindle et al. 2001, Land et al. 2002). Home ranges of resident adults tend to be stable unless influenced by the death of other residents, however, several males have shown significant home range shifts that may be related to aging (D. Jansen, National Park

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Service [NPS], pers. comm. 2005). Home-range overlap is extensive among resident females and limited among resident males (Maehr et al. 1991a).

Activity levels for Florida panthers are greatest at night with peaks around sunrise and after sunset (Maehr et al. 1990a). The lowest activity levels occur during the middle of the day. Female panthers at natal dens follow a similar pattern with less difference between high and low activity periods.

Telemetry data indicate that panthers typically do not return to the same resting site day after day, with the exception of females with dens or panthers remaining near kill sites for several days. The presence of physical evidence such as tracks, scats, and urine markers confirm that panthers move extensively within home ranges, visiting all parts of the range regularly in the course of hunting, breeding, and other activities (Maehr 1997a, Comiskey et al. 2002). Males travel widely throughout their home ranges to maintain exclusive breeding rights to females. Females without kittens also move extensively within their ranges (Maehr 1997a). Panthers are capable of moving large distances in short periods of time. Nightly panther movements of 12 mi (20 km) are not uncommon (Maehr et al. 1990a).

Intraspecific Interactions--Interactions between panthers occur indirectly through urine markers or directly through contact. Urine markers are made by piling ground litter using a backwards-pushing motion with the hind feet. This pile is then scent-marked with urine and occasionally feces. Both sexes make urine markers, apparently males use them as a way to mark their territory and announce presence while females advertise their reproductive condition.

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Adult females and their kittens interact more frequently than any other group of panthers. Interactions between adult male and female panthers last from one to seven days and usually result in pregnancy (Maehr et al. 1991a). Aggressive interactions between males often result in serious injury or death. Independent subadult males have been known to associate with each other for several days and these interactions do not appear to be aggressive in nature. Aggression between males is the most common cause of male mortality and an important determinant of male spatial and recruitment patterns based on radio-collared panthers (Maehr et al. 1991a, Shindle et al. 2003). Aggressive encounters between radio-collared males and females also have been documented (Shindle et al. 2003, Jansen et al. 2005).

Food Habits--Primary panther prey are white-tailed deer (Odocoileus virginianus) and feral hog (Sus scrofa) (Maehr et al. 1990b, Dalrymple and Bass 1996). Generally, feral hogs constitute the greatest biomass consumed by panthers north of the Alligator Alley section of Interstate 75 (I-75) while white-tailed deer are the greatest biomass consumed to the south (Maehr et al. 1990b). Secondary prey includes raccoons (Procyon lotor), nine-banded armadillos (Dasypus novemcinctus), marsh rabbits (Sylvilagus palustris) (Maehr et al. 1990b) and alligators (Alligator mississippiensis) (Dalrymple and Bass 1996). No seasonal variation in diet has been detected. A resident adult male puma generally consumes one deer-sized prey every 8 - 11 days; this frequency would be 14 - 17 days for a resident female; and 3.3 days for a female with three 13-month-old kittens (Ackerman et al. 1986). Maehr et al. (1990b) documented domestic livestock infrequently in scats or kills, although cattle were readily available on their study area.

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Infectious Diseases, Parasites, and Environmental Contaminants--

Viral Diseases--Feline leukemia virus (FeLV) is common in domestic cats (Felis catus), but is quite rare in non-domestic felids. Routine testing for FeLV antigen (indicating active infection) in captured and necropsied panthers has been negative since testing began in 1978 to the fall of 2002. Between November 2002 and February 2003, however, two panthers tested FeLV antigen positive (Cunningham 2005). The following year, three more cases were diagnosed. All infected panthers had overlapping home ranges in the Okaloacoochee Slough ecosystem. Three panthers died due to suspected FeLV-related diseases (opportunistic bacterial infections and anemia) and the two others died from intraspecific aggression. Testing of serum samples collected from 1990 - 2005 for antibodies (indicating exposure) to FeLV indicated increasing exposure to FeLV beginning in the late 1990s and concentrated north of I-75. There was apparently minimal exposure to FeLV during this period south of I-75. Positive antibody titers in different areas at different times may indicate that multiple introductions of the virus into the panther population may have occurred. These smaller epizootics were apparently self-limiting and did not result in any known mortalities. Positive antibody titers, in the absence of an active infection (antigen positive), indicate that panthers can be exposed and overcome the infection (Cunningham 2005). Management of the disease includes vaccination as well as removal of infected panthers to captivity for quarantine and supportive care. As of June 1, 2005, approximately one-third of the population had received at least one vaccination against FeLV (FWC and NPS, unpublished data). No new positive cases have been diagnosed since July 2004.

Pseudorabies virus (PRV) (Aujeszky’s disease) causes respiratory and reproductive disorders in adult hogs and mortality in neonates, but is a rapidly fatal neurologic disease in carnivores. At

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least one panther died from PRV infection presumably through consumption of an infected feral hog (Glass et al. 1994). At least one panther has also died of rabies (Taylor et al. 2002). This panther was radiocollared but not vaccinated against the disease.

Feline immunodeficiency virus (FIV) is a retrovirus of felids that is endemic in the panther population. Approximately 28% of Florida panthers were positive for antibodies to the puma lentivirus strain of FIV (Olmstead et al. 1992); however, the prevalence may be increasing. Between November 2004 and April 2005, 13 of 17 (76%) were positive (M. Cunningham, FWC, unpublished data). The cause of this increase is unknown but warrants continued monitoring and investigation. There is also evidence of exposure to Feline panleukopenia virus (PLV) in adult panthers (Roelke et al. 1993a) although no PLV-related mortalities are known to have occurred.

Serological evidence of other viral diseases in the panther population includes feline calicivirus, feline herpes virus, and West Nile virus (WNV). However these diseases are not believed to cause significant morbidity or mortality in the population. All panthers found dead due to unknown causes are tested for alphaviruses, flaviviruses (including WNV), and canine distemper virus. These viruses have not been detected in panthers by viral culture or polymerase chain reaction (FWC, unpublished data).

Other Infectious Diseases--Bacteria have played a role in free-ranging panther morbidity and mortality as opportunistic pathogens, taking advantage of pre-existing trauma or FeLV infections (FWC, unpublished data). Dermatophytosis (ringworm infection) has been diagnosed in several panthers and resulted in severe generalized infection in at least one (Rotstein et al. 1999). Severe

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infections may reflect an underlying immunocompromise, possibly resulting from inbreeding depression or immunosuppressive viral infections.

Parasites--The hookworm, Ancylostoma pluridentatum, is found in a high prevalence in the panther population. Other parasites identified from live-captured or necropsied panthers include eight arthropod species, eight nematode species, three cestode species, two trematode species, and three protozoa species (Forrester et al. 1985, Forrester 1992, Wehinger et al. 1995, Rotstein et al. 1999, Land et al. 2002). Of these only an arthropod, Notoedres felis, caused significant morbidity in at least one panther (Maehr et al. 1995).

Environmental Contaminants--Overall, mercury in south Florida biota has decreased over the last several years (Frederick et al. 2002). However, high mercury concentrations are still found in some panthers. At least one panther is thought to have died of mercury toxicosis and mercury has been implicated in the death of two other panthers in ENP (Roelke 1991). One individual panther had concentrations of 150 parts per million (ppm) mercury in its hair (Land et al. 2004). Elevated levels of p, p’– DDE were also detected in fat from that panther. The role of mercury and / or p, p’– DDE in this panther’s death is unknown and no cause of death was determined despite extensive diagnostic testing. Elevated mercury concentrations have also been found in panthers from Florida Panther National Wildlife Refuge (FPNWR). Two sibling neonatal kittens from this area had hair mercury concentrations of 35 and 40 ppm. Although other factors were believed to have been responsible, these kittens did not survive to leave their natal den. Consistently high hair mercury values in ENP and FPNWR and the finding of elevated values in some portions of BCNP warrant continued monitoring (Land et al. 2004). Other environmental

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contaminants found in panthers include polychlorinated biphenyls (Arochlor 1260) and organochlorines (p, p’–DDE) (Dunbar 1995, Land et al. 2004).

