A State of Knowledge of the Natural, Cultural, and Economic Resources

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Natural Resource Stewardship and Science A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Natural Resource Report NPS/NRSS/BRD/NRR—2016/1243 ON THIS PAGE Photograph of Ibis, Mobile Bay, Alabama. Photograph courtesy of Hunter Nichols (http://www.hnproductions.com)

ON THE COVER Photograph of Bald Cypress, Mobile-Tensaw Delta, Alabama. Photograph courtesy Hunter Nichols (http://www.hnproductions.com). A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Natural Resource Report NPS/NRSS/BRD/NRR—2016/1243

Edited by: Gregory A. Waselkov1, C. Fred Andrus2, Glenn E. Plumb3

1Professor of Anthropology, Director Center for Archaeological Studies University of South Alabama 6052 USA Drive South Mobile, AL 36688

2Associate Professor of Geology, Chair Department of Geological Sciences University of Alabama Tuscaloosa, AL 35487

3Chief Wildlife Biologist National Park Service Biological Resources Division, Wildlife Conservation Branch 1201 Oak Ridge Drive, Suite 200 Fort Collins, CO 80525

Design by: Tani Hubbard National Park Service 12661 E. Broadway, Blvd. Tucson, AZ 85748

July 2016

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Please cite this publication as:

Waselkov G. A., C. F. Andrus, and G. E. Plumb, editors. 2016. A state of knowledge of the natural, cultural, and economic resources of the Greater Mobile-Tensaw River Area. Natural Resource Report NPS/NRSS/BRD/ NRR—2016/1243. Biological Resources Division, National Park Service, Fort Collins, Colorado.

NPS 909/133419, July 2016

ii A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Contents Page

Figures . . . . . �� ix Tables . . . . . �� xv Abstract . . . . . ��xvii Acknowledgments . . . . . ��xviii List of Acronyms . . . . . ��xviii Foreword Edward O. Wilson, Museum of Comparative Zoology, Harvard University . . . . . �� xix

INTRODUCTION . . . . . �� 1 1. Introduction Gregory A. Waselkov, University of South Alabama; C. Fred Andrus, University of Alabama; Glenn E. Plumb, National Park Service . . . . . �� 3

2. A Brief History of the Mobile-Tensaw River Bottomlands National Natural Landmark Glenn E.Plumb, National Park Service . . . . . �� 5 The National Natural Landmarks Program . . . . . �� 5 The Mobile-Tensaw River Bottomlands National Natural Landmark . . . . . �� 5 Further Study . . . . . �� 6 Summary . . . . . �� 7 Acknowledgments . . . . . �� 9

PHYSICAL LANDSCAPE . . . . . �� 11 3. Geology of the Lower Mobile River Basin Antonio B. Rodriguez, University of North Carolina at Chapel Hill . . . . . �� 13 Geologic Setting of the Lower Mobile River Basin . . . . . �� 13 Late Quaternary Sea Level Fluctuations and River Erosion . . . . . �� 15 Transgression and Valley Fill . . . . . �� 16 Mobile Bayhead Delta . . . . . �� 17

4. Hydrology and Water Quality in the Mobile-Tensaw Delta G. Milton Ward, Amelia K. Ward, and Alex Maestre, University of Alabama . . . . . �� 19 Geomorphology . . . . . �� 20 Delta Movement during the Holocene . . . . . �� 21 Hydrology . . . . . �� 22 Hydrological Alterations . . . . . �� 23 Water Quality . . . . . �� 24 Summary . . . . . �� 25

Contents iii Contents (continued) Page

BIOTIC LANDSCAPE . . . . . �� 27 5. Paleobotany and Paleoclimate Debra Z. Stults and Brian J. Axsmith, University of South Alabama . . . . . �� 29 Mobile River Terrace (Late Pleistocene—85,000 to 82,000 Thousand Years Ago) . . . . . �� 29 The Citronelle Formation Flora (Late Pliocene—3.4 to 2.7 Million Years Ago) . . . . . �� 30 Mauvilla Flora (Late Miocene—7.3 to 6.8 Million Years Ago) . . . . . �� 32 Bucatunna Formation (Early Oligocene—30.5 to 30.3 Million Years Ago) . . . . . �� 33 Gosport Formation, Claiborne Group (Middle Eocene—41 to 38 Million Years Ago) . . . . . �� 33 Summary . . . . . �� 33

6. Landscape and Disturbance Ecology of the Greater Mobile-Tensaw River Area—Geologic, Hydrologic, and Pyric Patterns and Processes William A. Finch, Earthword Services; and Glenn E. Plumb, National Park Service . . . . . �� 35

Overview of Ecosystems of the Greater Mobile-Tensaw River Area . . . . . �� 35 Geological and Geomorphological Patterns and Processes . . . . . �� 36 Hydrologic and Precipitation Patterns and Processes . . . . . �� 37 Fire . . . . . �� 39 Vulnerabilities and Challenges . . . . . �� 41

7. Ecoregional Basis for Biodiversity Richness and Endemism R. Scot Duncan, Birmingham Southern University . . . . . �� 43

Biogeographic Realms . . . . . �� 43 Climate . . . . . �� 43 Ecological Change . . . . . �� 44 Subregions Within the Greater Mobile-Tensaw River Area . . . . . �� 44 Anthropogenic Change . . . . . �� 46

8. Vegetation Communities L. J. Davenport, Samford University . . . . . �� 47

William Bartram’s Lyrical Descriptions . . . . . �� 47 Descriptions of Charles T. Mohr . . . . . �� 47 Later Studies . . . . . �� 52 The Delta Today . . . . . �� 53 Access . . . . . �� 53

9. Exploring the Unknown—The Insects of the Mobile-Tensaw River Delta John W. McCreadie and Peter H. Adler, University of South Alabama . . . . . �� 57

iv A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Contents (continued) Page

What Is Known of Insects in the Delta? . . . . . �� 57 The Total Insect Bio-inventory Project of the Mobile-Tensaw Delta . . . . . �� 58 Sampling and Collecting . . . . . �� 58 Insects . . . . . �� 59 Summary—More than 800 Species and Counting . . . . . �� 65 Acknowledgments . . . . . �� 65

10. Vector-Borne Disease Systems Nathan Burkett-Cadena, Florida Medical Entomology Laboratory, University of Florida . . . . . �� 67

Yellow Fever and the Greater Mobile-Tensaw River Area . . . . . �� 67 Vector-borne Diseases and the Greater Mobile-Tensaw River Area Today . . . . . �� 68 Tick as Vectors of Human Pathogens . . . . . �� 69 Concluding Thoughts . . . . . �� 70

11. Freshwater Mollusks of the Greater Mobile-Tensaw River Area Michael M. Gangloff, Appalachian State University . . . . . �� 73

Geographical Scope . . . . . �� 73 Historical Surveys . . . . . �� 73 Recent Surveys . . . . . �� 75 Mollusk Assemblage Structure . . . . . �� 76 Freshwater Bivalves . . . . . �� 80 Freshwater Gastropods . . . . . �� 81 Current Status and Threats to Mollusk Resources . . . . . �� 83 Author’s Note . . . . . �� 84

12. Crustaceans James A. Stoeckel and Brian S. Helms, Auburn University . . . . . �� 85

Current Knowledge—Natural, Cultural, and Economic Importance . . . . . �� 85 Key Drivers and Stressors . . . . . �� 88 Resource Conservation Implications . . . . . �� 92 13. Fishes Dennis R. DeVries and Russell A. Wright, Auburn University . . . . . �� 93

The Landscape for Fishes . . . . . �� 93 Fishes in the Greater Mobile-Tensaw River Area . . . . . �� 94 Salinity . . . . . �� 98

Contents v Contents (continued) Page

Threats to Fishes in the Lower Alabama River and Mobile-Tensaw River Delta . . . . . �� 98 Resource Conservation Implications . . . . . �� 99

14. Reptiles and Amphibians Jen Williams, National Park Service; JJ Apodaca, Warren Wilson College; and Craig Guyer, Auburn University . . . . . �� 101

Reptile Conservation . . . . . �� 101 Amphibian Conservation . . . . . �� 108 Thwarting Extinction . . . . . �� 109

15. Avifauna of the Lower Alabama River and Mobile-Tensaw Delta Joel A. Borden and C. Smoot Major, University of South Alabama . . . . . �� 111

Significance to Avifauna of the Greater Mobile-Tensaw River Area . . . . . �� 111 Species Overview . . . . . �� 112 Species of Special Concern . . . . . �� 116 Neotropical Migrants . . . . . �� 119 Migration Corridor . . . . . �� 119 Impacts of Birding . . . . . �� 119 Conservation Implications . . . . . �� 119 Summary . . . . . �� 120

16. Mammals Troy L. Best, Auburn University . . . . . �� 121

Overview . . . . . �� 121 Opossums, Manatees, and Armadillos . . . . . �� 121 Rodents . . . . . �� 123 Hares, Pikas, and Rabbits . . . . . �� 124 Shrews, Moles, Desmans, and Relatives . . . . . �� 124 Bats . . . . . �� 124 Carnivores . . . . . �� 125 Even-toed Ungulates . . . . . �� 126 Ever-increasing Threats . . . . . �� 126

CULTURAL LANDSCAPE . . . . . �� 129 17. Archeology—Arrival to AD 1550 David W. Morgan, National Park Service . . . . . �� 131

vi A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Contents (continued) Page

Bottle Creek . . . . . �� 131 Little Lizard Creek . . . . . �� 135 Earliest Human Use of the Greater Mobile-Tensaw River Area . . . . . �� 135 Life in the Greater Mobile-Tensaw River Area during the Woodland Stage . . . . . �� 137 Summary . . . . . �� 139

18. Archeology and History, AD 1550 to 1950 Gregory A. Waselkov, University of South Alabama . . . . . �� 141

Evidence of the Earliest Mobilians . . . . . �� 141 Establishment of the Louisiane Colony and Old Mobile . . . . . �� 141 Salt and Trade . . . . . �� 142 African Slaves . . . . . �� 143 The British Colonial Period . . . . . �� 143 From British to Spanish to US Control . . . . . �� 145 The Redstick War and Alabama Fever . . . . . �� 145 Native Americans in the Greater Mobile-Tensaw River Area . . . . . �� 146 Impact of the Civil War . . . . . �� 147 Following the Civil War . . . . . �� 147 The Logging Economy . . . . . �� 148 On-going Connections . . . . . �� 149

19. Modern Cultures of the Mobile-Tensaw Delta Ben Raines, Independent Environmental Reporter . . . . . �� 151

The People of the Delta . . . . . �� 151 Stories from Delta Locals . . . . . �� 151 “Delta Rats” . . . . . �� 152 Looking to the Future . . . . . �� 153 Suggested Readings . . . . . �� 154

20. National Historic Landmarks David B. Schneider, Schneider Historic Preservation; Bonnie L. Gums, University of South Alabama; Gregory A. Waselkov, University of South Alabama; George W. Shorter, Jr., retired archeologist, Mo- bile, Alabama; Richard S. Fuller, retired archeologist, Mobile, Alabama . . . . . �� 155

Bottle Creek Site (Archeological Site 1BA2; 1994 NHL Designation) . . . . . �� 155 Government Street Presbyterian Church (1992 NHL Designation) . . . . . �� 157 Fort Morgan (1960 NHL Designation) . . . . . �� 159

Contents vii Contents (continued) Page

Mobile City Hall (1973 NHL Designation) . . . . . �� 161 USS Alabama (1986 NHL Designation) . . . . . �� 163 USS Drum (1986 NHL Designation) . . . . . �� 165 Summary . . . . . �� 166

21. A Socioeconomic Overview of the Greater Mobile-Tensaw River Area Ermanno Affuso, University of South Alabama . . . . . �� 167

Socioeconomic Analysis . . . . . �� 168 Overview of Economic Activities . . . . . �� 168 Major Economies of the Greater Mobile-Tensaw River Area . . . . . �� 169 Minor Economies of the Greater Mobile-Tensaw River Area . . . . . �� 175 Summary . . . . . �� 176

22. Conservation and Tourism Development in the Greater Mobile-Tensaw River Area Michele L. Archie, The Harbinger Consulting Group . . . . . �� 177

Tourism is Underdeveloped in the Greater Mobile-Tensaw River Area . . . . . �� 178 Factors Contributing to Underdevelopment of Tourism in the Greater Mobile-Tensaw River Area . . . . . �� 178 Opportunities to Align Conservation and Tourism/Economic Development . . . . . �� 179 Potential Economic Benefits of the Bartram Canoe Trail . . . . . �� 181 Managing for Shared Benefits . . . . . �� 181 Inhabiting the Ecosystem . . . . . �� 182

SYNTHESIS . . . . . �� 183 23. A Greater Mobile-Tensaw River Area—Connections and Consequences Glenn E. Plumb, National Park Service; Gregory A. Waselkov, University of South Alabama; C. Fred Andrus, University of Alabama; Bill Finch, Earthword Services . . . . . �� 185 Lay of the Land . . . . . �� 185 As Goes the Water . . . . . �� 186 Connectivity, the Whys and Wherefores . . . . . �� 187 Historical Connections . . . . . �� 189 A Greater Mobile-Tensaw River Area . . . . . �� 192

Literature Cited . . . . . �� 195 List of Contributors . . . . . �� 233

viii A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figures Page

Figure 2-1. Boundary delineation of the Mobile-Tensaw River Bottomlands National Natural Landmark...... �� 8 Figure 3-1. Geologic map showing the different geologic provinces (arrows) and the scale of the Mobile River drainage basin, which comprises the Tombigbee and Alabama river valleys (above)...... �� 13 Figure 3-2. Digital elevation model of the Lower Mobile River Basin...... �� 14 Figure 3-3. Sea level curve for the northern Gulf of Mexico after Anderson et al. (2004)...... �� 15 Figure 3-4. Cross sections through the lower Mobile River Basin and Mobile Bay showing two stacked erosional surfaces that formed during periods of river incision...... �� 16 Figure 4-1. Mobile River Basin and river network...... �� 19 Figure 4-2. Mobile-Tensaw Delta landscape...... �� 21 Figure 4-3. Monthly mean, minimum, and maximum discharge (in ft3/s or cfs) into the Mobile River based on summation of discharge from lower Alabama and lower Tombigbee rivers, 1976-2014...... �� 22 Figure 5-1. Representative plant fossils from a Pleistocene terrace deposit in Mobile County...... �� 29 Figure 5-2. Representative Citronelle Formation plant fossils of taxa still living in the region today...... �� 30 Figure 5-3. Representative regionally extinct Citronelle Formation plant fossils . . . . . �� 31 Figure 5-4. Representative Mauvilla site plant fossils...... �� 32 Figure 5-5. Representative Bucatunna Formation fossils...... �� 33 Figure 8-1. Cypress-gum-pine border, Byrnes Lake Boat Landing north of Spanish Fort...... �� 48 Figure 8-2. Hardwood pine border, Byrnes Lake Boat Landing north of Spanish Fort...... �� 49 Figure 8-3. Marsh with Mobile skyline, Five Rivers Delta Resource Center, Spanish Fort...... �� 49 Figure 8-4. Marsh and cypress-gum edge, Historic Blakeley State Park, Spanish Fort...... �� 50 Figure 8-5. Boardwalk along Tensaw River marsh edge, Historic Blakeley State Park, Spanish Fort...... �� 50 Figure 8-6. Virginia salt-marsh mallow, Kosteletskya virginica, Bayfront Park, Daphne...... �� 51 Figure 8-7. Pickerel-weed, Pontederia cordata, Bayfront Park, Daphne...... �� 51 Figure 8-8. Beach morning-glory, Ipomoea imperati, on dry sand spit, Village Point Park Preserve, Daphne...... �� 52 Figure 8-10. The Jackson Oak, Village Point Park Preserve, Daphne...... �� 53 Figure 8-9. Water’s edge, Byrnes Lake Boat Landing north of Spanish Fort...... �� 53 Figure 8-12. American alligator, Alligator mississippiensis, viewed from boardwalk, Bayfront Park, Daphne...... �� 54 Figure 8-11. Warning sign, Village Point Park Preserve, Daphne...... �� 54 Figure 8-13. E. O. Wilson Boardwalk, Historic Blakeley State Park, Spanish Fort...... �� 55 Figure 9-1. The water scorpion Ranatra with a barnacle attached to its thorax...... �� 57 Figure 9-2. Malaise trap, the primary means of collecting insects in the Delta...... �� 58 Figure 9-3. Larva of the blue dasher dragonfly (Pachydiplax longipennis)...... �� 59 Figure 9-4. A species of the genus Anthomyia (family Anthomyiidae)...... �� 60