F. Habitat Characteristics / Ecosystem

Landscape Composition--Noss and Cooperrider (1994) considered the landscape implications of maintaining viable panther populations. Assuming a male home range size of 215 mi 2 (558 km2) (Maehr 1990a), an adult sex ratio of 50:50 (Anderson 1983), and some margin of safety, they determined that a reserve network as large as 15,625 – 23,438 mi2 (40,469 - 60,703 km2) would be needed to support an effective population size of 50 individuals (equating to an actual adult population of 100 - 200 panthers [Ballou et al. 1989]). However, to provide for long-term persistence based on an effective population size of 500 individuals (equating to 1,000 - 2,000 adult panthers [Ballou et al. 1989]), could require as much as 156,251 - 234,376 mi2 (404,687 - 607,031 km2). This latter acreage corresponds to roughly 60 - 70% of the Florida panther’s historical range. Although it is uncertain whether this much land is needed for panther recovery, it does provide some qualitative insight into the importance of habitat conservation across large landscapes for achieving a viable panther population (Noss and Cooperrider 1994).

Between 1981 and 2003, more than 55,000 locations on more than 100 radio-collared panthers were collected. Belden et al. (1988), Maehr et al. (1991a), Maehr (1997a), Kerkoff et al. (2000), and Comiskey et al. (2002) provide information on habitat use based on various subsets of these data. Since almost all data from radio-collars have been collected during daytime hours (generally 0700 - 1100), and because panthers are most active at night (Maehr et al. 1990a),

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daytime radio locations are insufficient to describe the full range of panther habitat use (Beyer and Haufler 1994, Comiskey et al. 2002, Beier et al. 2003, Dickson et al. 2005, Beier et al. in press).

The FWS created the Multi-species/Ecosystem Recovery Implementation Team (MERIT) to assist with implementation of the MSRP after it was signed in 1999. The Florida Panther Subteam of MERIT developed a landscape-level strategy for the conservation of the panther population in south Florida which was not finalized. Many of the Panther Subteam members refined the methodology, further analyzed the data, and better defined the results of this landscape-level strategy into a spatial model (Kautz et al. in press). In developing the model, data from radio-collared panthers collected from 1981 through 2000 were used to evaluate the relative importance of various land cover types as panther habitat, thus identifying landscape components important for panther habitat conservation. Those components were then combined with a least cost path analysis to delineate three panther habitat conservation zones for south Florida: (1) Primary Zone – lands essential to the long-term viability and persistence of the panther in the wild; (2) Secondary Zone - lands which few panthers use contiguous with the Primary Zone, but given sufficient habitat restoration could accommodate expansion of the panther population south of the Caloosahatchee River; and (3) Dispersal Zone - the area which may facilitate future panther expansion north of the Caloosahatchee River (Kautz et al. in press) (Figure 3). The Primary Zone is currently occupied and supports the breeding population of panthers. Although panthers move through the Secondary and Dispersal Zones, they are not permanently occupied. The Secondary Zone could support panthers with sufficient restoration.

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These zones vary in size, ownership, and land cover composition. The Primary Zone is 3,548 mi2 (9,189 km2) in size, 73% of which is publicly owned (R. Kautz, Dennis, Breedlove, and Associates, pers. comm. 2005), and includes portions of the BCNP, ENP, Fakahatchee Strand Preserve State Park (FSPSP), FPNWR, Okaloacoochee Slough State Forest, and Picayune Strand State Forest. This zone’s composition is 45% forest, 41% freshwater marsh, 7.6% agriculture lands, 2.6% prairie and shrub lands, and 0.52% urban lands (Kautz et al. in press).

The Secondary Zone is 1,269 mi2 (3,287 km2) in size, 38% of which is public land (R. Kautz, pers. comm. 2005). This zone’s composition is 43% freshwater marsh, 36% agriculture, 11% forest, 6.1% prairie and shrub lands, and 2.3% low-density residential areas and open urban lands (Kautz et al. in press).

The Dispersal Zone is 44 mi2 (113 km2) in size, all of which is privately owned (R. Kautz, pers. comm. 2005). This zone’s composition is 49% agriculture (primarily improved pasture and citrus groves), 29% forest (wetland and upland), 8.8% prairie and shrub land, 7.5% freshwater marsh, and 5.1% barren and urban lands (Kautz et al. in press).

Diurnal Habitat Use--Diurnal panther locations appear to be within or closer to forested cover types, particularly cypress swamp, pinelands, hardwood swamp, and upland hardwood forests (Belden 1986, Belden et al. 1988, Maehr 1990a, Maehr et al. 1991a, Maehr 1992, Smith and Bass 1994, Kerkhoff et al. 2000, Comiskey et al. 2002). Day beds and denning sites are important habitat features that should be considered in panther management. Dense understory

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vegetation comprised of saw palmetto provides some of the most important resting and denning cover for panthers (Maehr 1990a). Shindle et al. (2003) show that 73% of panther dens were in palmetto thickets.

Radio-collar data and ground tracking indicate that panthers use the mosaic of habitats available to them as resting and denning sites, hunting grounds, and travel routes. These habitats include cypress swamps, hardwood hammocks, pine flatwoods, seasonally flooded prairies, freshwater marshes, and some agricultural lands. Although radiocollar monitoring indicates that forest is a preferred cover type for daytime rest, it is clear that panthers utilize non-forest cover types, including areas disturbed by humans (e.g., Belden et al. 1988, Maehr et al. 1991a, Comiskey et al. 2002). Compositional analyses by Kautz et al. (in press) confirmed previous findings that forest patches comprise an important component of panther habitat in south Florida, but that other natural and disturbed cover types are also present in the large landscapes that support panthers (Belden et al. 1988, Maehr et al. 1991a, Comiskey et al. 2002). Kautz et al. (in press) found that the smallest class of forest patches (i.e., 9 - 26 ac [3.6 - 10.4 ha]) were the highest ranked forest patch sizes within panther home ranges; this indicates that forest patches of all sizes appear to be important components of the landscapes inhabited by panthers, not just the larger forest patches.

Nocturnal Habitat Use--Maehr et al. (1990a) provide the only descriptions of panther nocturnal activities and represent the available radiocollar data collected during night time hours. Unfortunately, this paper does not provide analyses of nocturnal habitat use. Dickson et al. (2005) examined the movements of 10 female and seven male puma at 15-minute intervals

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during 44 nocturnal periods of hunting or traveling in southern California. They found that traveling puma monitored over nocturnal periods used a broader range of habitats than what they appeared to use based on diurnal locations alone. These findings support the argument of Comiskey et al. (2002) that analyses of diurnal locations provide limited information about puma patterns of habitat selection during the time of day when puma are most active (Dickson et al. 2005). The use of Global Positioning System (GPS) radiocollars is now being investigated to determine if this technology will be suitable to answer questions regarding Florida panther nocturnal habitat use.

Prey Habitat Use--Panther habitat selection is related to prey availability (Janis and Clark 1999, Dees et al. 2001) and, consequently, prey habitat use. Adequate cover and the size, distribution, and abundance of available prey species are critical factors to the persistence of panthers in south Florida and often determine the extent of panther use of an area. Duever (1986) calculated a deer population of 1,760 in BCNP, based on Harlow (1959) deer density estimates of 1 / 210 ac (85 ha) in pine forest, 1 / 299 ac (121 ha) in swamps, 1 / 1,280 ac (518 ha) in prairie, 1 / 250 ac (101 ha) in marshes, and 1 / 111 ac (45 ha) in hammocks. Schortemeyer et al (1991) estimated deer densities at 1 / 49 - 247 ac (20 - 100) ha in three management units of BCNP based on track counts and aerial surveys. Labisky et al. (1995) reported 1 / 49 ac (20 ha) in southeastern BCNP. Using track counts alone, McCown (1994) estimated 1 / 183 - 225 ac (74 - 91 ha) on the FPNWR and 1 / 133 - 200 ac (54 - 81 ha) in the FSPSP.