Contents ix Figures (continued) Page

Figure 9-5. Green bottle flies (family Calliphoridae) depositing eggs on a dead snake...... �� 60 Figure 9-6. A biting midge of the genus Palpomyia (family Ceratopogonidae) that preys on other insects...... �� 61 Figure 9-7. A male nonbiting midge (family Chironomidae)...... �� 61 Figure 9-8. A typical metallic long-legged fly (family Dolichopodidae)...... �� 62 Figure 9-9. Flies in the same family as the house fly (family Muscidae)...... �� 62 Figure 9-10. A moth fly (family Psychodidae)...... �� 63 Figure 9-11. A black fly (family Simuliidae) that feeds on the blood of mammals...... �� 63 Figure 9-12. A member of the large family of parasitoid flies (family Tachinidae)...... �� 64 Figure 9-13. A typical crane fly (family Tipulidae)...... �� 64 Figure 10-1. Historic marker at Old Church Street Cemetery, Mobile, AL, commemorating the interment of early residents of the greater Mobile-Tensaw River area who fell victim to vector-borne diseases, particularly yellow fever...... �� 67 Figure 10-2. Transmission cycle of a mosquito-borne disease: Eastern equine encephalitis...... �� 69 Figure 10-3. Medically important tick species of the greater Mobile-Tensaw River area . . . . . �� 69 Figure 10-4. Life cycle of the lone star tick, Amblyomma americanum...... �� 70 Figure 11-1. Alabama River main channel habitat looking upstream from Claiborne Landing, Monroe County, Alabama...... �� 73 Figure 11-2. Little River headwaters, Claude D. Kelly State Park...... �� 74 Figure 11-3. Bassett Creek near Jackson, Alabama is a fast-moving, gravel-bottomed tributary to the Tombigbee River...... �� 74 Figure 11-4. Leatherwood Creek is more typical of low-gradient swampy channels in the Salt Springs region of Clarke County...... �� 75 Figure 11-5. Limestone Creek on the Fred Stimpson Sanctuary is characterized by a highly incised and high-gradient, third-order channel with relatively coarse gravel substrates...... �� 75 Figure 11-6. Floodplain wetland or backwater habitat adjacent to the main stem of the Tombigbee River on Fred Stimpson Sanctuary, Clarke County, Alabama...... �� 76 Figure 11-7. Lampsilis straminea from Bassett Creek, Clarke County, Alabama...... �� 81 Figure 11-8. Quadrula apiculata collected from Donnelley Reservoir, an impoundment on the Alabama River in Wilcox County...... �� 81 Figure 11-9. Plectomerus dombeyanus collected from Donnelley Reservoir, an impoundment on the Alabama River in Wilcox County...... �� 82 Figure 11-10. Anodontoides radiatus from a tributary of the Alabama River in Wilcox County...... �� 84 Figure 12-1. Select crayfish from the Mobile-Tensaw Delta, lower Alabama River, lower Tombigbee, and adjacent watersheds...... �� 87 Figure 12-2. Connectivity between aquatic and semi-terrestrial habitats is critical for crayfishes...... �� 88

x A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figures (continued) Page

Figure 12-3. Ohio Shrimp (Macrobrachium ohione) male (left) and female (right)...... �� 89 Figure 12-4. Importance of linkages between freshwater, brackish, and saltwater habitats for crabs and shrimp as illustrated by life history-stage migrations...... �� 90 Figure 13-1. Byrne’s Boat Ramp—a typical backwater habitat in the greater Mobile-Tensaw River area at the mouth of a blackwater tributary to the main channel...... �� 93 Figure 13-2. Cypress root habitat—a complex freshwater swamp habitat created by bald cypress knees...... �� 93 Figure 13-3. Cypress root habitat—a lowland swamp and marsh habitat. Note the high water marks on the cypress trunks...... �� 94 Figure 13-4. Lower delta habitat—extensive shallow water marshes with both emergent and submersed vegetation are the primary habitat type in the lower Delta just above the causeway, at the head of Mobile Bay...... �� 94 Figure 13-5. D’Olive Bay—emergent grasses are present at the lowermost end of the Delta, at the head of Mobile Bay...... �� 95 Figure 13-6. Millers Ferry Lock and Dam—showing the gated spillways to the left and the navigational lock chamber to the right...... �� 95 Figure 13-7. Large largemouth bass—this example of a largemouth bass from the Mobile-Tensaw Delta weighs more than 2.3 kg (5 lb)...... �� 97 Figure 13-8. Southern flounder—collected in the Delta...... �� 97 Figure 13-9. Two red drums—sometimes called “redfish”, collected from the Delta...... �� 98 Figure 13-10. Excessive sediment from erosion is considered among the most important non-point source pollutants in Alabama...... �� 100 Figure 13-11. Sunset on the Mobile-Tensaw River Delta...... �� 100 Figure 15-1. Purple Gallinule hatchling...... �� 111 Figure 15-2. Flight of Blue-Winged Teal...... �� 112 Figure 15-3. Male Wood Duck...... �� 112 Figure 15-4. Reddish Egret...... �� 113 Figure 15-5. Green Heron...... �� 113 Figure 15-7. Black-and-White Warbler...... �� 114 Figure 15-6. Prothonotary Warbler...... �� 114 Figure 15-8. Carolina Chickadee...... �� 115 Figure 15-10. Pileated Woodpecker...... �� 115 Figure 15-9. White-Crowned Sparrow...... �� 115 Figure 15-11. Osprey...... �� 116 Figure 15-12. Barred Owl...... �� 116 Figure 15-13. Swallow-tailed Kite...... �� 117

Contents xi Figures (continued) Page

Figure 15-14. White Ibis...... �� 119 Figure 16-1. Virginia opossum (Didelphis virginiana) is a common species in the greater Mobile-Tensaw River area of Alabama...... �� 121 Figure 16-2. The Rafinesque’s big eared bat (Corynorhinus rafinesquii) is a rare species that occupies caves, abandoned buildings, and hollow trees in the greater Mobile-Tensaw River area of Alabama...... �� 125 Figure 16-3. The long tailed weasel (Mustela frenata), once a common species throughout the south eastern United States, is currently uncommon in the greater Mobile-Tensaw River area and elsewhere in Alabama...... �� 125 Figure 17-1. Topographic map of the Bottle Creek mounds (1BA2) (Waselkov 1993)...... �� 131 Figure 17-2. Timeline of cultures in the greater Mobile-Tensaw River area. (adapted from Fuller 1998, p 4, Figure 2)...... �� 133 Figure 17-3. Field Sketch of the Crossroads Mound (1BA356) when initially recorded in 1998 by Richard S. Fuller, David W. Morgan, and Jack W. Morgan...... �� 134 Figure 17-4. Archeologist Richard S. Fuller, then of the Alabama Museum of Natural History, standing next to the Bryant’s Landing 4 site (1BA175) site...... �� 136 Figure 17-5. Example of a small, circa AD 400 to 700 Weeden Island pot excavated from the Late Woodland period Seymour (Parkway) Mound (1BA80) complex in the 1930s by the Alabama Museum of Natural History...... �� 138 Figure 17-6. The Briar Lake Slough Site (1BA270) prior to excavation in 1998...... �� 139 Figure 18-1. Detail from the French map titled “Carte d’une partie du cours de la riviere de La Mobille et de celle des Chicachas,” attributed to Valentin Devin, circa 1725...... �� 142 Figure 18-2. “Salines, or Salt Springs, along the lower Tombigbee River [in modern day Clarke County], as drawn by Thomas Freeman, Surveyor General for Lands South of Tennessee, September 30, 1816” (Lowrie 1834)...... �� 143 Figure 18-3. Numbered long lot plantations in the Mobile-Tensaw Delta area, detail from “A Plan of part of the Rivers Tombecbe, Alabama, Tensa, Perdido & Scambia in the Province of West Florida,” by David Taitt, circa 1771...... �� 144 Figure 18-4. “Scene on the Alabama River, Loading Cotton.” From Ballou’s Pictorial 13(22), p 1, November 28, 1857...... �� 146 Figure 18-5. Partially excavated Civil War-era furnace with steamship boiler converted for brine boiling, Furnace AA, Upper Salt Works, Clarke County, Alabama, 2010...... �� 147 Figure 18-6. Four-yoke team of oxen pulling a high-wheel cart, or log cart, hauling a large clamped log from a longleaf pine forest near Mobile, Alabama...... �� 148 Figure 18-7. Turpentine gathering containers and dip iron, collected by Robert Leslie Smith from the Mobile-Tensaw Delta...... �� 149 Figure 20-1. Location of Bottle Creek Site National Historic Landmark, Baldwin County, Alabama . . . . . �� 155 Figure 20-2. Topographic map of Bottle Creek (Waselkov 1993)...... �� 156

xii A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figures (continued) Page

Figure 20-3. Locations of National Historic Landmarks in Mobile, Mobile County, Alabama: Government Street Presbyterian Church, Mobile City Hall (Southern Market), USS Alabama, USS Drum . . . . . �� 157 Figure 20-4. South façade detail, Government Street Presbyterian Church, Mobile, Mobile County, Alabama...... �� 158 Figure 20-5. Interior sanctuary, Government Street Presbyterian Church, Mobile, Mobile County, Alabama...... 158 Figure 20-6. Location of Fort Morgan National Historic Landmark, Baldwin County, Alabama . . . . . �� 159 Figure 20-7. Entrance at north rampart, Fort Morgan, Baldwin County, Alabama...... �� 160 Figure 20-8. Interior vaulted casemates, Fort Morgan, Baldwin County, Alabama...... �� 160 Figure 20-9. Officer’s quarters, Fort Morgan, Baldwin County, Alabama...... �� 161 Figure 20-10. Mobile City Hall, Mobile County, Alabama...... �� 162 Figure 20-11. Interior first-floor lobby of the History Museum of Mobile, in Mobile City Hall, with 1936 Works Progress Administration murals by John Augustus Walker...... �� 162 Figure 20-12. The USS Alabama...... �� 163 Figure 20-13. The USS Alabama...... �� 163 Figure 20-14. Galley of the USS Alabama...... �� 164 Figure 20-15. USS Drum at Battleship Alabama Memorial Park, Mobile, Alabama...... �� 164 Figure 20-16. View across the foredeck of USS Drum...... �� 165 Figure 20-17. After torpedo room of USS Drum...... �� 165 Figure 21-1B. Demography of the study area...... �� 167 Figure 21-1A. Geography of the study area...... �� 167 Figure 21-2. A disaggregated geospatial analysis of A) per capita income and B) unemployment rate for the region...... �� 168 Figure 21-3. Geographic distribution of industrial sectors...... �� 172 Figure 21-4. Metropolitan area in Mobile-Tensaw watershed...... �� 172 Figure 21-5. Macroeconomic indicators of Mobile metropolitan area: A) nominal and real GDP expressed in 2009 constant dollars; B) inflation rate and economic growth rate from 2002 to 2013...... �� 173 Figure 21-6. Comparison of the industry of A) Mobile metropolitan area, and B) Daphne-Fairhope-Foley micropolitan area...... �� 174 Figure 21-7. Total value of agricultural production in constant 2012 dollars...... �� 175 Figure 22-1. Most popular recreational activities based on visitor studies in 4 water- and wetlands- focused National Park Service areas . . . . . �� 180 Figure 23-1. The greater Mobile-Tensaw River area...... �� 188 Figure 23-2. A cast of a flint clay stone pipe, created in the form of a crayfish effigy at Cahokia in 11th century AD and discovered in 1922 by Theodore R. Vaughan, Sr., on Dauphin Island...... �� 190

Contents xiii Figures (continued) Page

Figure 23-3. A stylized crayfish effigy on the rim of a Weeden Island Punctated ceramic vessel, Bayou St. John site (1BA21), Late Woodland period, Orange Beach, Alabama...... �� 190 Figure 23-4. Photograph of Cudjo Lewis, enslaved in Africa and transported to Mobile in 1861 aboard the Clotilda, at home in Africatown, circa late 1920s...... �� 191 Figure 23-5. “Mobile, taken from the marsh opposite the city near Pinto’s residence,” watercolor by William Todd, 1841...... �� 193

xiv A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Tables Page

Table 4-1. Annual minimum and maximum flux of nitrate-N, ammonium-N, and orthophosphate-P (103 kg/yr [103 lb/yr]) for selected time periods in the lower Alabama (USGS 02428400) and lower Tombigbee rivers (USGS 02469762) ...... �� 24 Table 11-1. Systematic list of freshwater bivalve taxa collected, conservation status, and habitats occupied during historical (1900-1990) and recent (1990-2015) freshwater mollusk surveys in the greater Mobile-Tensaw River area...... �� 77 Table 11-1. Systematic list of freshwater bivalve taxa collected, conservation status, and habitats occupied during historical (1900-1990) and recent (1990-2015) freshwater mollusk surveys in the greater Mobile-Tensaw River area . . . . . �� 78 Table 11-2. Systematic list of freshwater gastropod taxa collected, conservation status, and habitats occupied during historical (1900-1990) and recent (1990-2015) freshwater mollusk surveys in the greater Mobile-Tensaw River area...... �� 79 Table 12-1. Crayfish found in the Mobile-Tensaw Delta (MD), lower Alabama River (AL), Lower Tombigbee (LT), and adjacent local watersheds...... �� 86 Table 13-1. List of common names and scientific names of organisms referenced in the text...... �� 96 Table 14-1. Reptile and amphibian species documented (D), likely to occur (L), or previously occurring (i.e., extirpated [E]) in the greater Mobile-Tensaw River area and their federal, state, and Interna- tional Union for Conservation of Nature (IUCN) classifications...... �� 102 Table 14-2. Federal, State, and International Union for Conservation of Nature (IUCN) classification definitions for reptiles and amphibians...... �� 106 Table 15-1. Ranking species is conducted by NatureServe on Global (G rank), National (N rank), and a State rank (S rank) and are considered very reliable by most scientists, government agencies, and conservationists...... �� 117 Table 15-2. Species rankings for the IUCN red list (IUCN 2012)...... �� 118 Table 15-3. High conservation state of Alabama concern wetland species within the greater Mobile- Tensaw River area...... �� 118 Table 15-4. High conservation state of Alabama concern for species inhabiting uplands adjacent to swamps and bottomlands of the greater Mobile-Tensaw River area...... �� 118 Table 16-1. Mammals of the greater Mobile-Tensaw River area of Alabama (Best and Dusi 2014) and a qualitative estimate of their current status in the region...... �� 122 Table 16-1. Mammals of the greater Mobile-Tensaw River area of Alabama (Best and Dusi 2014) and a qualitative estimate of their current status in the region, cont...... �� 123 Table 21-1. Business pattern in the greater Mobile-Tensaw River area: number of establishments in 2013...... �� 170 Table 21-2. 2013 Annual payroll (expressed in thousands of dollars) of industrial sectors in the greater Mobile-Tensaw River area, shown by county...... �� 171 Table 22-1. Concentration of jobs in tourism-related industries, 2013...... �� 178 Table 22-2. Everglades National Park, visitor group participation in selected guided and self-guided activities for winter and spring 2008...... �� 180

Contents xv

Abstract

Red Hills, Monroe County, Alabama. Photograph courtesy of Hunter Nichols.

From the convergence of the Alabama and Tombigbee rivers in southern Alabama, arise the sister Mobile and Tensaw Rivers, and one of the great natural and cultural wonders of North America. The Greater Mobile-Tensaw River Area has been called “America’s Amazon” because of its obvious natural bounty, wondrous complexity, and profound diversity. Dramatic bluff lands and pinelands plunge down to the Mobile-Tensaw Bottomlands and Delta comprised of ever-changing levees, islands, channels, and bayous. The region’s unique geology and hydrology underpin its dynamic biotic systems as, likewise, do associated ecological processes that range from the lingering influences of ancient and far-off continental glaciation to the daily rise and fall of tides and changes in water salinity. Flora and fauna of the area are at once fragile and bountiful—the area contains many endangered, threatened, and special concern species, but also tree species diversity that ranks among the highest in North America, a diverse assemblage of freshwater crustaceans, over 200 species of birds, and likely the greatest turtle diversity in the world. The Greater Mobile-Tensaw River Area is steeped in human history, as well, from the originating Native American tribes and the mysteries of their ancient mounds awaiting exploration; through the area’s critical role in European settlement of America and, later, the American Civil War; through today’s cultural and economic vibrancy of its adjacent urban center, Mobile. Here people’s lives are woven among the dynamic rhythms of the area’s lands and waters. Yet, much remains elusive about how the area’s places are connected ecologically, socially, and economically. This holistic volume combines science, natural and cultural history, economics, and personal reflection to call attention to the connectivity of the area, to acknowledge challenges from human encroachment, and to serve as a foundation for a discussion of shared ecological, cultural, and economic stewardship of the Greater Mobile-Tensaw River Area.