Hardwood hammocks and other forest cover types are important habitat for white-tailed deer and other panther prey (Harlow and Jones 1965, Belden et al. 1988, Maehr 1990a, Maehr et al.

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1991a, Maehr 1992, Comiskey et al. 1994, Dees et al. 2001). Periodic understory brushfires (Dees et al. 2001) as well as increased amounts of edge (Miller 1993) may enhance deer use of hardwood hammocks, pine, and other forest cover types.

However, wetland and other vegetation types can support high deer densities. In the Everglades, for example, deer appear to be adapted to a mosaic of intergrading patches comprised of wet prairie, hardwood tree islands, and peripheral wetland habitat (Fleming et al. 1994, Labisky et al. 2003). High-nutrient deer forage, especially preferred by females, includes hydrophytic marsh plants, white waterlily (Nymphaea odorata), and swamp lily (Crinum americana) (Loveless 1959, Labisky et al. 2003). Wetland willow (Salix spp.) thickets provide nutritious browse for deer (Loveless 1959, Labisky et al. 2003).

Marshes, rangeland, and low-intensity agricultural areas support prey populations of deer and hogs. The importance of these habitat types to panthers cannot be dismissed based solely on use or lack of use when daytime telemetry are the only data available (Comiskey et al. 2002, Beier et al. 2003, Comiskey et al. 2004, Beier et al. in press).

Travel and Dispersal Corridors--In the absence of direct field observations / measurements, Harrison (1992) suggested that landscape corridors for wide-ranging predators should be half the width of an average home range size. Following Harrison’s (1992) suggestion, corridor widths for Florida panthers would range 6.1 - 10.9 mi (9.8 - 17.6 km) depending on whether the target animal was an adult female or a transient male. Beier (1995) suggested that corridor widths for transient male puma in California could be as small as 30% of the average home range size of an

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adult. For Florida panthers, this would translate to a corridor width of 5.5 mi (8.8 km). Without supporting empirical evidence, Noss (1992) suggests that regional corridors connecting larger hubs of habitat should be at least 1.0 mi (1.6 km) wide. Beier (1995) makes specific recommendations for very narrow corridor widths based on short corridor lengths in a California setting of wild lands completely surrounded by urban areas; he recommended that corridors with a length less than 0.5 mi (0.8 km) should be more than 328 ft (100 m) wide, and corridors extending 0.6 - 4 mi (1 - 7 km) should be more than 1,312 ft (400 m) wide. The Dispersal Zone encompasses 44 mi2 (113 km2) with a mean width of 3.4 mi (5.4 km). Although it is not adequate to support even one panther, the Dispersal Zone is strategically located and expected to function as a critical landscape linkage to south-central Florida (Kautz et al. in press). Transient male panthers currently utilize this Zone as they disperse northward into south-central Florida.

G. Habitat and Prey Management

Land management agencies in south Florida are implementing fire programs that mimic a natural fire regime through the suppression of human-caused wildfires and the application of prescribed natural fires. Chemical, biological, and mechanical controls of invasive plants are also conducted to maintain and restore native habitat types. Management for panther prey consists of a variety of approaches such as habitat management and regulation of hunting and off-road vehicle (ORV) use.

Eight public land areas within the Primary Zone are managed by five Federal or State agencies and one non-governmental organization (NGO). The annual prescribed fire goals of these public land areas in south Florida total 166 mi2 (430 km2). Two-to-five year fire rotations and burn

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compartments less than 10 mi2 (25 km2) are recommended to increase habitat heterogeneity (Schortemeyer et al. 1991). However, fire prescriptions vary based on fuel conditions, weather conditions, and historic fire frequency. Compartment size will vary based on site conditions, including the use of existing fire breaks or reluctance to establish new fire breaks that would reduce native habitats, fragment native habitats, and serve as vectors for the spread of invasive plants. For example, FPNWR, the only area managed specifically for panthers, uses existing swamp buggy trails and highways as burn compartment boundaries. The Refuge is divided into 54 burn compartments that range in size from 0.47 – 1.72 mi2 (1.22 – 4.45 km2). A range of 8 - 12 mi2 (20 - 32 km2) is burned annually depending on weather conditions. The fire program at BCNP averages 47 - 62 mi2 (121 - 162 km2) burned annually (4 - 5% of the total area) as many habitats are adapted to long fire intervals.

Invasive non-native vegetation has the capacity to replace native plant communities and drastically change the landscape both visually and ecologically. The invasive plants of most concern in south Florida are melaleuca (Melaleuca quinqueneervia), Brazilian pepper (Schinus terebinthifolius), old-world climbing fern (Lygodium microphyllum), cogongrass (Imperata cylindrica), and downy rose-myrtle (Rhodomytrus tomentosus). The effect of invasive plants on panther habitat utilization, particularly melaleuca, is unknown. However these species may reduce the panther’s prey base by disrupting natural processes such as water flow and fire and by significantly reducing available forage. All public lands in south Florida have active invasive plant treatment programs. As of 2002, over 243 mi2 (630 km2) of invasive plants had been treated, with an estimated 579 mi2 (1,500 km2) yet untreated. No studies have been conducted to determine the effects of invasive plant management on panthers. 28

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Prey management has been accomplished by regulating harvest using a variety of strategies. ENP, FSPSP, and FPNWR are closed to hunting. Corkscrew Regional Ecosystem Watershed, Picayune Strand State Preserve, Okaloacoochee Slough State Forest, and BCNP allow hunting. Only BCNP allows ORV use. It also has the longest deer and hog hunting season (95 days), whereas the other three areas allow hunting for 35 days or less annually. A combination of hunter and vehicle use quotas, restrictions on hunting methods, and harvest limits are used in BCNP to regulate impacts on the panthers’ prey base. Over the past 25 years, the annual deer and hog harvest reported at check stations has averaged 210 and 127, respectively, representing a sample of deer and hogs actually harvested. Hunter pressure during that time period has averaged 15,809 “hunter-days” annually (Adams and Bozzo 2002).

H. Response to Management Activities

Few studies have examined the response of panthers to various land / habitat management activities. Dees et al. (2001) investigated panther habitat use in response to prescribed fire and found that panther use of pine habitats was greatest for the first year after the area had been burned and declined thereafter. Prescribed burning is believed to be important to panthers because prey species (e.g., deer and hogs) are attracted to burned habitats to take advantage of changes in vegetation structure and composition, including exploiting hard mast that is exposed and increased quality or quantity of forage (Dees et al. 2001). Responses of puma to logging activities (Van Dyke et al. 1986a) indicate that they generally avoid areas within their home range with intensification of disturbance.

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There is the potential for disturbance to panthers from recreational uses on public lands. Maehr (1990a) reported that indirect human disturbance of panthers may include activities associated with hunting and that panther use of Bear Island (part of BCNP) is significantly less during the hunting season. Schortemeyer et al. (1991) examined the effects of deer hunting on panthers at BCNP between 1983 and 1990. They concluded that, based on telemetry data, panthers may be altering their use patterns as a result of hunting.