Contents xvii Acknowledgments

We are thankful for knowledge, advice, aid, and encouragement from numerous people in the preparation of this report. Especially we acknowledge Dr. Leslie Rissler who while at the University of Alabama helped develop and initiate this undertaking. Elaine Leslie, Dr. Herbert Frost, Dr. Gary Machlis, Linda Drees, and Darrell Echols with the National Park Service provided counsel, encouragement, and programmatic support for the project. Dr. Robert F. Olin, Dean of the College of Arts and Sciences at the University of Alabama provided leadership and encouragement in developing cooperative relationships with the National Park Service and the University of South Alabama that served as the foundation of this project. Dr. Paula Ehrlich, President and CEO of the E.O. Wilson Biodiversity Foundation shared funding support and a powerful vision of possibilities. Mrs. Mary Ivy Burks of The Alabama Conservancy and Mrs. Verda Horne of the Alabama League of Women Voters more than 40 years ago were vocal champions of conservation of the Mobile-Tensaw delta and bottomlands. Hunter Nich- ols very kindly provided photographs that illustrate the beauty and wonder of the Greater Mobile-Tensaw River Area. Finally, we gratefully acknowledge the conscientious and skillful technical editing of the entire report by Scott Bischke of MountainWorks, Inc. (www.emountainworks.com).

List of Acronyms

ADCNR—Alabama Department of Conservation and Natural Resources

IUCN—International Union for the Conservation of Nature

NHL—National Historic Landmark

NNL—National Natural Landmark

NOAA—National Oceanic and Atmospheric Administration

NPS—National Park Service

USDA—United States Department of Agriculture

USEPA— United States Environmental Protection Agency

USFWS—United States Fish and Wildlife Service

USGS—United States Geological Survey

xviii A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Foreword Edward O. Wilson, Museum of Comparative Zoology, Harvard University

Mobile-Tensaw River Delta, Alabama. Photograph courtesy of Hunter Nichols.

1 March 2016

The greater Mobile-Tensaw River area, containing the two powerful rivers feeding between them, inward Delta and the Red Hills immediately to the north, is from shoreline bluffs, a labyrinth of streams, oxbow both uniquely beautiful and one of the least known lakes, freshwater and brackish marshes, flooded natural areas in America. Although located on the flat woodland, and raised islets of secure ground. The Gulf of Mexico plain, its habitats match—or exceed— Delta is a miniature wilderness. Even with a boat, in diversity those in any of America’s established map, and GPS, travel is difficult. national parks. In the Red Hills, ridges with remnant longleaf pine savannas drop to deep, cool ravines that The Delta may also be the easiest natural land in Amer- harbor relict “ice-age” floras left behind during the ica in which to get lost. Yet at the same time it is the retreat of the last continental glacier. Southward they most accessible of any known wildland. Its bayfront give way to the Mobile-Tensaw Delta, second largest border, lined with a causeway, can be reached by auto- delta in America after the Mississippi, enclosed by the mobile from downtown Mobile in 15 minutes.

Contents xix Because of the great variety of the habitats in which Actually, my attachment to this immediate area runs they live, and their still mostly undisturbed condi- even more deeply. When I was a boy, living in Mobile tion, the fauna and flora of the greater Mobile-Tensaw and also nearby Brewton, Alabama, I fell in love with River area may well be most diverse of comparable what has come to be called “America’s Amazon.” I was geographical areas in North America. The 28+ species predisposed to do so because Mobile was my ances- of oaks in the Red Hills, for example, is evidently a tral home. My forebears were among the settlers of national record. The number of turtle species is one of Mobile County and adjacent Baldwin County during the highest in the world. The full roster of plants and the American era, from its 1819 statehood. They all animals includes many endangered species. Others worked close to and around the Delta, variously as are certain to exist still unknown to science. merchants, marine engineers and bar pilots on the Bay. One couple helped establish the doomed community The pre-European human history of the Mobile- at Blakeley. After its abandonment, they moved with Tensaw Delta has also just begun to yield its secrets. their infant daughter to the village of Mobile. Archaeological evidence exists of settlements that go back thousands of years, with a likelihood of incep- The Delta had a powerful influence on my develop- tion during the Late Archaic Period, at least 6000 years ment as a scientist. As I roamed around its edges, before the present. The best and most striking settle- collecting butterflies and snakes, I dreamed of some- ment in Lower Alabama is the Delta’s Bottle Creek day taking an expedition into the interior to discover site, which was occupied between AD 1250 and 1550. all of the insects and other animals. Now I understand Still mostly unexcavated by archaeologists, it contains that it will take a lot of expeditions. a high mound facing a plaza, along with smaller residential mounds scattered about. The largest of its kind Remarkably little has changed in the Red Hills and on the central Gulf Coast, it appears to have been the Mobile-Tensaw Delta since my boyhood in the 1940s. headquarters of elites who controlled the population Industries and housing are pressing in on the habit- for miles inside the Delta and to the Gulf. able edges, and the pollution of the river waters is rising inexorably. The populations of Mobile and the And there is substantial history as well. Through the Bay coasts are growing and spreading rapidly. Yet for a 1820s at the edge of the Tensaw River, Blakeley, a while longer this small wilderness will persist, a price- frontier boomtown flourished, only to be dispersed less window on deep history, and a national treasure following two yellow fever epidemics. It was then at multiple levels. replaced by Fort Blakeley, the site of the last major battle of the Civil War, fought unknowingly on the The value of the greater Mobile-Tensaw River area same day as Lee’s surrender at Appomattox. The is great, unique, and if lost cannot be recreated. Is Confederate ramparts still stand, untouched to the there any other place in America where residents and present day. I am personally involved a bit in these visitors can live in a modern city and yet travel into battlements, since one of my great-grandfathers, a an authentically wild area within an hour? Or where Confederate artilleryman, escaped capture by fleeing students can experience a natural subtropical envi- across the Delta to reach his home in Mobile. ronment along with unaltered historical sites spanning 500 years in a single day’s excursion?

xx A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area INTRODUCTION

Alligator, Mobile-Tensaw Delta, Alabama. Photograph courtesy of Hunter Nichols.

Introduction - Chapter 1 1

1. Introduction Gregory A. Waselkov, University of South Alabama; C. Fred Andrus, University of Alabama; Glenn E. Plumb, National Park Service

This report is the product of a cooperative effort We have convened 35 contributing authors in an between the University of South Alabama, the Univer- unprecedented effort to assess the state of our knowl- sity of Alabama, and the National Park Service through edge on key natural, cultural, and economic conser- the Gulf Coast Cooperative Ecosystem Studies Unit, vation resources for the Mobile-Tensaw Delta, lower a consortium that facilitates cooperative science and Alabama and Tombigbee rivers, and adjacent contrib- scholarship between federal agencies and regional uting watersheds, blufflands, and pinelands. The area universities. The principal goal of contributors to under consideration begins at the head of Mobile Bay this report is to share with the public the state of our and extends north through the deltaic floodplain, knowledge on the natural, cultural, and economic including locally rising watersheds and streams, and resources of the greater Mobile-Tensaw River area. the forests sheltering those watersheds and streams, As defined for this project, this area encompasses up through the Red Hills region that overlooks the the large landscape and watershed that includes the northern extent of the Delta. This area—the greater Mobile-Tensaw Delta. Mobile-Tensaw river area—includes approximately the northeastern quadrant of Mobile County, the Alabama and the National Park Service have a strong northwestern quadrant of Baldwin County, most of history of cooperation via parks, national natural southern Clarke County, the southeastern corner landmarks, national historic landmarks, national of Washington County, the westernmost section of historic sites, and community assistance programs. Escambia County, and the southwestern quadrant of One National Natural Landmark (Mobile-Tensaw Monroe County. River Bottomlands) and 6 National Historic Land- marks (Fort Morgan, Mobile City Hall and South- The 23 chapters of this wide-ranging report will, we ern Market, USS Alabama, USS Drum, Govern- hope, enhance understanding and stewardship of ment Street Presbyterian Church, and Bottle Creek these natural and historic landmarks, and aid plan- Archeological Site) are located in or near the greater ning by the citizens and State of Alabama and its part- Mobile-Tensaw River area. The National Natural and ners. Among the contributors are many preeminent Historic Landmarks programs are administered by specialists who know their corners of the greater the National Park Service. Designation as National Mobile-Tensaw River area better than anyone else. Landmarks recognizes and encourages the conser- Here they introduce us to their topics in plain English vation of sites that contain outstanding natural and and share information not generally available outside cultural resources regardless of landownership type. of the technical literature. Every chapter illustrates the These are the only programs of national scope that area’s rich natural and cultural diversity, and identifies recognize the best examples of natural and cultural challenges and opportunities to preserve the integ- resources and values in both public and private rity and economic viability of these resources. This ownership. Partnerships among public and private report highlights the history of the Delta, its commu- land stewards allow participants to share information, nities and habitats, from the Delta’s geological origins solve problems cooperatively, and conserve outstand- through changes in land use by humans from earliest ing sites that illustrate the rich and diverse tapestry of to modern times. We have endeavored to share with the country’s natural and cultural landscapes. Integra- readers an up-to-date picture of the greater Mobile- tion of knowledge about and stewardship of natural Tensaw River area that can contribute to the area’s and cultural landscapes is crucial to preserving the biological health and sustainability by enhancing ecological and cultural integrity of our communities. understanding and stewardship of its resources, and inform conservation planning by Alabama and its partners.

Introduction - Chapter 1 3

2. A Brief History of the Mobile-Tensaw River Bottomlands National Natural Landmark Glenn E. Plumb, National Park Service

The National Natural Landmarks The Mobile-Tensaw River Bottomlands Program National Natural Landmark The National Natural Landmarks (NNL) Program was During the early 1970s, Richard Goodwin and established by the Secretary of the Interior in 1962, William Niering from Connecticut College conducted under authority of the Historic Sites Act of 1935 (16 a review of important inland wetlands of the United USC 461 et seq.), to identify and encourage the pres- States (see Goodwin and Niering 1975). By early ervation of the full range of geological and biological 1972, this comprehensive national study involved features that are determined to represent nationally discussion with Walter Beshears of the Alabama State significant examples of the nation’s natural heritage. Department of Conservation and Natural Resources Potential sites are evaluated by qualified scientists (Vance 1972; Mount 1973) regarding important and, if determined nationally significant, recom- inland wetlands of Alabama. Beshears had previously mended to the Secretary of the Interior for designa- co-published an overview of Alabama’s estuarine tion. Once a landmark is designated it is included on areas (Beshears and Bird 1959), including the Mobile- the National Registry of Natural Landmarks, which Tensaw River Bottomlands. Available records indicate currently includes nationally significant geological that by the early 1970s, it was Mr. Beshears who first and biological features in 48 states, American Samoa, suggested to Goodwin and Niering that the Mobile- Guam, Puerto Rico, and the US Virgin Islands. Tensaw River Bottomlands be considered for designation as an NNL (Vance 1972; Mount 1973; Goodwin National Natural Landmarks are nationally significant and Niering 1975). In response to this suggestion, sites owned by a variety of land stewards, and their Richard Vance, Park Naturalist with the Natchez participation in the NNL program is entirely volun- Trace Parkway, visited the area with Dr. M.W. Galland tary. National Natural Landmark designation is not a and Douglass Freeman on June 6, 1972, and engaged land withdrawal, does not change the ownership of a in additional correspondence with Dr. Arthur Garrett site, and does not dictate use, activity, or land acquisi- from the University of South Alabama and Mrs. Verda tion (NPS 2009). Of the nearly 600 NNLs designated D. Horne of Fairhope, Alabama (Vance 1972). In a to date and listed on the Registry, approximately one short report Vance (1972) described the highlights half are administered solely by public agencies (e.g., of the Mobile-Tensaw River Bottomlands and wrote federal, state, county, or municipal governments), that, “The area has problems. However, the tremen- nearly one third are owned entirely by private parties, dous amount of nationally significant natural values and the remaining are owned or administered by a offset most of the encroachments. Also, there are mixture of public and private owners. For the many many influential groups and people who are working natural landmarks that are privately owned or not to preserve this area.” He further concluded that, “It is managed for public access, owner permission must my recommendation that this site is eligible for inclu- be obtained prior to visitation. Designation in no sion in the National Registry of Natural Landmarks.” way confers any right of public access. The National Park Service (NPS) administers the NNL program Following Vance’s 1972 visit, the National Park Service and works in partnership with landmark owners to requested a formal evaluation by Dr. Robert H. Mount, encourage and advocate for conservation of landmark Professor of Zoology-Entomology at Auburn Univer- resources. Upon landowner request, the NPS can also sity, who had visited the area on numerous occasions. provide assistance for landowner conservation efforts. On June 28, 1973 Mount toured the area by boat with Dr. James Harper of Auburn University, and subsequently made contact with the 4 largest landowners in the area (Scott Paper Company, International Paper

Introduction - Chapter 2 5 Company, Baer and Company, and Alabama Public recommendations of the Advisory Board and, pursu- Lands Division). Mount also consulted with faculty ant to the Historic Sites Act of 1935, formally desig- colleagues from Auburn University, Mrs. Mary Ivy nated the Mobile-Tensaw River Bottomlands NNL Burks (Executive Secretary of The Alabama Conser- (NPS 1974c). vancy), and Mrs. Verda D. Horne (Conservation Chairman of the Alabama League of Women Voters). Further Study In his written report, Mount (1973) included detailed Public Law 94-458 (enacted October 7, 1976) directed descriptions of the location, size, ownership, land use, the Secretary of Interior “to investigate, study, and natural resource values, and threats to integrity of the continually monitor the welfare of areas whose area. He concluded that: resources exhibit qualities of national significance The delta of the Mobile bay drainage is one of and which may have potential for inclusion in the the most important wetland areas in the coun- National Park Service.” In the mid- to late-1970s state try. It contains a variety of habitats, ranging listings showed that 19 endangered or threatened from mesic floodplains and freshwater swamps animal species and 18 endangered plant species were to open, brackish-water marshes. Despite man’s known to be present in Mobile and Baldwin counties, intrusions into the area and his depositions of which encompass the Mobile-Tensaw River Bottom- pollutants in the streams that furnish its water, lands NNL (NPS 1979). In June 1977, an NPS field the delta still retains many of its natural values. inspection echoed the state’s findings and declared The area is, in my opinion, of considerable the Mobile-Tensaw River Bottomlands NNL “endan- national importance. Its designation as a Regis- gered.” It was subsequently listed as “threatened” in tered Natural Landmark will enhance efforts the 1977 NPS Report to Congress (see NPS 1979, p being made to protect it from further degrada- 5) on damaged and threatened natural landmarks. tion. Numerous references and knowledgeable Further action by NPS occurred a year later, in 1978, people were consulted during the study of the when the agency acted on the recommendation made Mobile Bay delta and in the preparation of this 4 years previously by the Advisory Board to conduct report. All of the individuals with whom I talked “further study to determine the best means of protect- concurred with the opinions expressed herein. ing the ecological values of the site” (NPS 1974b). The NPS undertook a “Study of Alternatives, Mobile- In April 1974, at its 70th meeting at Washington D.C., the Tensaw River Bottomlands” (NPS 1979) focused on Department of Interior Advisory Board on National two purposes: Parks, Historic Sites, Building and Monuments (here- ● to evaluate resource significance through devel- after, “Advisory Board”) identified 35 sites across the ● opment of a resource base, including maps, and United States eligible for designation as Registered report sections delineating natural resources, Natural Landmarks. Those identified included 3 sites cultural resources, water and air quality, land use in Alabama: Beaverdam Creek Swamp, the Dismals, and ownership, socioeconomic environment, and the Mobile-Tensaw River Bottomlands (NPS and regional context; and 1974a). On April 24, 1974 Lady Bird Johnson, Acting Chairman of the Advisory Board and wife of Presi- ●● to delineate preservation/management alterna- dent Lyndon Johnson, formally wrote to the Secretary tives in terms of specific institutional arrange- of Interior to recommend these 35 sites for official ments required to protect and properly utilize designation (NPS 1974a). She further informed the values of the area. Secretary of Interior, by a separate letter in that same capacity, “The Board recommends that the National The study involved extensive stakeholder consulta- Park Service conduct further study to determine the tion with federal, state, county, and non-governmen- best means of protecting the ecological values of this tal organizations, including specific contact with the site [i.e., the recommended Mobile-Tensaw River Director of the Alabama Division of Game and Fish, Bottomlands NNL]. This should be a cooperative and the Commissioner and Assistant Commissioner study carried out by the National Park Service and of the Alabama Department of Conservation and Bureau of Sport Fisheries and Wildlife” (NPS 1974b). Natural Resources. Other agencies and organizations On May 30, 1974, Secretary of Interior Roger C. B. contacted by the NPS included Alabama Air Pollution Morton under President Richard Nixon acted on the Control Commission; Alabama Coastal Area Board;