Janis and Clark (2002) compared the behavior of panthers before, during, and after the recreational deer and hog hunting season (October through December) on areas open (BCNP) and closed (FPNWR, FSPSP) to hunting. Variables examined were: (1) activity rates, (2) movement rates, (3) predation success, (4) home range size, (5) home range shifts, (6) proximity to ORV trails, (7) use of areas with concentrated human activity, and (8) habitat selection. Responses to hunting for variables most directly related to panther energy intake or expenditure (i.e., activity rates, movement rates, predation success of females) were not detected (Janis and Clark 2002). However, panthers reduced their use of Bear Island, an area of concentrated human activity, and were found farther from ORV trails during the hunting season, indicative of a reaction to human disturbance (Janis and Clark 2002). Whereas the reaction to trails was probably minor and could be related to prey behavior, decreased use of Bear Island most likely reflects a direct reaction to human activity and resulted in increased use of adjacent private lands (Janis and Clark 2002).

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I. Reasons for Listing / Threats Assessment

The Florida panther was listed as endangered throughout its range in 1967 (32 FR 4001), pursuant to the Endangered Species Preservation Act, and received Federal protection under the passage of the ESA in 1973. The 1967 document did not address the five factor threats analysis. However, we address these factors in the summary below to organize the threats to the panther in a manner consistent with current listing and recovery analyses under the ESA.

Overview of Species Decline / Causes of Decline--Historically, the panther occurred throughout the Southeast (Figure 1). Persecution, prey decline, habitat loss and fragmentation, and problems associated with small population size including inbreeding depression and reduced genetic health, resulted in a population in danger of extinction. Habitat loss, degradation, and fragmentation are the most significant threats to the continued survival of the panther throughout its range. In addition, human-related disturbance and mortality, disease, genetic problems, and contaminants are adversely affecting the panther population.

Threats Assessment--A detailed threats assessment for the panther was conducted by the Florida Panther Recovery Team using The Nature Conservancy’s (TNC) planning approach (TNC 2000) (Appendix B). Using this approach, the stresses (the types of degradation and impairment) for each factor were identified and evaluated in terms of severity and scope; sources of stresses were evaluated in terms of contribution and irreversibility. Separate analyses were conducted for the panther population in south Florida and for reintroduction in the Southeast.

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Factor A: The present or threatened destruction, modification, or curtailment of its habitat or range--The panther’s current occupied range is significantly reduced from its historic range from Louisiana and Arkansas east to South Carolina and southward through Florida. The panther occurs only in south Florida, less than 5% of its historic range (Figure 1). Because of their wide-ranging movements and extensive spatial requirements, panthers are sensitive to habitat fragmentation (Harris 1984).

Land Use Changes in Florida--Habitat loss, fragmentation, and degradation, and associated human disturbance are the greatest threats to panther survival and among the greatest threats to its recovery. These threats are expected to continue in Florida and throughout the Southeast. Throughout Florida, between 1936 and 1987, cropland and rangeland increased 6,609 mi2 (17,118 km2) or 30%, urban areas increased by 6,172 mi2 (15,985 km2) or 538%, while herbaceous wetlands declined by 6,063 mi2 (15,702 km2) or 56% and forests declined by 6,719 mi2 (17,402 km2) or 21% (Kautz et al. 1993, Kautz 1994). Assuming that all of the forest lost was panther habitat, Kautz (1994) estimated that the 21% loss of forests was the equivalent of 35 - 70 male panther home ranges and 100 - 200 female panther home ranges. Between 1985 – 1989 and 2003 an additional 5,019 mi2 (13,000 km2) (13%) of natural and semi-natural lands (including panther habitat) in the state were converted to urban / developed and agricultural uses (Kautz et al. in draft).

Continued expansion of urban areas on the coasts and the spread of agricultural and urban development in the interior of Florida continue to replace, degrade, and fragment panther habitat, placing the panther at greater risk. Agricultural development continues to replace and fragment

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panther habitat. Over 83% of the 2,500 mi2 (6,475 km2) of agricultural land in southwest Florida has been categorized as rangeland. Between 1986 and 1990, row crop acreage increased by 14 mi2 (36 km2) or 21%; sugarcane increased by 25 mi2 (65 km2) or 21%; citrus increased by 84 mi2 (219 km2) or 75%; and rangeland, much of it suitable for panther occupation, decreased by 250 mi2 (647 km2) or 10% (Townsend 1991). Rangeland losses were about evenly divided between agricultural and urban development (Townsend 1991).

The extent of land use conversions for southwest Florida (Collier, Lee, Hendry, Charlotte, and Glades Counties) between 1986 and 1996 was estimated using a change detection analysis performed by Beth Stys (FWC, unpublished data). The area of disturbed lands increased 31% in these five counties between 1986 and 1996, with the greatest increases in disturbed lands occurring in Hendry and Glades Counties. Most (66%) of the land use change over the 10-year period was due to conversion to agricultural uses. Forest cover types accounted for 42% of land use conversions, dry prairies accounted for 37%, freshwater marsh accounted for 9%, and shrub and brush lands accounted for 8%. Randy Kautz (FWC, pers. comm. 2003) estimated panther habitat loss to be 0.8% per year between 1986 and 1996 using a composite of three different methodologies. These included: (1) review of U.S. Forest Service forest data between 1936 and 1995 using loss of forest as an index of the rate of panther habitat loss, (2) analysis to detect changes in land cover in five south Florida counties (Charlotte, Collier, Glades, Hendry, Lee) between 1986 and 1996 using classified Landsat imagery, and (3) using the Cox et al. (1994) panther habitat model, and based on 1986 Landsat data, 1996 Landsat landcover data was overlaid and then areas originally mapped as panther habitat and subsequently converted to other uses over the 10-year period were tabulated. Kautz (Breedlove, Dennis, and Associates, pers.

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comm. 2005) believes the estimated annual habitat loss since 1996 may be 2 to 3 times higher than that calculated for the previous period.

More recently, Stys calculated the extent of semi-natural and natural lands that have been converted to agricultural and urban / developed in Florida between 1985 - 1989 and 2003 (B. Stys, FWC, pers. comm. 2005). Based upon this analysis, approximately 570 mi2 (1,476 km2) of natural and semi-natural lands in Glades, Hendry, Lee, Collier, Broward, Monroe, and Miami-Dade Counties were converted during this time period (FWC, unpublished data). Of these, approximately 340 mi2 (880 km2) were conversions to agricultural uses and 230 mi2 (596 km2) to urban uses. Nearly 42% (142 mi2 or 369 km2) of the conversions to agriculture occurred in Hendry County. These conversions have been offset to some degree (19 mi2 [49 km2]) by habitat conservation elsewhere in south Florida, particularly in recent years.

Rapid development in southwest Florida has compromised the ability of landscapes to support a self-sustaining panther population (Maehr 1990b, 1992). Maehr (1990b) reported that there were approximately 3,401 mi2 (8,810 km2) of occupied panther range in south Florida and that approximately 50% is comprised of landscapes under private ownership. In 2005, Kautz found that approximately 22% of the land in the Primary Zone, 60% of the land in the Secondary Zone, and 100% of the land in the Dispersal Zone is in private ownership (R. Kautz, pers. comm. 2005). Maehr (1990b) indicated that development of private lands may limit panther habitat to landscapes under public stewardship. Given the panther’s reliance on public land, the rising cost of land is an impediment to habitat protection and therefore panther conservation and recovery.

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Highways in wildlife habitat are known to result in loss and fragmentation of habitat, traffic related mortality, and avoidance of associated human development. As a result, small populations may become isolated, subjecting them to demographic and stochastic factors that reduce their chances for survival and recovery. Two-lane 108 ft (33 m) and four-lane 328 ft (100 m) cleared rights-of-way, respectively, occupy 2.0 and 6.2% of each 640 ac (259 ha) of land through which they pass (Ruediger 1998). Highways can also stimulate land development as far away as 2 mi (3.2 km) on either side (Wolf 1981). Thus, for each 1 mi (1.6 km) a highway is extended, 2,500 ac (1,012 ha) are potentially opened to new development (Wolf 1981).