6 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Alabama Cooperative Extension Service; Alabama of the Mobile-Tensaw River Bottomlands NNL. Water Improvement Commission; US Department of These potential alternatives included (NPS 1979): Agriculture Forest Service; Department of Defense, Department of the Army; Corps of Engineers-Mobile 1. maintaining the status quo, with no change in District; Environmental Protection Agency; Mobile ownership and management; Bureau of Environmental Health; South Alabama 2. imposing state regulation, with no change in Regional Planning Commission; University of South ownership and management; Alabama; and 5 Alabama non-government organiza- 3. creating a state park in the core area of the tions (The Alabama Conservancy, Birmingham; Bald- Mobile-Tensaw River Bottomlands; win County Wildlife Association, Robertsdale; Bald- win County Wildlife and Conservation Association, 4. creating a state park under state management; Bay Minette; Mobile Audubon Society, Mobile; and 5. designating an Area of National Concern, with a the Mobile County Wildlife Association, Mobile). core area of federal ownership and mixed state and federal management; The NPS released the “Study of Alternatives, Mobile- Tensaw River Bottomlands” as a public document in 6. establishing a US Fish and Wildlife Service 1979, which at the time was the first integrated synthe- (USFWS) Refuge for most of the Mobile-Tensaw sis of natural resource, cultural resource, and socio- River Bottomlands NNL; economic information for the area (NPS 1979). The 7. establishing an NPS preserve, wherein hunting report included detailed and summary state-of-the- and fishing and other uses would continue under knowledge information, tables, figures, and maps, as state management; and well as key findings about resource status/condition. 8. establishing a mixed NPS/USFWS approach. According to this report (NPS 1979): While the report articulated potential alternatives for While the delta still retains its natural values to a long-term management of the resources in the land- significant degree, threats to the integrity of the mark area, it did not recommend any alternative for natural environment do exist. If these threats further action by any agency. According to National are left unchecked, it is very likely that the Natural Landmarks Program authorities, no further significant natural values would be lost, and the NPS action regarding the proposed alternatives was integrity upon which the Secretary originally taken on the report following release. The site remains based his determination of national significance a designated landmark and NNL Program staff have would be destroyed. maintained contact with various landowners at the The report identified the major threats to the areas as: site over time. Recently, in an effort to digitize all landmark boundaries nationwide, the NNL Program has 1. existing industrial development along the west augmented the descriptions by Mount (1973) and bank of the Mobile River with further expansion NPS (1979) with aerial photo interpretation and, as planned northward and along the Tensaw River, a consequence, clarified that the designated Mobile- which would produce cumulative degradation; Tensaw River Bottomlands NNL includes approxi- 2 2. cumulative effects of pollution from silt, agricul- mately 178,000 acres (720 km ) (Figure 2-1; NPS tural chemicals, and to a lesser degree industrial 2011). waste as an ongoing problem; Summary 3. potential widespread expansion of timber harvesting practices; It has been over 40 years since the Mobile-Tensaw River Bottomlands was championed by Alabama citi- 4. uncontrolled wildlife harvest that reduces some zens and scientists as an area of national significance species; and leading to National Natural Landmark designation. 5. cumulative effects from dredging and spoilage. Since this designation, Alabama conservation agencies, Alabama academic scientists and scholars, Alabama public and private conservationists, and popular The report included descriptions of potential alterna- media have continued to highlight the extraordinary tives and various methods of preserving the resources values and characteristics of the Mobile-Tensaw River

Introduction - Chapter 2 7 MOBILE-TENSAW RIVER BOTTOMLANDS Landmark Boundary NATIONAL NATURAL LANDMARK

Baldwin, Mobile, and Washington Counties, Alabama Data Source: NPS NNL Data

Map Produced Jan 2016 0 2.5 5 by National Park Service Intermountain Region Calculated Acreage: 178,828.6 Miles ± Geographic Resources Division

Figure 2-1. Boundary delineation of the Mobile-Tensaw River Bottomlands National Natural Landmark. Source: National Park Service (2011), as informed by Mount (1973).

8 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Bottomlands as a place of local, regional, national, and (Anas fulvigula maculosa). Mammals include beaver, international significance. As denoted in a 2011 NNL raccoon, river otter, and deer. The area is also one of Program Brief (NPS 2011), the Mobile-Tensaw River the few locations in Alabama where the Florida black Bottomlands NNL a) remains an expansive wetland bear (Ursus americanus floridanus) can be found. and bottomland forest ecosystem of mixed ownership and jurisdictions originating near the confluence of the This chapter briefly summarizes the history of the Alabama and Tombigbee Rivers and extending south- designation of the Mobile-Tensaw River Bottomlands ward for about 60 km (37 mi) to the head of Mobile NNL and subsequent further study by the NPS. It Bay; and b) continues to be one of the most important is part of a comprehensive multidisciplinary report wetlands in the nation. The area combines a variety of through the Gulf Coast Cooperative Ecosystem Stud- habitats, ranging from mesic flood plains and fresh- ies Unit that is intended to inform enduring NPS water swamps to open, brackish water marshes, with responsibilities to monitor the condition and encour- a network of interconnecting streams and lakes and age and support the conservation of this landmark interspersed low, forested islands. It is an important site. habitat for wildlife, and the vastness of the delta and its limited accessibility continues to support its basic Acknowledgments integrity. Significant reptiles include the American alli- The author gratefully acknowledges Heather gator (Alligator mississippiensis), the federally endan- Eggleston and Carolyn Davis of the National Park gered Alabama red-bellied turtle (Pseudemys alabam- Service, National Natural Landmarks Program, and ensis), and the delta map turtle (Graptemys nigrinoda David Morgan, PhD, National Park Service, Southeast delticola). Diversity of birdlife is exceptional, includ- Archeological Center, who helped locate key docu- ing the Swallow-tail Kite (Elanoides forficatus) and ments and provided manuscript review. the Mississippi Kite (Ictinia mississippiensis), as well as abundant waterfowl, such as the rare mottled duck

Introduction - Chapter 2 9

PHYSICAL LANDSCAPE

Mobile-Tensaw Delta, Alabama. Photograph courtesy of Hunter Nichols.

Physical Landscape - Chapter 3 11

3. Geology of the Lower Mobile River Basin Antonio B. Rodriguez, University of North Carolina at Chapel Hill

Shaped like a dowsing rod pointing south to the ocean, chert, separated by valleys of less resistant shale and the Tombigbee and Alabama river valleys merge into carbonate rocks (Raymond et al. 1988). The Piedmont the extensive Mobile River Valley, which forms a delta and Blue Ridge, in the northeastern part of the basin, at the head of Mobile Bay (a bayhead delta). The loca- is characterized by igneous and metamorphic rocks tion and morphology of the Valley is controlled, in with elevations reaching greater than 3000 ft (910 m) part, by tectonic events related to the creation of the above sea level. More than half of the Mobile River Gulf of Mexico. Over the past few hundred thou- Basin is in the Coastal Plain, which is primarily uncon- sand years, the valley experienced multiple episodes solidated or poorly consolidated sands, gravels, clays, of erosion and deposition related to climate cycles and limestones. and associated fluctuations in sea level that repeatedly exposed and inundated the continental shelf In the lower coastal plain, the floodplains of the of the northern Gulf of Mexico. Reconstructing the Tombigbee and Alabama rivers widen dramatically paleogeography of the Mobile Bayhead Delta over the last 9000 years shows that it is highly sensitive to small increases in the rate of sea level rise, which calls into question the sustainability of the bayhead delta’s current extent in the face of global warming.

Geologic Setting of the Lower Mobile River Basin The extensive Mobile River Basin (41,600 mi2 [107,700 km2]) is made up of the Tombigbee and Alabama rivers (Figure 3-1). The basin extends across the broad flat coastal plain in Mississippi and Alabama and into the rugged southern Appalachian Moun- tains of Alabama, Tennessee, and Georgia in the north, including the Appalachian Plateaus, Valley and Ridge, and Piedmont and Blue Ridge provinces (Figure 3-1). The Appalachian Plateaus are composed mainly of limestone, sandstone, and shale and make up the northwestern part of the basin. The Valley and Ridge is made up of a series of northeast Figure 3-1. Geologic map showing the different geologic provinces (arrows) and trending subparallel ridges of the scale of the Mobile River drainage basin, which comprises the Tombigbee erosion resistant sandstone and and Alabama river valleys (above). Location of the Mobile River Basin is shown in the lower image. (Modified after Moore 1971 and Planert et al. 1993).

Physical Landscape - Chapter 3 13 from around 2.5 to 7 km (1.6 to 4.3 mi) just upstream 3300 ft thick (1000 m) (Wilson 1975; Miller 1982). from where they merge forming the Mobile River The salt was subsequently buried beneath denser (Figure 3-2). Downstream from that junction the sediment and its buoyancy helped to promote move- Mobile River Valley is extremely straight, trends to ment, which resulted in the formation of a variety of the north, and is consistently around 12 km (7.5 mi) complex structures including salt domes positioned wide to Mobile Bay, a distance of 37 mi (60 km; Figure between 1000 and 6000 ft (300 and 1800 m) from the 3-2). The general morphology and location of the surface. The Mobile Graben, a low area between two Mobile Valley has been attributed, in part, to geologic normal faults, is another structural element related to events dating back to the initial formation of the salt movement (Figure 3-1). The sides of the Mobile Gulf of Mexico (Planert et al. 1993). The continen- Valley and Bay generally line up with the north trend- tal crust extended (stretched) in this area during the ing faults that mark the eastern and western boundar- breakup of the super continent Pangea, around 180 ies of the Mobile Graben and the eastern extent of the million years ago, forming a rift basin that was peri- salt basin (Figure 3-1; Moore 1971). Many geologists odically connected to the ocean during Jurassic time have hypothesized that movement along the faults (160 to 140 million years ago; Salvador 1987). The within this system resulted in the confluence of the arid climate and restricted basin resulted in persistent Alabama and Tombigbee rivers and formation of the evaporation of sea water and deposition of salt up to broad Mobile Valley and Bay (Murray 1960; Tanner

Figure 3-2. Digital elevation model of the Lower Mobile River Basin. Resolution for areas outside and inside the black box is 1 arc-second and 1/3 arc-second, respectively. Data are from the National Oceanic and Atmospheric Administration’s National Centers for Environmental Information, formerly the National Geophysical Data Center.

14 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 1966; Smith 1988; Planert et al. 1993). While the loca- Martinsen 1996). For the Mobile River system, the tion of the valley may be a product of the structural gradient of the continental shelf is about 4 times the geology of the area, the depth of the valley and the coastal plain gradient. As a river crosses that apex, a sediments that fill it are largely controlled by fluctua- knickpoint forms, which is defined by a sharp change tions in sea level. in river slope. The river gradient will equilibrate by the process of landward knickpoint migration, which Late Quaternary Sea Level erodes the apex (Schumm et al. 1984; Schumm and Fluctuations and River Erosion Brackenridge 1987). The knickpoint moves upstream High magnitude (330 ft [100 m]) sea level cycles of only several miles for small, high gradient fluvial 20,000- to 100,000-year duration have been the pattern systems, whereas large, low gradient fluvial systems, for at least the last million years. The main cause like the Mobile River, adjust their profiles over 100 mi for these sea level fluctuations is the expansion and (160 km) upstream (Blum and Törnqvist 2000). The retreat of glaciers. During the last glacial maximum great landward extent of the valley, in part, is due to 21,000 years ago, water was stored on the continents the apex of the coastal prism being exposed during in glaciers and massive ice sheets, which lowered sea multiple sea level cycles, and the Mobile River erod- level to 394 ft (120 m) below present and caused north- ing to adjust its profile. ern ice sheets to reach 1.8 mi (2.9 km) in thickness. Evidence for multiple cycles of valley incision are As the climate warmed, glaciers and ice sheets melted preserved at the valley edge as a series of downward and their water returned to the oceans. Currently we stepping terraces above the present floodplain (Figure are experiencing a sea level highstand and there is 3-2; Smith 1997), and in the valley-fill stratigraphy relatively little glacial ice covering the planet. During as multiple stacked erosional surfaces (Greene et al. the last comparable event, around 120,000 years ago, 2007). The topography of those terraces show relict global mean sea level reached +4 to +9 m (+13 to +30 channels, point bars, and scroll bars and the terraces ft) (Cuffey and Marshal 2000; Lambeck et al. 2012; are remnants of former higher elevation floodplains. Kopp et al. 2013; Simms et al. 2013). Evidence of this Each terrace represents an episode of floodplain previous highstand is preserved as a 120,000-year-old deposition followed by river erosion. The downstream beach ridge on the back side of Fort Morgan Peninsula gradient of terraces on the western side of the valley that averages 4 to 5 m (13 to 16 ft) in elevation (Figure is extremely low, less than 1.32 ft/mi (0.25 m/km), 3-2) (Otvos and Howat 1992; Rodriguez and Meyer similar to the modern downstream gradient of the 2006). Given that the seaward edge of the continen- lower Mobile River floodplain of less than 0.53 ft/mi tal shelf offshore Alabama is 141 ft (43 m) below sea level, Mobile Bay and the continental shelf were exposed during most of the last sea level cycle (Figure 3-3) and for many of the sea level cycles during the past 1 million years. When the continental shelf was exposed, the Mobile River extended across the 46-mi (74-km) wide shelf, which influenced the morphology of the lower Mobile River Valley.

At the coast, rivers erode valleys when sea level fall exposes a surface with a steeper gradient than the river profile, such as the apex of the coastal prism, which is the transition between the flat coastal plain and steeper continental shelf (Helland-Hansen and Figure 3-3. Sea level curve for the northern Gulf of Mexico after Anderson et al. (2004).

Physical Landscape - Chapter 3 15 (0.10 m/km), suggesting that the current morphology ft) below sea level at the modern bayhead delta loca- of the lower Mobile River has been repeated multiple tion. The resulting erosional surface is clearly defined times in the past. in cores as a sharp contact between recent mud and older highly compacted and oxidized sand and clay. High resolution seismic data and cores from Mobile Oxidation results from sediment being exposed to air Bay and the Mobile Bayhead Delta show multi- for a long period. The increase in sediment compac- ple stacked erosional surfaces that formed by river tion across the erosional surface makes it easy to processes during periods when sea level was lower image and map regionally with seismic data (Rodri- and the area was exposed (Figure 3-4; Greene et guez et al. 2008). That mapping shows the area is al. 2007). The most recent episode of river erosion defined by multiple channels that extend onto the began as the climate was progressing towards the continental shelf. Similar to the valley terraces, each last glacial maximum and the inner continental shelf erosional surface represents an episode of river inci- was exposed, which resulted in incision to 20 m (66 sion followed by deposition. The more recent valleys generally reoccupied earlier valley positions, which further supports the interpretation that the modern morphology of the lower Mobile River is a reflection of previous cycles.