Belden and Hagedorn (1993) observed that Texas pumas introduced into northern Florida established home ranges in an area with one-half the road density of the region in general, and tended to avoid crossing heavily traveled roads. Of 26 western puma home ranges examined by Van Dyke et al. (1986b), 22 (85%) included unimproved dirt roads, 15 (58%) included improved dirt roads, and only six (23%) included hard-surfaced roads. Female panthers rarely establish home ranges in areas bisected by highways (Maehr 1997b). Because home ranges of resident males typically encompass the ranges of up to six female panthers, males are less likely than females to find sufficiently large areas devoid of major roads. Males tend to cross highways more frequently than females and suffer more vehicle-related injuries and mortalities (see Factor E).

In addition to a direct loss and fragmentation of habitat, constructing new and expanding existing highways may increase traffic volume and impede panther movement within and between frequently used habitat blocks throughout the landscape (Swanson et al. in review). Increases in

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traffic volume, increasing size of highways (lanes), and habitat alterations adjacent to key road segments may limit the panther’s ability to cross highways and may ultimately isolate some areas of panther habitat (Swanson et al. in review). The addition of wildlife crossings and fencing has ameliorated this threat in the immediate vicinity of these structures. The addition of more wildlife crossings, especially in areas with a history of collisions and where traffic is projected to increase, can help address this significant threat.

Past land use activity, hydrologic alterations, and lack of fire management (Dees et al. 1999) have also affected the quality and quantity of panther habitat. The effect of invasive plants on panther habitat utilization, particularly melaleuca, is unknown. As the remaining forested uplands are lost, sloughs containing cypress, marsh, and shrub wetlands comprise a greater percentage of the remaining habitat available to panthers, relative to habitat historically available to the species.

Human Population Growth--Insight can be gained into expected rates of habitat loss in the future by reviewing human population growth projections for the south Florida region. Smith and Nogle (2001) developed low, medium, and high population growth projections for all Florida counties from 2000 through 2030. Using their medium projections, which they believe provide the most accurate forecasts, Smith and Nogle (2001) estimate that the human population of the 10 counties in south Florida will increase from 6.09 - 9.52 million residents by 2030, an increase of 56%.

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Human population in the southeastern U.S. has increased 10-fold since 1850, expanding from 4.7 million to over 48 million in 2000 (cited in Swanson et al. in review). In Florida, the population increased from 87,000 to over 17 million (cited in Swanson et al. in review, U.S. Census Bureau 2004). From 1990 - 2004, the population in Collier County increased from 152,099 to 296,678 (U.S. Census Bureau 2002, 2004). During the same time period, the population in Lee County increased from 335,113 to 514,295 (U.S. Census Bureau 2002, 2004). The population of southwest Florida, particularly Collier and Lee Counties, is projected to increase 21% by 2010 (cited in Swanson et al. in review).

Land Use Changes in Southeastern States--Based on the current trends of urbanization across the southeast, it is likely that forested habitats will continue to be permanently altered, and the amount of available forest habitat will decrease in some areas (Wear and Greis 2002). Compared to earlier periods, land use in the southeast has been fairly stable since 1945, with the most notable exception of Florida, where developed land uses have expanded substantially (Wear and Greis 2002). Two dominant forces strongly influenced recent land use changes: (1) urbanization driven by population and general economic growth and (2) changing relative returns to agriculture and timber production; both of these influences are expected to continue (Wear and Greis 2002). As a result of anticipated population and economic growth, rural land will be converted to urban uses. Forecasts of land uses indicate that the southeast could experience a net loss of from 12,500 - 18,750 mi2 (32,375 - 48,562 km2) of forest land (roughly 5 - 8%) between 1992 and 2020 (Wear and Greis 2002).

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Potential panther habitat throughout the Southeast continues to be affected by urbanization, residential development, conversion to agriculture and silviculture, mining and mineral exploration, lack of land use planning, and other sources of stress (Appendix B). With human population growth and increased human disturbance, the extent of potentially suitable habitat remaining in the Southeast is expected to decrease. Habitat loss, fragmentation, degradation, and disturbance from human activity throughout the Southeast are expected to remain among the greatest threats to reintroduced panther populations. As development pressure and population growth continues, the opportunity for panther reintroduction in the Southeast could be diminished.

Factor B: Overutilization for Commercial, Recreational, Scientific, or Educational Purposes—There are no commercial or recreational uses of panthers. In rare cases where a panther is unable to survive in the wild, it may be captured and used for educational purposes. However, panthers are routinely captured and monitored for scientific purposes. Risks are associated with capture and monitoring, but the overall threat to the panther is considered low (Appendix B). Capturing and radiocollaring panthers and handling neonate kittens at dens may result in unintentional take relative to three factors. First, mortality or injury may result from the capture event because of capture induced trauma or an adverse reaction to immobilizing chemicals. Routine capture activities include the use of trained hounds to pursue and tree the panther and the subsequent anesthetization of the animal with remotely-injected immobilizing drugs. These activities may result in hyperthermia, hypothermia, dog bite wounds, drowning, fractures, lacerations, seizures, head and spinal trauma, penetration of the abdomen or thorax with dart, vomiting, aspiration, pneumothorax, respiratory depression or arrest, shock, cardiac arrest, or complications associated

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with treatment of the above conditions. However, the incidence of these injuries, especially serious injuries and mortalities, has been low over the last 25 years of panther capture work in part because of stringent capture and handling protocols developed by FWC, NPS, and FWS. Since 1981, the FWC has captured and immobilized 133 panthers over 296 times with only one fatality, two panthers suffering broken legs that resulted in their temporary removal to captivity for rehabilitation and the successful return to the wild, and the holding of one other panther for 24 hours to treat an injury involving a needle embedded in bone (D. Land, FWC, pers. comm. 2004). NPS staff in BCNP have been capturing adult panthers and handling kittens at dens since 2003. Between 2003 and 2005, the NPS handled 19 adult or dependent juvenile panthers with no injury or mortality (Jansen et al. 2005).

Second, capture and handling events can result in abandonment of kittens, other disruptions of family structure, or injury to a kitten that requires its removal from the wild for rehabilitation. Further, the injury or death of an adult female with dependent-aged kittens (those less than 1 year of age) could result in the death of the kittens or the need to raise them in captivity. Neonate kittens are handled at den sites when the kittens are older than 2 weeks of age and when the mother is not present. These activities do not require anesthesia of the kittens. Handling activities could result in injury or death to the kitten or the abandonment of one or more of the kittens. From 1986 - 2004, the FWC has captured and radiocollared 59 dependent-aged kittens ranging in age from 4 - 18 months (D. Land, pers. comm. 2004). These captures resulted in the abandonment of two kittens. One was subsequently reared in captivity and released. The other died of an infection in captivity shortly after its capture. Early break-up of family groups may have occurred on a few other occasions. For this reason, dependent-aged kittens less than one

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year are no longer captured. Between 1992 and 2005, FWS and NPS handled 195 kittens at 82 dens with no injury, mortality, or den abandonment (Jansen et al. 2005, Lotz et al. 2005).

Third, the loss of contact with or access to young radio-collared panthers whose collars need to be resized to accommodate growth may result in the collar becoming embedded in the panther’s neck. If the panther cannot be recaptured to remove (e.g., if a radiocollar prematurely fails) or resize the collar, infection and eventual death could occur. In September 2001, the FWC and NPS began fitting young panthers with break-away radiocollars. This change in protocol has greatly reduced the risks associated with radiocollaring young panthers (D. Land, pers. comm. 2004).

If stringent capture and handling protocols continue to be followed and refined, injury levels are expected to remain low and are not expected to significantly affect important demographic parameters at the population level, including mortality and reproductive rates or recruitment of juveniles. Handling panthers is important for research, management, and monitoring of the population, and overall the risks are low.