Transgression and Valley Fill The last deglaciation, from around 21,000 to 6000 years ago, caused sea level to rise approximately 330 ft (100 m) and the shoreline to move landward. Because the bases of valleys have a gradual slope, rising sea level penetrates many miles into the coastal plain during inundation. As a result, drowned river mouth estuaries, such as Mobile Bay, form. Mobile Bay is restricted from marine influence at its mouth by Dauphin Island in the west, and Morgan Peninsula in the east. The middle part of Mobile Bay is a large (400 mi2 [1036 km2]) and relatively shallow (10 to 13 ft [3 to 4 m]) basin where thick (up to 65 ft [20 m]) gray mud is deposited. The Mobile Figure 3-4. Cross sections through the lower Mobile River Basin and Mobile Bay show- River forms a bayhead delta ing two stacked erosional surfaces that formed during periods of river incision. The where it meets the bay and is erosional surface on top of the black unit formed during the last glacial maximum. Valley fill sediments shown in gray accreted during the last 9000 years. Figure after the main source of freshwa- Greene et al. (2007). ter into the estuary. During

16 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area sea level rise, the estuary and bayhead delta deposi- silt with abundant organic material such as wood and tional environments move landward up the valley and leaf litter. Overlaying delta plain deposits, the core a record of this is left behind in the stratigraphy of sampled 23 ft (7 m) of delta-front sand with abundant the estuary as thin bayhead delta deposits overlain by shell material. Delta plain below delta front environ- thick middle bay deposits. ments shows that the bayhead delta’s movement landward in response to sea level rise outpaced sedimenta- Before Mobile Bay existed, that area was occupied by tion. Over millennial time scales, the old delta plain, the bayhead delta and/or the river floodplain depo- seen at the base of the core, accumulated at a deceler- sitional environment. Radiocarbon dates from cores ating rate from 0.18 to 0.05 inch/yr (4.6 to 1.3 mm/yr) collected in upper and lower Mobile Bay show that and the delta front accumulated at 0.05 inch/yr (1.3 the estuary formed sometime between 8900 and mm/yr), about the same as sediment accumulation 8100 years ago (Rodriguez et al. 2008) and the entire rates in the middle of Mobile Bay (0.07 inch/yr; 1.8 modern extent of the estuary inundated rapidly mm/yr) (Rodriguez et al. 2008). These sedimentation during a 100-year period (Rodriguez et al. 2010). The rates are less than the present rate of relative sea level low gradient of the Mobile River Basin and a brief rise measured at Dauphin Island, 0.12 inch/yr (3 mm/ increase in the rate of sea level rise 8200 years ago are yr); however, sediment accumulation rates at decadal likely causes for the high rate of flooding and rapid time scales are over twice the millennial rate (Smith formation of Mobile Bay. This increase in rate of sea et al. 2013). This higher rate of recent sedimentation level rise occurred in response to the abrupt drainage is due, in part, to anthropogenic modifications to the of glacial lakes Agassiz and Ojibway (Alley and Ágúst- delta system, importantly, construction of the cause- sdóttir 2005; Kendall et al. 2008), which also caused way in the 1920s that restricts sediment and water marine inundation in Sabine Lake, Galveston Bay, exchange with Mobile Bay (Smith et al. 2013). Corpus Christi Bay, and Baffin Bay (Anderson and Rodriguez 2008; Rodriguez et al. 2010). The rapid transition from bayhead delta to estuary that occurred around 8200 years ago in the area of Mobile Bayhead Delta modern Mobile Bay demonstrates that bayhead deltas The Mobile Bayhead Delta extends 28 mi (45 km) from are highly sensitive to increases in the rate of sea level the first distributary channel in the lower Mobile River rise. During sea level rise, bayhead deltas, in general, valley to the upper bay. The Mobile River bifurcates are predisposed to episodes of rapid landward retreat into the Mobile and Tensaw distributary channels less as a result of the dendritic morphology of their river than 5 mi (8 km) downstream from where the Tombig- valleys (Simms and Rodriguez 2014). Tributary junc- bee and Alabama rivers merge (Figure 3-2). Those two tions, such as the one that exists below the modern main distributary channels of the delta discharge into Mobile Bayhead Delta location (Figure 3-4; Greene the western (Mobile) and eastern (Tensaw) sides of et al. 2007), act as pinning points for the transgress- Mobile Bay. The delta plain contains numerous highly ing shoreline. This is mainly due to sharp increases in sinuous distributary channels, levees, interdistribu- valley gradient and decreases in valley width across tary bays, and delta lobes. tributary junctions. With accelerating sea level rise, the Mobile Bayhead Delta will likely stabilize at The bayhead delta has existed at its modern location the next tributary junction located 35 mi (56 km) since around 9800 years ago. This conclusion is based upstream from the modern bayhead delta front where on a 62 ft- (19 m-) long core collected from the delta the Alabama and Tombigbee rivers merge. The timing front (Rodriguez et al. 2008). At the base of the delta, for this next (final?) bayhead delta backstepping event the core sampled 29.5 ft (9 m) of delta plain clay and is uncertain.

Physical Landscape - Chapter 3 17

4. Hydrology and Water Quality in the Mobile- Tensaw Delta G. Milton Ward, Amelia K. Ward, and Alex Maestre, University of Alabama

The Mobile-Tensaw River Delta (hereafter, “the the lower half of the Mobile Basin, water flows through Delta”) is the terminus of a river system with the 4th geologically younger, unconsolidated sediments. This largest discharge to a coastal area in North America geologic diversity produces, in many cases, differing (Wilson and Iseri 1969). The Mobile River, which amounts of runoff and distinct chemical composi- flows into the Delta from the north, is relatively short, tions of water, which contribute to the high biological only 50 mi (80 km) long. However, it is formed from diversity in the basin (Ward et al. 1991, 1992). the merging of two large rivers, the Alabama and Tombigbee, which drain (with their tributaries) the Abundant rainfall and runoff in the basin supply the eastern and western subbasins of the total Mobile Delta with ample water each year. Indeed, it is this River Basin. At 44,000 mi2 (114,000 km2) this basin abundance within the basin that ultimately creates the is the 6th largest in the US and the largest east of the vast wetland landscape we now observe as the Delta. Mississippi River that drains into the Gulf of Mexico Long-term average rainfall throughout the basin is (Atkins et al. 2004; Ward et al. 2005). It covers most of Alabama, parts of Missis- sippi and Georgia, and a small portion of Tennes- see. Therefore, the Delta receives water and materials that emerge from this expansive river system and serves as a critical buffer between it and coastal areas to the south, including Mobile Bay (Figure 4-1). Geomorphology, hydrology, and water quality of the Delta are highly influenced by features of the upgradient landscape and river systems of the Mobile River Basin.

Water that reaches the Delta travels through a landscape that exhibits great geologic and physiographic diversity. The northeastern portion of the basin presents higher elevations and very Figure 4-1. Mobile River Basin and river network. (Courtesy of Grover Ward and old rock (Cambrian and Amelia Ward, University of Alabama; image credit: Sarah Mattics, University of South Precambrian) formations. In Alabama.)

Physical Landscape - Chapter 4 19 relatively evenly distributed across seasons, although the confluence of the Alabama and Tombigbee rivers, low rainfall summer seasons are well known. Typi- the Mobile River splits to form the Tensaw River, cal summer precipitation is the result of convectional which runs along the eastern side of the Delta, and rainfall, but heavy rainfall from tropical storms and the Mobile River, which runs along the west side. hurricanes often produce large amounts of rain in The Tensaw River branches several times, forming normally drier summer months. Over the long term, Middle, Raft, Blakeley, and Apalachee rivers. Eventu- these heavy rainfall years even out seasonal rainfall ally 5 channels reach upper Mobile Bay. Spanish River, variability. Rainfall and runoff are not evenly distrib- which drains Grand Bay, one of the larger floodbasins, uted across the basin. The Delta and coastal areas receives water from Raft River before reaching Mobile typically receive average annual precipitation of 60 Bay (Figure 4-2). to 64 inches (150 to 160 cm), whereas further north in the basin annual rainfall amounts range from 52 to The Delta landscape is a complex of terrestrial land- 58 inches (130 to 150 cm) (Johnson et al. 2002). Not forms and aquatic habitats, created by thousands of surprisingly, annual runoff values are higher near years of geological, fluvial, and geomorphic interac- the coast (26 inches [66 cm]) relative to areas further tions. Smith (1988) provided detailed analyses of the north within Alabama (Johnson et al. 2002). many geomorphology components of the Delta landscape; the more important ones are noted below. In Geomorphology addition to active distributary channels, other major landscape components include islands, floodbasins, Flow from the major tributaries is responsible for levees, bays, abandoned channels, lakes, sloughs, and sustaining the highly diverse landscape within the bayous. Levees, observed frequently along distribu- Delta. Published values for the size and areal cover- tary channels, develop as a result of the capture of age vary somewhat, depending on boundary defini- sand and silt suspended in flood waters that over- tions employed, but the Delta is about 45 mi (72 km) top river banks and build in height over time from long, from 6 to 16 mi (10 to 26 km) wide and 300 to repeated flooding. 400 mi2 (780 to 1040 km2) in area. The US Department of Commerce (1979) estimated the area as 185,000 Floodbasins, a major landscape component of the acres (750 km2), identifying 20,000 acres (80 km2) of Delta, are large areas of floodplain forest surrounded open water; 10,000 acres (40 km2) of marsh; more by distributary channels and their associated levees. than 70,000 acres (280 km2) of swamp; and more than These basins receive flood water when levees are over- 85,000 acres (340 km2) of bottomland forest. Fortu- topped, or receive inflow through breaches (crevasse nately, federal, state, and private conservation efforts channels) when water levels are below the height of have resulted in a large amount of land now protected the levee. Much of the inflowing water will later drain and managed (Smith 2014). back through these breaches into distributary channels, or over low points in levees. These landscape The Delta is considered geologically to be a drowned units typically contain a network of stream channels alluvial plain and valley, one of several along the that permit drainage out of the floodbasin after flood northern Gulf Coast (Smith 1988; Kindinger 2015). events. Examples include Gravine Island and the However, only the Mobile and Pascagoula rivers Mifflin Lake basin. exhibit well-developed alluvial plains. Because the lower Alabama and Tombigbee rivers flow through a Within the Delta, relict stream and abandoned river relatively flat landscape, these channels exhibit broad channels are common landscape features. Former meanders, and water in river channels in such terrain channels once carried river water, but flood waters cut frequently overtops river banks. Water elevations new routes and left them abandoned. Sedimentation from the confluence of the Alabama and Tombigbee at the terminal ends has isolated them from riverine rivers to Mobile Bay decline by roughly 0.16 ft/mi (3 flow, creating floodplain lakes that retain stream chan- cm/km; Smith 1988). Thus, over time, flood flows nel shapes. Notable examples include Tensaw Lake, moving south to Mobile Bay have carved a complex Mifflin Lake, Stiggins Lake, Big Chippewa Lake, and network of interconnected channels, described as a Little Chippewa Lake. distributary network. Many shallow basins, termed bays, exist at the south- The Mobile-Tensaw River distributary network ern end of the Delta. Strictly speaking, bays are conducts water through 7 major channels. South of restricted to embayments at the terminus of the Delta

20 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figure 4-2. Mobile-Tensaw Delta landscape. (Courtesy of Grover Ward and Amelia Ward, University of Alabama; image credit: Sarah Mattics, University of South Alabama.)

impacted by tidal fluxes. For example, Grand Bay, Delta Movement during the Holocene Pole Cat Bay, Chacaloochee Bay, and Bay Minette are The Delta has been in its present location for roughly actually relict floodbasins that have been inundated the last 4000 years, but over geological time has by rising sea level. Stream channels that are a) tribu- migrated north and south from its present loca- taries to bays or inundated floodbasins and b) subject tion coincident with the rise and fall of sea level to tidal influences are termed bayous (e.g., Bayou Sara (Hummel and Parker 1995). Sea levels 30,000 to and Catfish Bayou). 35,000 years before present were thought to be similar

Physical Landscape - Chapter 4 21 to present day levels. However, from 16,000 to 18,000 ft3/s (1760 m3/s). That discharge is made up of the years before present, global sea level was 427 ft (130 Mobile River (95%) and several small independent m) lower than at present (Milliman and Emery 1968). tributaries (Loyacano and Busch 1979). The total flow During the last glaciation, the Gulf of Mexico shore- of the Mobile River is derived from the Alabama River line was perhaps 60 mi (100 km) south of its present (52%) and the Tombigbee River (48%) (Johnson et al. position (Smith 1988). After glaciers began melting 2002). and prior to 7000 years before present, the head of the Delta system lay in what is now southern Mobile Bay, Inflows vary substantially, both seasonally and annu- extending out onto the continental shelf. As is the case ally. Analyses of United States Geological Survey currently, the Delta contained a distributary network (USGS) discharge data for 1976 to 2014 from gauges of channels, the larger of which flowed through an at Coffeeville Lock and Dam (Tombigbee River) and area now occupied by the Fort Morgan Peninsula Claiborne Lock and Dam (Alabama River) were and another through present-day Main Pass. Tribu- combined to estimate mean flow into the Delta (Figure taries that currently flow directly into Mobile Bay 4-3). Highest average monthly flows occur from Janu- 3 would have discharged instead into the main stem ary to April each year, typically exceeding 95,000 ft /s 3 of the Mobile River. At this time, the Mobile River (2700 m /s). Discharge falls rapidly over the late spring 3 was perhaps 100 to 125 ft (31 to 38 m) lower than at and early summer, reaching flows less than 25,000 ft /s 3 present (Smith 1988), having downcut through sedi- (710 m /s) in August through October. Detailed and ments of Miocene, Pliocene, and Pleistocene age. accurate knowledge of hydrological characteristics is River-derived sediments and ocean sediment depos- one of the most important physical features of aquatic its sustained terrestrial habitats as the Delta migrated. ecosystems. This is important for wetlands because As glaciers melted, sea levels rose more rapidly, and by the biological structure of the system is determined 7000 years before present the water in the main chan- by the quantity, quality, and timing of water move- nel of the Mobile River flowed through what is now ment through the system. Unfortunately, knowledge Mobile Bay, likely flanked by marshlands and perhaps of wetland hydrology lags well behind that of other a wide bottomland floodplain forest.

By 6000 years before present, rising sea levels (to a point 15 ft [5 m] below current level) caused a northward migration of the Delta as more than 75% of Mobile Bay was inundated with seawater and filled with sediment (Smith 1988). Approximately 4000 years ago sea levels stabilized, Mobile Bay assumed its current location, and the southern edge of the Delta was established. The landforms, bays, bayous, and distributary drainage pattern we now observe in the Delta are, thus, the result of a river valley slowly drowned as a result of sea level rise. As the southern edge of the Delta disappeared due to inundation by the more saline waters of Mobile Bay, typical deltaic landforms developed at its northern edge.

Hydrology Figure 4-3. Monthly mean, minimum, and maximum discharge (in ft3/s or The mean annual discharge of water cfs) into the Mobile River based on summation of discharge from lower into the Delta is approximately 62,100 Alabama and lower Tombigbee rivers, 1976-2014. (Courtesy of Grover Ward and Amelia Ward, University of Alabama; image credit: Sarah Mattics, University of South Alabama.)

22 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area aquatic systems, and the larger and more complex the years 2003 to 2006 to construct a preliminary water wetland, the more difficult it is to determine sources, budget for the Delta. Not surprisingly, not all of the routing, and losses of water. The complex topography water flowing from the Alabama and Tombigbee rivers and diversity of land and water features in the Delta could be accounted for by summing data for flows in make an accurate estimate of inflow and water rout- the Mobile and Tensaw rivers. Indeed, approximately ing challenging because so many measurements are 12.5% of the annual input of water from the Alabama required to characterize the many braided channels, and Tombigbee rivers was not measured as flow bays, and off-channel lakes where water is naturally through the channels of the Mobile or Tensaw rivers. stored. Also, due to low elevation gradients over- This analysis suggested that when the combined flow land flow typically occurs during floods, but direct of the Alabama and Tombigbee rivers was less than measurement of this movement is difficult to obtain. 130,000 ft3/s (3700 m3/s), flow values in the Mobile and Tensaw rivers were similar to the summed flow of USGS gauging stations on the lower Alabama and these two rivers. However, flows greater than 130,000 Tombigbee rivers have been in operation for several ft3/s (3700 m3/s) did not result in similar flows. O’Neil decades and provide flow and other data necessary hypothesized that the “missing” discharge likely origi- to compute riverine flow, nutrient, and other mate- nated in the lower Alabama River, and during flood rial input to the Mobile River. Two other gauging events likely flowed overland to the south where it stations located within the Delta, with shorter periods entered the Tensaw River, completely missing the two of record but still useful for these calculations, are on river gauges. Given the geomorphic structure of the the Mobile River at Bucks, Alabama, 31 mi (50 km) Delta, this idea is plausible. north of Mobile Bay, and on the Tensaw River near Mt. Vernon, Alabama, east of the Mobile River. Overland flow in the Delta appears to be an annual occurrence. Analysis (for this report) of USGS flow An accurate hydrological budget for the Delta first data for the lower Alabama and Tombigbee rivers over requires an accurate measurement of combined flows the period 1976 to 2014 revealed that flows greater from the Alabama and Tombigbee rivers as an estimate than 130,000 ft3/s (3700 m3/s) occur every year. On of the total water input to the Delta from the Mobile average, flows of this magnitude or greater occur 48 River Basin. Gauges on the lower Alabama and days per year, ranging from a low of only 6 days in the Tombigbee rivers provide the long-term flow record drought years of 1988 and 2007 to a high of 134 days needed for these estimates. A second requirement for in 1983. a robust hydrological budget is quantification of flow through the Mobile River, on the western edge of the Hydrological Alterations Delta, as well as the Tensaw River that flows along the eastern boundary. All other distributary channels Humans frequently modify hydrologic patterns in within the Delta receive the majority of their flow from wetlands and rivers. Such modifications often impact these two major rivers. However, flow measurements water movement, and especially hydrologic connec- and calculations for these rivers are confounded for tivity with adjacent ecosystems. Over the decades, several reasons. Accurate estimates of flow require a several bridges and roadways across the Delta and substantial period of record, but flow records from Mobile Bay have been constructed. Bridge supports gauges on the Mobile River near Bucks and the Tensaw tend to obstruct downstream flow as well as seawa- River near Mt. Vernon have only been in place for 12 ter exchange between the Delta and Mobile Bay. years. The most difficult estimates to acquire are for However, the most significant obstruction to connec- flow during floods, when water overtops river banks. tivity between the Delta and Mobile Bay is the Mobile In addition, measuring the quantity and distribution Bay Causeway (US Highway 90/98). Constructed of overland flow (flow outside river channels) across in 1926 as the first direct road connection between the Delta would be technically difficult. Lastly, flow Mobile and Baldwin counties, it consists of a series of calculations must account for the fact that tidal fluxes raised earthen embankments and concrete bridges, in Mobile Bay impact the measurement of river flow and it served as the primary transportation artery even at the northern end of the Delta. over Mobile Bay until the Interstate 10 Jubilee Park- way (locally known as the “Bayway”) was completed O’Neil (2007) examined data from the Tensaw River in 1978. The causeway’s embankments have impeded (Mt. Vernon) and Mobile River (Bucks) gauges for the exchange of water between Mobile Bay and the

Physical Landscape - Chapter 4 23 lower Delta, largely by reducing the cross-sectional functions as a nutrient retention and transformation area through which higher salinity estuarine water zone between the Mobile River and Mobile Bay, thus can flow from Mobile Bay into the lower reaches lessening nitrate-nitrogen export to Mobile Bay and of the Delta. The loss of hydrological connectiv- fragile coastal areas. ity (between the Delta and Mobile Bay) reduced the salinity of important nursery habitats (e.g., Chocolatta Although the Mobile River Basin is noted for its nitro- Bay and Tensaw River). As a result, species composi- gen retention capabilities, human activities threaten to tion of fish, crustaceans, and shellfish communities saturate them, which would result in greater export of has been altered within these habitats. Lowered salin- nitrogen, and possibly other nutrients, to the Delta and ity resulted in loss of nursery habitat and population coastal areas. The Delta is the last nutrient retention densities for commercially important species, such defense before Mobile Bay; thus, if more nutrients are as Blue Crab, white shrimp, and southern flounder exported into the Delta from upgradient rivers, then (Rozas et al. 2013). more pressure is put on retention mechanisms within the Delta.

Water Quality Over 95% of both nitrogen and phosphorus inputs to The Mobile River delivers 95% of the freshwater input the Mobile River Basin derive from nonpoint sources, to the Delta and Mobile Bay from the merged flows with major contributions from animal wastes, row of the Tombigbee and Alabama rivers (Schroeder and crops, and urban areas (Atkins et al. 2004). A compar- Wiseman 1999). It is also a primary nutrient source ison of nutrient flux from the lower Alabama and to both (Pennock et al. 1999). Reported high levels of Tombigbee rivers shows that export of nitrate-N, nitrate input from Midwestern agricultural areas into ammonium-N, and orthophosphate-P (soluble reac- the Mississippi River that cause “dead zones” in the tive phosphorus) into the Delta is generally higher northern Gulf of Mexico (e.g., Rabelais et al. 2002) from the Tombigbee River than from the Alabama have stimulated interest in nitrogen dynamics in other River, western and eastern subbasins of the Mobile watersheds and riverine-dominated estuaries of the River Basin, respectively (Table 4-1). Also, in both northern Gulf Coast. Compared to other watersheds rivers, nitrate-N flux is higher than ammonium-N that have been studied across the US, the Mobile flux and, thus, represents a higher proportion of over- River Basin appears to be nitrogen retentive. Total all dissolved inorganic nitrogen transported into the nitrogen exported from the Basin (i.e., discharged Delta, based on minimum to maximum ranges of each into the Mobile River), based on nitrogen flux from over multiple years. the lower Tombigbee and Alabama rivers, is about 5 to 7% of total nitrogen input to the Basin, whereas A few recent studies have investigated nutrient other watersheds typically export 25 to 30% of basin concentrations within the Delta (Valentine and inputs to coastal areas (Carey et al. 2003). Nitrogen Sklenar 2006; O’Neil 2007). O’Neil (2007) reported retention mechanisms within the Mobile River Basin turbidity, suspended solids, and total nitrogen are have not been well studied. However, nitrogen is likely reduced in Tensaw Lake and the Tensaw River chan- retained in forest soils, as well as in floodplains and nel compared to Mobile, Alabama, Tombigbee, and other features of large rivers (Tatariw et al. 2013). The Middle rivers. These data suggest that these compo- Delta, which receives nutrients from this vast upgra- nents are filtered out of overland flow in the eastern dient river system via the Mobile River, also likely Delta. Nevertheless, concentrations for chlorophyll a,

Alabama River Tombigbee River Nutrient Min Max Years Min Max Years Nitrate-N 1540 (3395) 7160 (15,785) 2001-2011 2800 (6173) 11,300 (24,912) 2001-2011 Ammonium-N 560 (1235) 1980 (4365) 1986-1993 169 (373) 1574 (3470) 2001-2014 Orthophosphate-P 594 (1310) 3861 (8512) 1991-2013 1260 (2778) 7680 (16,932) 2001-2014

24 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area total nitrogen, and suspended solids collected from Trace elements most frequently detected in fish tissue the Mobile River (western Delta), Middle River, collected from sites throughout the Mobile River Tensaw Lake, and Tensaw River (all sites north of Basin suggest that aluminum, boron, copper, iron, Interstate 65 in the northern part of the Delta) (O’Neil manganese, selenium, strontium, and zinc are most 2007) indicate ongoing mesotrophic to mesoeutro- likely to bioaccumulate in riverine fish (Zappia 1998). phic conditions (Wetzel 2001). Values for total phosphorus from those same sites suggest mesoeutrophic Within the Delta, most detection of trace metals, to hypereutrophic conditions—in other words, highly including toxic ones, occurred in the Mobile River enriched and productive environments. Concentra- on the western side of the Delta (O’Neil 2007). Tita- tions of soluble nutrients important to plant growth, nium, arsenic, nickel, antimony, and lead were most such as orthophosphate-P, ammonium-N, and nitrate- frequently encountered. However, all pesticides and N, were low (orthophosphate-P was undetectable) herbicides analyzed in the overlying water were below to moderate. The combination of somewhat modest detection limits, and the only pesticide or herbicide concentrations of dissolved nutrients, but relatively detected in sediments was 2,4-D. Off-channel lakes high concentrations of nutrients in particulate form and embayments are sinks for metals because of (chlorophyll a, total nitrogen, and total phosphorus), higher organic matter and low flushing rates. Highest suggests rapid uptake of soluble inorganic nutrients levels of the toxic trace metals cadmium and mercury by phytoplankton, other microbes, and plants that occurred in sediments in Cedar Creek and Tensaw occupy the diverse habitats within the Delta. This Lake, which are not in the main river channels of the mechanism likely retains nutrients and also lessens Delta. Black water streams also accumulate metals nutrient transport downgradient to southern parts of because of higher acidity, increased organic matter, the Delta and Mobile Bay. and low flushing rate. Comparisons of nutrients in dissolved and particu- Summary late forms just north and south of the causeway, in the The Delta is one of the most important and valuable southern reaches of the Delta (Valentine and Sklenar natural resources in the US. It has existed for tens 2006), revealed slightly higher values than in northern of thousands of years, and currently receives water, parts of the Delta (O’Neil 2007). Since the two stud- sediments, and solutes from a substantial area of ies were completed several years apart, these differ- Alabama and adjoining states via the river system of ences may reflect inter-annual variability of nutrients the Mobile River Basin. Sea level changes and water in general in the Delta. Alternatively, areas of the Delta flow from the basin have over time carved an exten- near the causeway, the city of Mobile, and other urban sive and complex riverscape of channels, lakes, bays, developments may be affected by human activities bayous, and bottomland forests, which both harbor that increase nutrients proximate to those activities. extensive wildlife resources and act to sequester The lower part of the Delta is also influenced by inter- sediments, nutrients, metals, and pesticides draining actions with water in Mobile Bay, including saltwater from the basin upstream. It is important to recognize intrusions that may affect nutrient dynamics. that human activities in the Mobile River Basin have In addition to nutrients described above, the Delta profound impacts on the Delta downstream. We do receives inputs of other types of chemicals. Pesticides not currently have a thorough understanding of the are widespread in streams and rivers in the Mobile rates of water and material flow in the Delta nor of the River Basin, but in relatively low concentrations, with myriad roles of biota in mitigating nutrient and other heavily used atrazine, simazine, metolachor, tebuthi- chemical inputs that otherwise would be transported uron, and 2,4-D (the herbicide Dichlorophenoxyace- into Mobile Bay. Without this knowledge, fully effec- tic acid) most frequently detected (Atkins et al. 2004). tive management of the Delta will remain elusive.

Physical Landscape - Chapter 4 25

BIOTIC LANDSCAPE

Green tree frog, Mobile-Tensaw Delta, Alabama. Photograph courtesy of Hunter Nichols.

Biotic Landscape - Chapter 5 27

5. Paleobotany and Paleoclimate Debra Z. Stults and Brian J. Axsmith, University of South Alabama

Paleobotanical assemblages can be valuable resources burial (Wallinga 2008), provides an estimate of 83,000 for understanding how modern terrestrial plant years ago for this deposit, which correlates with the communities developed, but they are only now being last warm episode (interglacial) before the final glacial analyzed with modern techniques in the greater advance of the Pleistocene. Mobile-Tensaw River area. This oversight is due to the historical perception that this region is nearly devoid Common plant fossils from this deposit include of fossil plant localities, despite being underlain by members of the birch family (Betulaceae), includ- massive layers of sediments deposited over the last ing Betula nigra (river birch) and Carpinus caro- 40 million years (Graham 1999a). Ongoing research liniana (ironwood), and several oaks, including shows that this view is mistaken and that plant fossils Quercus lyrata (overcup oak), Q. michauxii (swamp can be obtained through persistent search. Although chestnut oak), and Q. virginiana (Virginia live oak). much of this work is in early stages, some important Other fossils include Carya aquatica (water hickory), findings are summarized here. This treatment focuses Liquidambar styraciflua (sweetgum), Nyssa aquatica on the fossil leaf and seed record, which is congruent (water tupelo), Pinus elliotii (slash pine), and Vitus with the stratigraphically limited pollen data. rotundifolia (muscadine grape) (Figure 5-1; Stults and Axsmith 2009). All of these plants are still common in The plant fossil record in the greater Mobile-Tensaw the modern greater Mobile-Tensaw River area, which River area begins with Eocene Epoch deposits indicates that this region did not experience the (approximately 40 million years ago) along the bluffs extensive community reorganizations that occurred in Monroe County, and extends into the last intergla- in more northern areas during the Pleistocene glacia- cial (approximately 85 thousand years ago) in Mobile tions (Delcourt and Delcourt 1993). County. This record is spotty, with the more recent floras the best studied. Therefore, this account starts with the youngest sites and works backwards.

Mobile River Terrace (Late Pleistocene—85,000 to 82,000 Thousand Years Ago) Pleistocene (ice age) terrace deposits resulting from sea level fluctuations are common around the greater Mobile-Tensaw River area, but their ages are disputed (Heinrich 2009). However, plant fossils from a terrace deposit along the Mobile River near Bucks, north of Mobile, have been reliably dated using optically stimulated luminescence (Stults and Axsmith 2009). This Figure 5-1. Representative plant fossils from a Pleistocene terrace deposit technique, which determines age by in Mobile County: A) seed cone of Pinus elliotii (slash pine); B) fruit of Liq- measuring radiation accumulated in uidambar styraciflua (sweetgum); C) acorn of Quercus lyrata (overcup oak); mineral crystals since their time of D) leaf of Betula nigra (river birch); E) leaf of Vitus rotundifolia (muscadine grape).

Biotic Landscape - Chapter 5 29 The Citronelle Formation Flora (Late (Figure 5-2) (Stults et al. 2002; Stults 2003, 2011; Stults Pliocene—3.4 to 2.7 Million Years Ago) and Axsmith 2009, 2011a). Some of the most abun- Quercus falcata The Citronelle Formation underlies much of the dant taxa include several oaks: (south- Q. virginiana Q. northern Gulf of Mexico Coastal Plain, extending ern red oak), (Virginia live oak), and laevis from east Texas to the Florida panhandle (Matson (turkey oak). Several species of hickory are also Carya aquatica 1916; Otvos 1998). Edward Berry (1916) long ago abundant, with (water hickory) being Platanus occidentalis described 18 plant fossil taxa from sites near Citro- most common. (American syca- Planera aquatica Betula nigra nelle in Mobile County and along Perdido Bay in more), (water elm), and Baldwin County. He attributed most of the fossils to (river birch) are also common at some localities. The Taxodium distichum species still living in the region, or considered them conifer (bald cypress), which is their immediate ancestors. One interesting excep- still an important tree of wetland habitats, occurs at tion is the strange horned fruit of the floating aquatic all localities (Stults et al. 2011). plant Trapa (water caltrop), which is extinct in North The Citronelle Formation flora also includes region- America but common in Africa and Eurasia. Little ally extinct forms, in addition to Trapa. Some of these additional work was done until we reinitiated study reveal the remarkable biogeographical fact that plant of the Citronelle flora. In addition to refining original communities of eastern North America are closely identifications, our investigations have revealed more related to those of eastern Asia, despite the thou- than 50 plant taxa. These findings are filling a major sands of kilometers separating them today (Manos gap in the fossil record, as floras of this age are rare and Meireles 2015). This knowledge indicates that throughout the continent. the forest types of these areas once extended more Berry’s (1916) conclusion that the Citronelle flora widely across the Northern Hemisphere. During the is dominated by plant groups still important in the Oligocene Epoch (approximately 35 million years region today is supported by our reinvestigations ago), climates became cooler and drier following the warmer temperatures of the Eocene. This trend continued with minor inter- ruptions through the Miocene and Pliocene, resulting in loss of deciduous forests in many areas. Such relictual forests now occur only in areas wet and warm enough to sustain them.