Factor C: Disease or Predation-- The Florida panther is susceptible to a number of infectious and parasitic diseases some of which are of population significance while others are important only to the individual. Some diseases have not been diagnosed in panthers but remain a potential threat. As a single contiguous population, there is potential for an infectious disease to have a catastrophic impact on the panther population.

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Although FeLV is common in domestic cats (Felis catus), it is quite rare in non-domestic felids. The recent outbreak of this disease in the panther population shows the potential of this disease to be of population significance. Another viral disease potentially of population significance is PRV. PRV causes respiratory and reproductive disorders in adult hogs and mortality in neonates, but is a rapidly fatal neurologic disease in carnivores. Approximately 35% of feral hogs are seropositive for PRV in Florida (van der Leek et al. 1993). The virus is actively shed by only a small percentage of infected hogs at any given time; however, stress can increase the percentage that shed the virus (Murphy et al. 1999). Feral hogs are an important prey species for panthers (Maehr et al. 1990b), and there is potential for significant mortality in panthers due to PRV.

Raccoons are a common prey item for panthers (Maehr et al. 1990b) and are the most important reservoir for rabies in the Southeast (Burridge et al. 1986). As panthers are now vaccinated against rabies at capture, only uncollared panthers are at significant risk.

Feline panleukopenia virus (PLV) causes significant mortality in domestic kittens. The virus is also carried by raccoons and is quite stable in the environment. However, kittens are at greatest risk of infection and causes of mortality in this cohort are largely unknown. An epizootic of PLV caused significant mortality among radio-collared bobcats in the late 1970s in south-central Florida (Wassmer et al. 1988), suggesting that the panther population may also be at risk.

Hookworm infections in domestic kittens can cause significant morbidity and mortality resulting from blood loss. The impact of this parasite on panther kittens in the wild is unknown.

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Some individual panthers have been shown to be at risk from exposure to mercury in the food chain (Newman et al. 2004). Mercury bioaccumulates through the aquatic food chain reaching high concentrations in higher trophic level carnivores such as raccoons and alligators. Panthers preying on these species are at risk for accumulating high tissue mercury concentrations. Neonates may be more susceptible to the toxic effects of mercury (Berglund and Berlin 1969).

Disease and parasites have not been documented to be a major mortality factor in the panther population (Maehr et al. 1991b, Taylor et al. 2002). However, this observation is largely based on the captured and vaccinated sample of the population. Disease expression and mortality events for the unmarked and unvaccinated segment of the population, including kittens, may be higher, especially for those diseases included in the vaccination regimen. Further, as the panther population density increases there is an increased risk of diseases transmitted by direct contact. The recent outbreak of FeLV demonstrated the potential impact of infectious diseases on the population. Should a virulent pathogen enter the population, such as occurred with FeLV, there is no absolute barrier in south Florida that could prevent such a disease from impacting the entire population (Beier et al. 2003). Consequently, until additional populations of panthers can be established elsewhere in their historic range, infectious diseases and parasites remain a threat to the south Florida population. Finally, infectious diseases, parasites, and environmental contaminants, even of low pathogenicity, may work synergistically to reduce panther fitness and reproduction.

Factor D: The Inadequacy of Existing Regulatory Mechanisms--The panther is federally listed as endangered and is on the State endangered lists for Florida, Georgia, Louisiana, and

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Mississippi. The protection provided by Federal (ESA, Clean Water Act [62 Stat. 1155, as amended; 33 U.S.C. 1251-1376] [CWA], National Environmental Policy Act of 1969 [83 Stat. 852, as amended; 42 U.S.C. 4321-4347] [NEPA], Fish and Wildlife Coordination Act [48 Stat. 401, as amended; 16 U.S.C. 661 et seq.] [FWCA]) and State (Florida protective provisions specified in Rules 68A-27.0011 and 68A-27.003) laws help conserve the panther and its habitat.

Section 7(a)(2) of the ESA requires that all Federal agencies consult with the Service to insure that any action authorized, funded, or carried out by the agency is not likely to jeopardize the continued existence of any listed species or result in the destruction or adverse modification of critical habitat. If a project will not jeopardize the continued existence of a species but may result in incidental take of the species, the Service works with the action agency and any applicants to find ways to minimize the effects of the take. Section 7(a)(1) requires all Federal agencies to utilize their authorities in furtherance of the ESA by carrying out programs for the conservation of listed species pursuant to section 4. Section 9 prohibits unlawful acts, including unauthorized take. Section 10(a)(1) allows for the issuance of permits for scientific or enhancement of survival purposes, provided that certain terms and conditions are met. Section 10(a)(2) allows for the issuance of permits, provided that the taking will be incidental to an otherwise lawful action, adequately minimized and mitigated, appropriately funded, and will not appreciably reduce the likelihood of survival and recovery of the species in the wild. Section 4(a)(3) requires the designation of critical habitat for listed species to the maximum extent prudent and determinable.

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As discussed previously in Factor A, development pressure in southwest Florida has been high; for example, data for Collier, Lee, and Hendry Counties, a stronghold for the panther population, indicate that from 1985 through 2003 more than 223 mi2 (578 km2) of natural and semi-natural lands were converted to agriculture (FWC, unpublished data). In addition, more than 145 mi2 (375 km2) of semi-natural and natural lands in this three-county area have also been lost to development (FWC, unpublished data) (see Factor A). While not all of these habitat losses and conversions involved panther habitat, many projects involved wetland impacts, requiring permit review by the U.S. Army Corps of Engineers (COE) pursuant to section 404 of the CWA and / or coordination among regulatory agencies pursuant to the FWCA. For projects with a Federal nexus, consultation pursuant to section 7 of the ESA was needed for actions that may affect the panther and other listed species. Especially in recent years, impacts have been offset by protection and restoration of important panther habitat.

Through 2005, no Habitat Conservation Plans (HCP) have been finalized under section 10(a)(2) of the ESA and no incidental take permits have been issued for the panther. Most panther habitat is interspersed with wetlands, which often require a section 404 permit from the COE and therefore are reviewed under section 7 of the ESA. Section 10, however, provides opportunities for large-scale and regional approaches to panther habitat conservation, and promises to be an especially valuable tool at the county level.

Florida Statute 373.414 requires that activities permitted in wetlands and surface waters of the state are not contrary to the public interest. If it is determined that an activity will adversely effect panthers or panther habitat, the governing board (Water Management District [WMD]) or

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the Florida Department of Environmental Protection (FDEP) can consider measures (e.g., on-site mitigation, off-site mitigation, purchase of credits from mitigation banks) that will mitigate the effects of the regulated activity.

In addition to the impacts of individual projects, the FDEP and WMD shall take into account cumulative impacts on water resources and manage those resources in a manner to ensure their sustainability (Chapter 373.016(2) F.S.). Cumulative impacts can be considered unacceptable when they provide significant impacts to functions of wetlands, including the utilization of the wetlands by wildlife species. In practice, evaluating cumulative impacts of development in southwest Florida on panthers has not been sufficient to prevent significant loss of panther habitat.

Since the majority of panther habitat in southwest Florida has significant wetland components, provisions of 373.414 are usually a part of the review of proposed development. However, the state wetlands permitting authorities currently lack comparable regulatory mechanisms to assess impacts to panthers or panther habitat on project sites that do not have a wetland component.

The FWC may exercise the regulatory and executive powers of the State with respect to wild animals, including panthers. The FWC has responsibility for conserving and managing these species and their habitat; however the FWC does not provide regulatory protection for listed species habitat. The FWC provides comments regarding potential impacts to panther habitat to FDEP and WMDs under the authority of Chapter 20.331 Florida Statutes.

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Because of the project-specific focus of regulatory programs and other constraints such as high workloads, local, State, and Federal regulatory agencies sometimes find it difficult to complete the cross-government review that would be ideal to thoroughly review and effectively assess all potential impacts to panthers. In addition, local, State, and Federal agencies sometimes have difficulty monitoring permit compliance and tracking the precise impact on species and habitat from authorized actions, as well as tracking the impact from unauthorized actions. Assessing current baseline conditions and accurately predicting future impacts are also challenging because the panther is a wide-ranging species that uses a wide array of habitat types. Furthermore, baseline conditions for the panther are continually changing (e.g., impacts from development, conservation actions). Through consultation under section 7(a)(2) of the ESA, the Service evaluates impacts to panthers from proposed projects against compensation offered to minimize those impacts. Through this process, over 19 mi2 (49 km2) were secured in the Primary and Dispersal Zones from September 2003 to May 2005. Rigorous assessments and close coordination and scrutiny of project impacts by local, State, and Federal agencies during the planning phase could help maximize conservation benefits for the panther.

Factor E: Other Natural or Manmade Factors Affecting its Continued Existence--

Mortality, Trauma, and Disturbance--Florida panthers were hunted for bounty during the 1800s and for sport up until the 1950s. Nine illegal shootings were documented in south Florida between 1978 and 2005, three of which were not fatal. Education, self-policing among hunters, and regulation are the tools by which shootings are minimized. All free-ranging puma in Florida are protected by a “similarity of appearance” provision pursuant to the ESA.

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Records on documented mortality of uncollared panthers have been kept since February 13, 1972. Records on mortality of radio-collared panthers have been kept since February 10, 1981. Eighty-four radio-collared panthers have died since 1981, and intraspecific aggression was the leading cause, accounting for 42% of these mortalities (Lotz et al. 2005). Unknown causes and collisions with vehicles accounted for 24% and 19% of mortalities, respectively. Other factors (7%), infections (5%), and diseases (4%) caused the remaining mortalities (Land et al. 2004).

One-hundred fifty-three panther mortalities have been documented from February 1972 through June 2004, with at least 58 (41%) of known deaths occurring in the latest four-year period (Land et al. 2004). Overall, documented mortality (n = 105) of radiocollared and uncollared panthers averaged 3.4 per year through June 2001. However, from July 2001 through June 2004, documented mortality (n = 48) increased with an average of 16.0 per year during these years (Land et al. 2004). This increase in panther mortality (e.g., intraspecific aggression, collisions with vehicles) corresponds with increases in the panther population observed in recent years. This increased mortality may indicate a population that is at or approaching carrying capacity (C. Belden, FWS, pers. comm. 2005).

From February 1972 through June 2004, 36 panthers were documented to have died from intraspecific aggression (Land et al. 2004). Although most of these encounters are male-male, from July 2001 through June 2004, at least nine females have been killed in encounters with males (Land et al. 2004). Defense of kittens and / or a kill is suspected in five of these instances that occurred through 2003 (Shindle et al. 2003).

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From February 1972 through June 2004, 27 panthers were documented to have died from unknown causes (Land et al. 2004). While a couple of deaths from unknown causes occur each year, five deaths occurred in various areas in 2000 and six deaths occurred in a localized area (Seminole game and safari pens) in 2003 (Land et al. 2004).

Eighty-six panther-vehicle collisions were documented between 1972 and 2005 of which 80 (52%) resulted in panther deaths (Lotz et al. 2005). However, panther-vehicle collisions were identified as the third most important source of mortality among radiocollared panthers (19%), a less biased sample (Land et al. 2004). Fifty-six percent (48) of panther-vehicle collisions have occurred since 2000 with all but two being fatal to the panther (Lotz et al. 2005). Approximately 53% of documented panther roadkills have occurred within the Primary Zone through 2004 (Swanson et al. in review). Panther-vehicle collisions are a significant source of mortality and pose a serious on-going threat to the species. In addition, new and existing roads, expansion of highways, and increases in traffic volume and speed contribute to a loss of panther habitat and impede movement within and between high use habitat blocks throughout the landscape (Swanson et al. in review) (see Factor A). New and expanded highways are likely to increase the threat of panther mortality and injuries due to collisions.

Wildlife crossings and continuous fencing were required during the conversion of two-lane SR 84 (Alligator Alley) into four-lane I-75. To date, no panthers have been killed in these protected areas since completion of I-75 in 1992. Similarly, six wildlife crossings and some fencing were required along SR 29 as a prerequisite to the SR 29 / I-75 interchange. Four of these crossings are now complete and the completed fencing-crossing areas have prevented panther-vehicle

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collisions. In addition, a crossing was required on CR 858 (Oil Well Road) to offset projected traffic increases from development. In the absence of crossings and fencing, the remaining stretches of SR 29 and I-75 as well as several other roads continue to pose a serious mortality risk to panthers, including U.S. 41 (Tamiami Trail), SR 82, and County Roads (CR) 850 (Corkscrew Road), 858, 846 (Immokalee Road), 832, and 833. Up through July 2004, 59 of 73 panther roadkills or injuries occurred along these unsecured roads (Swanson et al. in review).

Florida’s human population has been steadily growing and as a result, urban / suburban areas now interface with panther habitat. Extensive developments planned within two districts within Collier County’s Rural Lands Stewardship Plan have the ability to affect highway infrastructure within the Primary and Secondary Zones (Swanson et al. in review). Extensive developments planned within these districts, such as the Ave Maria University and associated town, will expand local road networks and extend the human / panther interface into primary panther habitat (Swanson et al. in review).

In recent years, there has been an increase in potential for panther-human interactions and disturbance associated with management responses to panthers that have interacted with humans. For example, in 2004, aversive conditioning was used on panthers observed near areas of human habitation in the Pinecrest area within BCNP, and a juvenile dependent male panther was subsequently relocated to Okaloacoochee Slough State Forest. If panther-human interactions increase, the potential for complaints from the public and, in some cases, the need for subsequent management responses could result in take of panthers in the form of harassment through aversive conditioning in an attempt to teach individuals to avoid humans. In extreme cases,

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permanent removal from the wild is possible. Currently, the Service, the FWC, and the NPS are working on a document titled Guidelines for Living with Florida Panthers and Florida Panther Response Plan. This plan will provide guidance on methods for minimizing the potential for panther-human interactions and help ensure consistency in use of potential management responses.

There is the potential for disturbance to panthers from recreational uses on public lands. Maehr (1990a) and Schortemeyer et al. (1991) reported that panthers may be altering their use patterns as a result of hunting. Janis and Clark (2002) compared the behavior of panthers before, during, and after the recreational deer and hog hunting season on areas open and closed to hunting. Responses to hunting for variables most directly related to panther energy intake or expenditure were not detected (Janis and Clark 2002). However, panthers reduced their use of an area of concentrated human activity, and were found farther from ORV trails during the hunting season, indicative of a reaction to human disturbance (Janis and Clark 2002). Whereas the reaction to ORV trails was probably minor and could be related to prey behavior, decreased use of areas of human activity most likely reflects a direct reaction and resulted in increased use of adjacent private lands. Additional habitat loss on those private lands could exacerbate the negative consequences of this pattern of use (Janis and Clark 2002).

Loss of Genetic Diversity--Natural genetic exchange with other panther populations ceased when the Florida panther became geographically isolated over a century ago (Seal 1994a). Isolation, reduced population size, and inbreeding have resulted in loss of genetic variability and diminished health. Data on polymorphism and heterozygosity, along with records of multiple

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physiological abnormalities, suggest that the panther population has experienced inbreeding depression (Roelke et al. 1993b, Barone et al. 1994). Inbreeding has been related to decreased semen quality, lowered fertility, reduced neonatal survival, and congenital heart defects in a variety of domesticated and wild species (Lasley 1978, Ralls and Ballou 1982, O’Brien et al. 1985, Roelke 1991). Genetic problems in the Florida panther included heart murmurs, a high rate of unilateral cryptorchidism, low testicular and semen volumes, diminished sperm motility, and a high percentage of morphologically abnormal sperm.