The fossil record indicates that floral similarity between eastern North Amer- ica and easterb Asia was even greater in the past, as many plants now considered Asian endemics (e.g., Ailanthus [tree of heaven] and Ginkgo) were once present in North America but became extinct here (Manchester 1999). A striking example from the Citronelle Formation includes discovery of fossil leaves and fruits of Pterocarya (wingnut), a member of the walnut family (Juglandaceae) represented today by 4 species in Asia. Nuts of Pterocarya bear Figure 5-2. Representative Citronelle Formation plant fossils of taxa still 2 wings for wind dispersal. Pterocarya living in the region today: A) branch of Taxodium distichum (bald cypress); fossils are common in Oligocene and B) branch of T. distichum with numerous pollen cones; C) leaflet of Carya aquatica (water hickory); D) fruit of Cyrilla racemiflora (titi); E) immature Miocene localities in North America, acorn of Quercus virginiana (Virginia live oak); F) leaf of Q. virginiana; Europe, and Asia. Citronelle Forma- G) leaf of Quercus falcata (southern red oak); H) leaf of Platanus occiden- tion Pterocarya fossils represent the last talis (American sycamore). occurrence of this genus in all of North

30 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area America (Stults 2011). Another remarkable find is a make to our understanding of the floristic history of fruit of Begonia (Stults and Axsmith 2011b). Although a region. Examples of regionally extinct taxa from the commonly known as a tropical houseplant, Begonia is Citronelle Formation can be seen in Figure 5-3. one of the most species-rich flowering plant genera. Yet the Citronelle Formation Begonia is the only fossil The abundance and quality of the Citronelle Forma- record for this genus. tion plant fossils makes them suitable for a variety of quantitative techniques for estimating the climate One of the most surprising findings from our Citro- under which they grew (Stults 2011; Stults and nelle Formation research involves the pines (Stults et Axsmith 2015). Such an effort is especially interesting al. 2011). The genus Pinus is composed of two subgen- for the late Pliocene, a time of recent global warmth era, Haploxylon (soft or white pines) and Diploxylon that interests researchers using fossil assemblages as (hard or yellow pines), distinguishable by leaf anatomy models for possible future climate change (Lunt et al. and pollen. Pines dominate many of the forests of the 2012). Quantitative paleoclimate methods using plant greater Mobile-Tensaw River area today, and they are fossils fall into two broad categories: a) those using all Diploxylon species. Pinus palustris (long leaf pine), specific structural features of leaves (physiognomy) P. taeda (loblolly pine), and P. elliotii (slash pine) have as paleoclimate indicators (Yang et al. 2011); and b) particular ecological and economic significance. The estimates based on climate tolerances of the nearest only Haploxylon pine in eastern North America today living relatives of the fossils (Utescher et al. 2014). is Pinus strobus (eastern white pine), which is common Physiognomic methods are based on the proposition in the northeast, but occurs in the southeast only in that leaf features (e.g., size, presence of teeth along the the Appalachian Mountains. Today, Haploxylon pines margins) are adaptations to specific climatic condi- are completely absent from Alabama. Based on the tions. Nearest-living-relatives approaches assume that similarity of the Citronelle Formation broadleaf flora the climatic requirements of fossil plants were similar to that of today, we anticipated a similar pattern with to those of their nearest living relatives. Two climate the pines, especially considering that their abundant, reconstruction analyses based on physiognomy robust needles and cones have excellent fossilization and two based on nearest-living-relatives analyses potential. However, Berry (1916) reported only one were performed on the Citronelle Formation fossils. possible needle fragment and a seed from his original Surprisingly, a synthesis of the results suggests that late collections, and we encountered none during our first several years of collecting. A few pine needle clusters (fascicles) were eventually found at one site, but based on their structure (5 needles/ fascicle, small fascicle base) they clearly represent a Haploxylon pine similar to Pinus strobus. We also found that that Haploxylon pine pollen grains dominate all of the pollen assemblages (Stults, Axsmith, and Liu 2010). These findings demonstrate that the current exclusive dominance of Haploxylon pine species in the greater Mobile-Tensaw River area developed relatively recently— sometime between 3 million years ago and 85,000 years ago. Such findings underscore the unique and sometimes surprising contri- Figure 5-3. Representative regionally extinct Citronelle Formation plant fossils: butions that paleobotany can A) fruit of Trapa alabamensis (water caltrop); B) fruit of Begonia; C) fruit of Pterocarya sp. (wingnut); D) leaflet ofPterocarya ; F) fascicle of white pine.

Biotic Landscape - Chapter 5 31 Plicente mean annual temperature (18°C [64°F]) was of grasslands (Graham 2011). One important excep- similar to modern levels or even slightly lower, and tion to this dearth of Miocene fossils is the Mauvilla annual precipitation levels (1190 to 1250 mm [46.9 to fauna, which was discovered on private land along 49.2 inch]) were lower despite the global warmth of Chickasaw Creek in Mobile County (Isphording and that time (Stults 2011; Stults and Axsmith 2015). Lamb 1971). The mammals have been described in detail and include horses, peccaries, rhinos, and many Some other proxies correlated with climate can be others. Several of the mammal taxa are age diagnostic estimated using plant fossils. For example, atmo- and indicate an age range of 6.8 to 0.3 million years spheric carbon dioxide (CO ), which is a greenhouse 2 ago (Hulbert and Whitmore 2006). gas associated with warming climates, can sometimes be estimated from fossil plants. This estimate is possi- The original account of the Mauvilla site mentioned ble because many plants reduce the numbers of gas wood fragments, but no other plants (Isphording exchange openings (stomata) on their leaves to mini- and Lamb 1971). We have discovered a layer above

mize water loss as atmospheric CO2 levels increase. the mammal fossils that also includes abundant seeds Individual species response to increasing atmospheric and leaves. Although in early stages of investigation,

CO2 can be determined from historical herbarium this flora includes many taxa still common in the area specimens (i.e., dried leaf collections) for the indus- today, including the conifers Chamaecyparis thyoides trial revolution until now, during which time atmo- (Atlantic white cedar) and Taxodium distichum (bald

spheric CO2 levels increased from 280 ppm (parts cypress), Cyrilla (titi), Nyssa (tupelo), several Quer- per million) to the current value of 400 ppm. Stoma- cus (oak) species, Sambucus (elderberry), and Vitus tal openings are sometimes well preserved on fossil (grape). Regionally extinct taxa have also been recov- plants, and these can be counted and compared with ered, including Trapa and fragments of the conifer the calibrated data set based on the modern plants, Podocarpus, which is now mainly a Southern Hemi-

thus providing an estimate of the atmospheric CO2 sphere genus. New taxa are still being found, and we concentration when the fossil plant lived (Woodward anticipate that this flora will soon be sufficiently well 1988). Using this procedure on Taxodium distichum known to allow climate analyses like those done for (bald cypress) leaves from the Citronelle Formation, the Citronelle Formation flora. Examples of Mauvilla

the atmospheric CO2 level during the late Pliocene plant fossils can be seen in Figure 5-4. was about 351 ppm, which is higher than the pre-industrial value, but lower than current levels (Stults et al. 2011). This observation suggests that the late Pliocene warm interval may have been caused by elevated

CO2, but other factors may also have contributed to greater climate sensitivity (Lea 2015).

Mauvilla Flora (Late Miocene—7.3 to 6.8 Million Years Ago) The greater Mobile-Tensaw River area is underlain by massive layers of sediment from the Miocene Epoch, but they are mostly poorly dated and few fossils of any kind have been described from them. This lack of fossils is unfortunate as the Miocene included important events like the continuation of post- Figure 5-4. Representative Mauvilla site plant fossils: A) branches of Eocene cooling and the expansion Chamaecyparis thyoides (Atlantic white cedar); B) seeds of Vitis sp. (grape); C) pit of Nyssa sp. (tupelo); D) immature acorn (Quercus sp.).

32 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Bucatunna Formation (Early Gosport Formation, Claiborne Group Oligocene—30.5 to 30.3 Million Years (Middle Eocene—41 to 38 Million Ago) Years Ago) Early Oligocene plant fossils have recently been The oldest plant fossils from the greater Mobile- collected from the Bucatunna Formation in south- Tensaw River area come from Eocene outcrops of eastern Monroe County, Alabama. This discovery is the Gosport Formation, which is part of the Clai- exciting because early Oligocene plant fossil sites are borne Group of formations, at the bluffs of Claiborne rare throughout the entire US, and this epoch was landing in southern Monroe County (Berry 1924). important for the onset of globally cooler tempera- Based on recent studies at Claiborne Group sites in tures following the extreme warmth of the Eocene, as Kentucky and Tennessee (Dilcher 2000; Dilcher and mentioned above (DeVore and Pigg 2010). Lott 2005; Devore and Pigg 2010; Wang et al. 2013), we know that many of Berry’s early identifications Numerous leaves and fruits have been collected from were incorrect. The flora now includes taxa related to the Bucatunna Formation, but are not yet well identi- modern plants in the region today, like Nyssa (tupelo), fied. At least 10 different leaf types are present, includ- and Gordonia (loblolly bay). However, there are many ing some with affinity to the Lauraceae (laurel family) extinct forms with affinities to species today in Central and Fagaceae (beech, chestnut, oak family). Although America, South America, Europe, Africa, and Asia. It hundreds of leaves have been collected, only one would be interesting to make additional collections at has marginal teeth, which is remarkable since a high the bluffs at Claiborne for updated comparisons with proportion of leaves with smooth margins is typi- the Kentucky and Tennessee sites, but the fossiliferous cally correlated with warmer climates (Chen et al. layer is now inaccessible due to extensive slumping of 2014). In this instance, however, we suspect that this overlying sediments. flora represents a specialized, marginal marine coastal community. Examples of Bucatunna Formation plant Summary fossils can be seen in Figure 5-5. It is now evident that the greater Mobile-Tensaw River area contains significant plant fossil resources. Recent studies of the plant fossil sites briefly described here are helping unravel the evolution, biogeography, and climate of this important floristic region. Although in early stages of investigation, two important gener- alizations are possible. First, plant communities have been remarkably stable, with a few interesting exceptions, since the late Miocene Epoch (approximately 7 million years ago), and probably for much longer. This plant community stability is likely due to relative climatic stability imposed by proximity to the Gulf of Mexico. Second, the marshy and deltaic environments common today have been typical of this region for millions of years (Noss et al. 2015). The Pliocene Citronelle flora is now well documented, and future work will focus on the Pleis- Figure 5-5. Representative Bucatunna Formation fossils: A) elongate leaf mor- tocene, Miocene, Oligocene, and photype; B) ovoid leaf morphotype; C) fruit with persistent sepals; D) rare leaf Eocene floras and their paleoclimate with toothed margin. implications.

Biotic Landscape - Chapter 5 33

Landscape ecology addresses ecological processes is unusually high in herbaceous diversity, and the and effects within a context of broad spatial land- eastern broadleaf forest, which is high in tree diver- scape patterning, including effects of dynamic spatial sity (Sorrie and Weakley 2001; Thorne 2008; Kartesz heterogeneity on biotic and abiotic processes, the role 2015a; Weakly 2015). For the purposes of this synthe- and consequences of periodic disturbance within sis we characterize the greater Mobile-Tensaw River and beyond a range of normal variability, and the area as part of the East Gulf Coastal Plain ecoregion, role of habitat fragmentation on species and ecologi- with a dizzying array of complex and heterogeneous cal processes (Turner 1989; Turner et al. 1989). The ecosystems organized within 3 primary systems, as components of an area’s disturbance regime include follows (Bailey et al. 1994; Comer et al. 2003). the frequency, return interval, size, intensity and severity, and residuals (see White and Pickett 1985; Turner 1) Southern Sandy and Sandy-loam Pinehills, et al. 1998; Turner 2010). While some scientific litera- Plains, and Savannas ture exists about some of these specific elements and These ecosystems, which correspond to Nature- components of the landscape and disturbance ecol- Serve’s East Gulf Coastal Plain Interior Upland Long- ogy of the greater Mobile-Tensaw River area, there is leaf and East Gulf Coastal Plain Near-Coast Pine yet no published description of the area’s overarch- Flatwoods, develop over fluvial sediments, generally ing and integrated large-scale landscape and distur- Miocene to Pleistocene in depositional origin. There bance ecology. This chapter offers an initial synthesis are myriad divisions within this category, including of how large landscape and disturbance ecology of steephead seeps, xeric uplands, pine plains and hills, the greater Mobile-Tensaw River area are organized flatwoods, and bogs. and expressed according to geologic/geomorphologic, hydrologic/precipitation, and pyric patterns and 2) Floodplain processes. This ecosystem includes deltaic and riparian zone floodplains and all estuarine areas, with two major Overview of Ecosystems of the divisions: Greater Mobile-Tensaw River Area Before delving into the specific geologic, hydro- ●● Brown water (redwater) stream floodplains and logic, and pyric patterns and processes that under- riparian zones, which correspond with Na- pin the landscape and disturbance ecology of the tureServe’s East Gulf Coastal Plain Large River greater Mobile-Tensaw River area, we should step Floodplain Forest and East Gulf Coastal Plain back and acknowledge the area as the largest inland Freshwater Tidal Wooded Swamp, and North delta complex in the United States, at the base of the Central Gulf of Mexico Salt and Brackish Tidal nation’s 4th largest river system (Sturm et al. 2006). Marsh. These areas have high flow, high con- The greater Mobile-Tensaw River area has, by far, the ductivity and fine grain sediment loads, and largest discharge (79,300 ft3/s [2250 m3/s]) of any Gulf slightly acidic to nearly neutral pH. This division of Mexico river system east of the Mississippi River includes deepwater swamps; bottomland hard- embayment, representing 35 times the discharge of all wood levees and terraces; transitional “bay” tidal the other eastern Gulf Coast river systems combined forests; freshwater to brackish water marsh; and (Tetra Tech 2012). The area lies at the intersection of submerged and floating-emergent vegetation on the greater longleaf pine savanna ecosystem, which the edges of stream channels and in deeper pools of “dead” channels and lakes.

Biotic Landscape - Chapter 6 35 ●● Blackwater stream floodplains and riparian zones, al. 1992; Shumm et al. 2000). This vertical relief, and which corresponds with NatureServe’s South- the complex mineralogy associated with it, has had a ern Coastal Plain Blackwater River Floodplain profound effect on the ecosystems surrounding the Forest, East Gulf Coastal Plain Small Stream and Delta and also on the development of the river and River Floodplain Forest and East Gulf Coastal Delta itself as it winds through these steep headlands. Plain. These areas arise in the immediate coastal As is the case with other rare areas of high relief or plain, with low conductivity and naturally very unusual soil characteristics within the coastal plain low, large-grained sediment loads. They are (e.g., the Apalachicola bluffs region), this combination highly acidic and poorly buffered. This division of factors has resulted in areas with both high total includes blackwater swamps and drains, white species diversity and unusual floral and faunal ende- cedar swamps, blackwater streamside scour mism, perhaps because these areas offered multiple zones and meadows, and submerged and float- habitat niches and remained relatively stable during ing-emergent vegetation zones. past climate change events (Graham 1999b; Fetter 2014; Noss et al. 2015). 3) Riparian Zone Bluffs, Coves, and Large- scale Erosional Cuestas The Delta is a fluvially dominated system with high These ecosystems correspond to NatureServe’s rates of discharge and high rates of sediment loading Southern Coastal Plain Mesic Slope Forest and South- comparable to other Appalachian-origin rivers in the eastern Coastal Plain Cliff and Southern Coastal Plain eastern US (Watson et al. 1986; Smith 1988; Isphord- Limestone Forest. They are typically areas of rela- ing et al. 1996). Thus, it is generally considered a tively high relief (shaped by the interaction of uplift “prograding” system, with its potential to move south- of ancient seabeds with small-scale and large-scale ward and seaward influenced by rates of sea level rise. fluvial processes) with complex mineralogy and multi- Like the Bay below it, the Delta developed over a Pleis- ple layers of highly exposed strata. These areas are tocene river valley incised in Mobile Bay some 100 ft generally occupied by highly diverse broadleaf forests (30 m) below current sea level (Smith 1988; Greene et characterized by some 25 species of oaks, 7 species of al. 2007). The surrounding headlands show succes- magnolias, high shrub and tree diversity, and wide- sive layers of Pliocene/Pleistocene alluvium laid over spread occurrence of “disjunct” Appalachian species a Miocene basement of marine sediments (Smith (e.g., beech and basswood). 1988; Raymond 1993; Isphording et al. 1996). The prehistoric river may have originally drained much of Geological and Geomorphological the Appalachians now drained by the Tennessee and Patterns and Processes other rivers. For most Appalachian slopes, sediment loss was an order of magnitude higher during the The Mobile-Tensaw Delta and the lower Alabama Pleistocene than today, so it is reasonable to assume River and its tributaries slice through unusually that the pre-Holocene Alabama River had discharge complex and highly exposed Cenozoic marine-origin rates and sediment loads much higher than at present formations, overlain with sandier Pliocene-Pleisto- (Delcourt and Delcourt 1987; Mongomery and Wohl cene alluvium (Smith 1988; Smith 1997). This process 2003). has resulted in surrounding zones of high relief, with steep riparian bluffs, coves, and—in the Red Hills Isphording et al. (1996) and Smith (1988) hypoth- region—large-scale erosional cuestas, occasionally esize that the Deltaic floodplain rose rapidly with the with sharp drops of several hundred feet to the river increase in sea level after the last low-stand 18,000 bed. Such features are highly unusual in the Southern years before present. In essence, the intersection of Coastal Plain and are believed to have been shaped Delta and Bay appears to have remained relatively by the interaction of uplift of ancient seabeds and stable for many thousands of years, in spite ongoing large- and small-scale fluvial processes. There is much sea level rise (estimated at 8 to 11 inch [20 to 28 cm] speculation on the cause of the uplift, but it appears per century) (Penland and Ramsey 1990; Williams et to be the result of active tectonics that produce the al. 1999). As Isphording (1996) describes, neither the highest rate of elevational change (approximately 4 Bay nor the Delta seem to have existed much north- mm/yr [0.2 in/yr]) of any place in the Southeastern ward of where they are now, at least since the last coastal plain (King and Beikman 1978; Leopold et interglacial. Rodriguez et al. (2010) provide a more