Geographic isolation, habitat loss, small population size, and associated inbreeding have resulted in the loss of the panther’s genetic diversity (Roelke 1990). Measured heterozygosity levels indicate that the Florida panther has lost about 60 - 90% of its genetic diversity (Culver et al. 2000). Measured levels of mDNA variation are the lowest reported for any similarly studied feline population, including leopards, cheetahs, and other puma subspecies. Electrophoretic analyses also indicated that the Florida panther has less genetic variation than any other puma subspecies. Panther DNA fingerprint variation is nearly as low as in the small, isolated population of Asiatic lions of the Gir Forest Sanctuary in India (Roelke et al. 1993b).

To address these threats, a genetic management program was implemented with the release of Texas puma into south Florida in 1995 (see Conservation Efforts Section). The initial results of genetic restoration have been promising (Hedrick 2004), with an increasing population, signs of increased genetic health, recolonization of areas in BCNP and ENP recently unoccupied, and increased dispersal (McBride 2000, 2001, 2002; Maehr et al. 2002a). To date, neither atrial septal defects nor cryptorchidism have been found in introgressed panthers (M. Cunningham, 51

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pers. comm. 2005). Semen examination of a couple of introgressed panthers indicated that sperm volume, motility, and count were higher than for an uncrossed Florida panther. The FWC indicates that representation of Texas puma genes is probably close to the original genetic restoration program goal of 20%, although two of the eight Texas females are overrepresented (Shindle et al. 2001, Land et al. 2004). Genetic introgression is also reducing the occurrence of kinked tails and cowlicks in intercross progeny (Land et al. 2004).

Human Dimension--Previous recovery plans have called for the establishment of additional populations within the historic range of the Florida panther (FWS 1981, 1987, 1995). The FWC studied the possibility of establishing additional populations within the historic range of the panther (Belden and Hagedorn 1993, Belden and McCown 1996). Between 1988 and 1995, 26 Texas puma were released near Okefenokee NWR and Osceola National Forest. Study animals, monitored by radiocollars at least three days per week, established large home ranges, killed large prey at expected frequencies, and generally adapted well to their new environment (Belden and McCown 1996). When these studies were terminated, the remaining panthers were captured and removed from the wild.

Experimental releases of Texas pumas indicated that habitat and prey availability in northern Florida and southern Georgia were sufficient to support a panther population (Belden and McCown 1996). However, although there appeared to be support for reintroduction among the general public in Florida, local landowners tended to oppose having panthers on their property. Political and social issues will be the most difficult aspect of panther reintroduction and must be

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resolved before further restoration efforts are initiated (Belden and Hagedorn 1993, Belden and McCown 1996).

Habitat assessment studies have been conducted to identify potential sites for reintroduction of the panther in the Southeast (Thatcher et al. 2003, Thatcher et al. in press). The purpose of these studies was to identify prospective sites for panther reintroduction within the historic range based on quantitative landscape assessments. Nine potential reintroduction sites of sufficient size to support a panther population have been found including: Ozark National Forest region, Ouachita National Forest region, southwest Arkansas, and Felsenthal NWR region in Arkansas; Kisatchie National Forest region in Louisiana; Homochitto National Forest region in Mississippi; southwest Alabama; Apalachicola National Forest region in Florida; and Okefenokee NWR region in Georgia (Thatcher et al. in press). Of the nine areas identified, the Okefenokee NWR, Ozark National Forest, and Felsenthal NWR regions ranked as the best prospective reintroduction sites based on the numerical combination of effective habitat area and expert model scores (Thatcher et al. in press).

Sociopolitical obstacles to large carnivore reintroduction are often more daunting than biological ones (Clark et al. 2002). A lack of public support and tolerance could prevent the reintroduction of panthers anywhere outside of Florida. Public opinion is the most critical impediment to reintroduction efforts and attainment of recovery goals.

Contaminants--Because the panther is a top carnivore, bioaccumulation of environmental contaminants remains a concern (Dunbar 1995, Newman et al. 2004), with the threat of mercury

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toxicity considered medium (See Appendix B). However, mercury in the Everglades ecosystem has decreased over the last several years (Frederick et al. 2002).

Other environmental contaminants found in panthers include polychlorinated biphenyls (Arochlor 1260) and organochlorines (e.g., p, p’–DDE) (Dunbar 1995, Land et al. 2004). Continued monitoring for contaminants, especially mercury and organochlorines, in panthers, their prey, and sentinel species is warranted (see E. Life History / Ecology).

Prey availability--The size, distribution, and abundance of available prey species are critical factors to the persistence of panthers in south Florida and often determine the extent of panther use of an area. A resident adult male puma generally consumes one deer-sized prey every 8 - 11 days; this frequency would be 14 - 17 days for a resident female; and 3.3 days for a female with three 13-month-old kittens (Ackerman et al. 1986).

Historically, hunting in the Big Cypress physiographic region has been a major recreational activity with many hunt camps throughout the region. With establishment of national and state parks, the numbers of hunt camps were minimized and additional hunting regulations that reduced hunting pressure on deer were implemented. Although deer densities are difficult to determine, the deer population appears to have steadily increased.

Using aerial surveys, Schemnitz (1974) estimated the deer population in the 3,438 mi2 (8,903 km2) in the area south of the Caloosahatchee River and Lake Okeechobee) at 20,000 in 1972, and stated that the deer population had decreased in the Water Conservation Areas (WCA) due to

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deeper water levels and submersion of tree islands. Fleming et al. (1994) compared deer density estimates in WCAs 2 and 3 in the 1950s with those from 1985 - 1988 and found a 67% reduction in the deer herd. They surmised that this reduction was due to habitat degradation from impoundment and associated water management.

ENP and portions of the WCAs are within the Primary Zone. If the implementation of the Comprehensive Everglades Restoration Program (CERP) results in higher water levels in the WCAs and ENP, the panthers’ prey base could be reduced. Smith and Bass (1994), however, stated that fire and water, which drive the Everglades system, appear to have little effect on the long-term dynamics of the ENP deer population.

Few studies have been done on the hog component of the panthers’ prey base (e.g., Maehr et al 1989b). However, the mean checked hog harvest of 29 in BCNP for the past three years (2003 - 2005) has fallen well below the previous 22-year average of 144, probably due to a combination of factors, including high water events and predation by panthers (D. Jansen, pers. comm. 2005).

Although the exact status of prey in different portions of the panther’s occupied range is not known at this time, assessment of overall panther health and their success in raising young indicate that the prey base is adequate to support the current panther population. Adequate prey elsewhere within the historic range would be needed to establish populations in other areas.

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J. Past and Current Conservation Efforts

Habitat Conservation and Protection--Habitat protection has been identified as being one of the most important elements to achieving panther recovery. While substantial efforts have been made to secure a sufficient habitat base (Figure 4), continued action is needed to obtain additions to and inholdings for public lands, assure linkages are maintained, restore degraded and fragmented habitat, and obtain the support of private landowners for maintaining property in a manner that is compatible with panther use. Conservation lands used by panthers are held and managed by a variety of entities including FWS, NPS, Seminole Tribes of Florida, Miccosukee Tribe of Indians of Florida, FWC, FDEP, Florida Division of Forestry (FDOF), WMDs, NGOs, counties, and private landowners.

Public Lands--Public lands in south Florida that benefit the panther are listed below and shown in Figure 4:

• In 1947, ENP was established with 2,356 mi2 (6,102 km2) and in 1989 was expanded with the addition of 163 mi2 (421 km2).

• In 1974, Congress approved the purchase and formation of BCNP, protecting 891 mi2 (2,307 km2), later 228 mi2 (591 km2) were added.

• In 1974, the State of Florida began acquiring land for the FSPSP, which encompasses over 125 mi2 (324 km2). Efforts are underway to acquire approximately 26 mi2 (68 km2).

• In 1985, acquisition of Picayune Strand State Forest and Wildlife Management Area