36 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area complex assessment of how bayhead deltas across the inch/yr [130 cm/yr]). While heavy rainfall events are northern Gulf of Mexico may have stepped back in common, sometimes with 5 inches (13 cm) or more in response to brief and rapid changes in sea level. a day, rainfall is well distributed throughout the year (with rainfall peaks in early spring and again in mid- In spite of high sediment loads, it is important to summer). Mobile also ranks high in the number of note that the Delta has low trap efficiency. That low days per year with at least 0.1 inch (0.25 cm) of rain. efficiency results because this delta is constrained by This typical abundance of rainfall has, not surpris- high headlands on either side and cannot create new, ingly, shaped the area’s geography, promoted the lower-elevation deltaic lobes that are more efficient development of extensive and diverse stream and sediment traps. Most sediment capture in the greater wetland habitats, and is critical to the area’s biological Mobile-Tensaw River area occurs only during select diversity and productivity. It is also worth noting that overbank flood events, whereas the lower Mississippi high water availability can control and even limit the Delta can spread freely and often “moves” sideways expansion and distribution of ecological communi- in search of lower elevation paths to the ocean. As a ties, just as lack of water does in drier environments. result, only about 30% of the suspended sediments High water availability also makes the area highly entering or originating in the Delta actually settle there; sensitive to small changes in rainfall, stream inflow, most are discharged into the Bay and Gulf (Isphord- groundwater levels, and water tables. ing 1996; Rodriguez et al. 2008). For this reason, Isphording (1996) suggests the Delta has been spared High precipitation generates high rates of freshwater much of the contamination that might otherwise have inflow from both local, coarse-grained, low-conduc- occurred as a result of mid- and late-20th-century agri- tivity blackwater systems and regional fine-grained, cultural and industrial water-borne pollution (with high-conductivity redwater rivers (Ward et al. 2005). some significant localized exceptions). Many of these Tidal and biological connectivity to the Gulf of Mexico contaminants are bound up in fine-grained and highly creates estuarine conditions virtually throughout the charged sediments that settle out into the Bay, where region’s waterways, with seasonal estuarine processes they have limited availability to the food chain. Essen- that create a freshwater-dominated Delta system tially, the only way to broadly increase the Delta’s trap during months with high freshwater runoff (January efficiency is to raise sea level, which lifts more sedi- through April) and an increasingly brackish-water ment out of the channels and brings it into contact system during the months with lower runoff (July with the Delta’s floodplain and filtering vegetative through November) (Schroeder and Wiseman 1999; structures. This appears to partially explain why the Valentine and Sklenar 2006; O’Neil 2007; Goecker et Delta continues a very slow progradation southward al. 2009; TetraTech 2012). In the southernmost sections and seaward in spite of current sea level rise. It may of the Delta floodplain, hydroperiod and water levels also suggest that the Delta’s floodplain and its natural are influenced more by tidal head than by river head. communities can respond to and survive a significant All but the very largest riverine flood events quickly degree of sea level rise. disperse and attenuate in the broad lower floodplain and in Mobile Bay, typically raising water levels by Hydrologic and Precipitation Patterns inches rather than feet (Tetra Tech 2012; USGS 2015). and Processes Freshwater flood levels and impacts steadily increase The greater Mobile-Tensaw River area has a distinctly upstream in the Delta floodplain, in the Alabama and seasonal climate, with large variation in average daily Tombigbee rivers, and in the surrounding blackwa- temperatures from season to season, but with an ter streams. Unusually active tectonics for the coastal unusually long growing season, typically 300 days. plain have promoted a high relief coastal zone and The northern Gulf Coast receives more rainfall than undulating to steep riparian and streamside zones, any other major region of the country. The city of exposing complex geological strata and encouraging Mobile receives more rainfall than any other metro- unusual fluvial geomorphologic processes (Byrnes et politan area in the US, at 67 inches (170 cm) annually, al. 2013). The Delta is embedded in a region where almost twice as much rainfall as Seattle. This rainfall long-term climate dynamics have been relatively stable rate is high even in comparison to many other areas compared to much of North America, resulting in the of the humid southeast US. (Rainfall on the Atlan- persistence of species (aquatic and terrestrial) eradi- tic Coastal Plain, for example, is typically under 50 cated by climate fluctuations in other parts of eastern

Biotic Landscape - Chapter 6 37 North America (Delcourt and Delcourt 1987; Graham of the Delta, where water conditions are primarily 1999b; Sorrie and Weakly 2001; Stults et al. 2009; Noss controlled by inflow from hundreds of miles upstream et al. 2015). Periodic tropical disturbances such as or by tidal inflow, water quality on most of the Delta’s hurricanes can alter the composition of ecosystems, blackwater streams is determined by local condi- though perhaps not as significantly as floods and fires tions and hydrological connectivity to the surround- (Isphording 1996; Park et al. 2007; Smith et al. 2013). ing uplands. Relatively small changes in freshwater Gilliam and Platt (2006) discuss the role large tropi- inflow as a result of house construction, road build- cal weather disturbances can have on longleaf ecosys- ing, or groundwater pumping can result in rapid loss tems and indicate that a loss of such disturbances, of stream head and rapid replacement by siltier waters which result in more open canopy conditions, could from the Delta’s redwater systems, or more saline reduce biodiversity. tidal waters. Such changes can dramatically alter floral and faunal communities. In the longleaf forests High water tables in the Delta and in various kinds of above these blackwater streams, mesic flatwoods perched wetland and flatwoods in the uplands have a and pitcher plant bogs exhibit extraordinarily high significant impact on the oxidation-reduction (redox) diversity (up to 60 species/m2 [6 species/ft2]) in part potential of soils, limiting not only the growth rate but because the plants have carefully partitioned habitat also the kinds of plant communities that occur there. as a result of minute (1 cm [0.4 inch]) changes in depth Because of this, changes of elevation of a few centi- to the water table. Road building and house construc- meters can have dramatic effects on floral and faunal tion can severely impair the sheet-flow hydrology that communities, much greater than changes in elevation feeds these highly diverse wetland systems, resulting of hundreds of feet in montane communities (Noss in the loss of hundreds of species of plants and insects et al. 2015). Likewise, seemingly minute increases or from relatively small areas. decreases in water levels can play a significant role in the success or failure of species and communities Interactions of tidal and freshwater influences are within and around the Delta. vital to the ecosystems in this region. Virtually every waterway from the Gulf of Mexico to the first dams Mature bald cypress (Taxodium distichum) are the on the Alabama and Tombigbee rivers, almost 100 signal trees of the Delta and are ecologically restricted water miles (160 km) inland from Mobile Bay, are to wet or frequently flooded sites. But seed germina- tidally influenced, and influenced by saline density tion and seedling growth requires moderately dry and currents much of the year. While tidal dynamics are highly aerobic topsoils, which occur in the Delta only moderate compared to many sites fronting the Atlan- after an extended period of low water and low water tic or Pacific, large-scale wind-driven water level “set- tables. Such a period may occur seasonally every ups” in the Gulf can greatly amplify daily tidal fluc- year in some areas, or once every few years in others. tuations, and do so throughout the year. Though the The frequency of these occurrences can be dramati- brief surges associated with tropical systems are most cally altered either by timber harvest (which results notorious, non-tropical set-ups can last for days or in longer hydroperiod and significantly higher water sometimes weeks and often have more impact (Schro- tables throughout the year) or by changes in seasonal eder and Wiseman 1999). flow variability of adjacent streams, either through increased runoff from development or through The Delta is often described as a seasonal estuary. It is seasonal manipulation of flow rates by upstream a freshwater system while evaporation and transpira- dams. Under such conditions, swamp tupelo (Nyssa tion are low and inflow from upstate is high. Because aquatica), whose seedlings are more tolerant of flood- the Delta estuarine system dissipates its freshwa- ing, can displace cypress. In many areas of the Delta, ter inflows over a large floodplain and bay system, flow rate manipulation appears to have been used the hydrological head of the Gulf plays a significant to promote pure stands of swamp tupelo, which are role in determining hydroperiods and flood levels, preferred for paper pulp processing (Messina and mitigating the effect of flood and drought on water Connor 1998; Evans et al. 2013). levels. Less often the opposite happens: dramatic increases and decreases in water levels occur that are Even more dramatic consequences of hydrological larger than those attributable to drought and flood. alteration affect the blackwater streams and pitcher The estuary becomes more brackish as inflow from plant bogs surrounding the Delta. Unlike the interior upstate decreases in summer and fall, and evaporation

38 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area and transpiration increase. This seasonal transition to of the bay by removing loose sediments (Isphording a brackish system greatly alters the aquatic commu- 1996). While hurricanes apparently play a significant nity, mixing marine predators (e.g., tarpon, speckled role in sediment accumulation in Mobile Bay salt trout) and prey species (e.g., shrimp, anchovies) with marshes, floods seem to play a more important role freshwater species such as bream and bass (Lowe et in the Delta itself (Smith et al. 2013). Small magnitude al. 2009; Norris et al. 2010; Carassou et al. 2011; Lowe but long duration wind-forced changes in water levels et al. 2011; DeVries 2013; Rozas et al. 2013). Seasonal may play an equally large role in community struc- transitions may also play a role in mediating domi- ture. In winter they can expose mud flats for weeks, nance in the herbaceous and woody plant communi- and in spring and summer they can result in weeks of ties in the lower Delta (Myers and Ewel 1990; Light et much higher than normal tides that flood usually dry al. 2002). habitats. Flooded or exposed, these are exceptionally productive habitats for mudflat crustaceans and In the lower Delta, hydroperiods are less significant mollusks, as well as shorebirds and even fish, includ- than changes in conductivity as a result of tidal influ- ing the threatened Gulf sturgeon (Peterson and Peter- ence. Moving upstream in the Delta floodplain, and son 1979; Randall et al. 2006). along the river and local stream channels, the influence of tidally influenced conductivity declines and Droughts are perhaps equally significant, resulting the impact of hydroperiod increases. As is typical in dewatering of fresh groundwater throughout the of estuarine systems, the tidal head forms persistent Delta and longer term elevation of salinity high up in density currents in the channel thalwegs—heavier the Delta. These drought effects, in turn, can increase saltwater currents that have the appearance of a salt- habitat and access to vegetative and hard structure for water wedge as they move up the channels (Schroeder estuarine-dependent species like white shrimp. At the and Wiseman 1999). During floods, these density same time they can reduce gross productivity, slow currents may be restricted to the deepest channels in down eutrophication of some water bodies, and slow the lowermost Delta. As the hydrological head of the the regeneration and advance of trees and shrubs into river decreases in late summer, the saltwater wedge freshwater marshlands (Rozas et al. 2012; Rozas et al. penetrates as far as the dams upstream (and likely 2013). once went even farther north) and mixing of fresh and salt water increases throughout the water column. Hydrological and landscape continuity is unusually In the lower Delta, the entire water column typically important in this region. Water quality and productiv- becomes brackish in fall. This is a critical breeding ity in the Delta is the product of the myriad streams and development time for many marine and estua- surrounding it, and the hydrology of these streams rine fauna—ranging from white shrimp to speckled is linked to percolation and water retention rates of trout—which take advantage of the increased salinity the surrounding uplands. Development can compro- to forage and develop in the midst of the complex and mise the surrounding landscape’s ability to serve as a productive structure of Delta vegetation, shell reefs, water reservoir, resulting in greater runoff immedi- and organic soils. ately following rains and less residual freshwater days after a rain. Cycles of flood and drought are ampli- Hurricanes and wind may also play a role in commu- fied. Because the blackwater streams surrounding the nity structure, at least temporarily, and occasionally Delta serve as important biological refugia and nurs- with long-term impacts. Wind impacts to terrestrial eries, loss of connectivity between the Delta’s brown systems (e.g., downed timber) are perhaps most obvi- water aquatic habitats and these blackwater streams ous, but the most significant impacts are usually the can directly impact the Delta’s biological diversity. result of wind-forced water surges. These brief but dramatic rises and falls in sea level can be devastating Fire to human structures, and can lead to higher salinities. Oddly enough, the area’s high rainfall is also respon- In most cases, impacts on the ecosystems are gener- sible for the ecosystem’s long and critical association ally short-lived (Park et al. 2007), but under the right with fire. Thunderhead development and rains during conditions hurricanes can significantly rearrange sedi- the growing season on the Gulf Coast are inevitably ments and even influence geomorphology. In 1979 accompanied by lightning. The highest flash density Hurricane Frederic, a moderately large storm, rapidly of lightning in North America (and one of the highest drained the bay and measurably increased the depth

Biotic Landscape - Chapter 6 39 densities in the world) occurs in a narrow band around on ecosystems (see Frost 1993; Noss 2013; Ryan et al. the Gulf Coast and in the Florida peninsula (Noss 2013). 2013). On moister north- or east-facing slopes, and on Given this high frequency of ignition, the only limit slopes and banks immediately surrounding streams, to the frequency of fire historically was the availabil- fire return intervals and fire intensity would have ity of combustible fuel. Because of high rainfall, fuel been somewhat less, typically once every 5 to 25 years loads (flammable vegetation) could build up quickly (Harper 1943; Myers and Ewell 1990; Frost 2000; Frost on most sites. Longleaf pine itself, along with the oil- 2006). These areas also typically overlap exposures of rich herbaceous grassland community it supported, multiple geological strata, each supporting a unique is responsible for “nursing” ignition, increasing the chemical and sedimentary signature, which allowed chances that fires would occur. Within the long- development of a very different forest community, one leaf pine forest ecosystem surrounding the Delta, more resembling the Appalachian forests of central fire return intervals were typically from 1 to 3 years. North America. Higher content of magnesium and Because of their high frequency, fires were usually calcium and a higher redox potential encourage colo- only of moderate intensity, and the vegetative commu- nization by plant species seldom encountered in the nity recovered rapidly, often in a matter of days. In more common, coarse-grained acidic soils. Broadleaf these areas, virtually all species were fire tolerant, or trees (e.g., oaks, magnolias, hickory, beech) domi- even fire dependent (Myer and Ewell 1990; Jose et al. nate the canopy, with a wide assortment of shrubs 2006; Palmquist et al. 2014). Many species in these and ephemeral herbaceous plants that respond to plant communities (e.g., wiregrass, multiflowered brief, seasonal exposure to sunlight. In areas of full calopogon orchid, toothache grass, American chaff- habitat development, along the lower Alabama River seed) flower only after growing season fires. These at the top of the Delta, tree species diversity may be frequent fires restricted development of common tall higher than almost any place else in North America, shrubs, vines, and trees, which opened up space for with some 25 species of oaks, 7 species of magno- an unusual diversity of plant species, up to 60 species/ lias, and many other species occupying a very small m2 (6 species/ft2) and 140 species per 1000 m2 (10,800 area. There is indication that the Red Hills of south- ft2) in some cases (Peet and Allard 1993). These west Alabama support the highest oak diversity (28+ frequently burned habitats represented some of the oak species) for North America (Kartesz 2015b). But densest concentrations of plant diversity on the planet even in these habitats, occasional fire trickling in from (Peet et al. 2014). the adjacent longleaf forests is necessary to maintain diversity, and loss of connectivity between these areas Even with abundant lightning, large-scale landscape and the surrounding longleaf ecosystem has resulted connectivity is critical to fire-maintained ecosys- in decline in oak regeneration and the decline or loss tems, and fragmentation of the landscape is unusually of many other species (Adams 1992; Barnes and Van devastating. Only one well-timed lightning strike was Lear 1998). necessary when fires burned uninterrupted “river- to-river.” Today roads, fields, exurban and suburban While fire was critical right up to the banks of the development, and active fire suppression limit the Delta, fires within the Delta would likely have been spread of any one strike. Some researchers estimate rare. Fires would certainly have occurred in cypress- that tens of thousands of acres of relatively unbro- tupelo swamps, and may be necessary for their long- ken pyrogenic landscapes may be required to support term health and reproduction, but the return interval natural fire frequencies with lightning as the sole would have been measured in centuries (Messina and source of ignition (Lindenmayer and Fischer 2006; Conner 1998). White cedar swamps, a subcompo- Bailey et al. 2007). Otherwise, prescribed burns must nent of blackwater swamps, require catastrophic fires, be used to supplement or supplant natural sources with a return interval of 50 to 100 years, for regenera- and spread of ignition. The relationship of fire ecol- tion. Lack of fire and loss of connectivity to longleaf ogy to pre-Columbian cultures is controversial and uplands is likely responsible for the continued loss of complex, but along the Gulf Coast, at least, natural fire this ecosystem (Laderman 1989; Messina and Conner frequency is so high, limited not by ignition but rather 1998; Atkinson et al. 2003). Fire may have played a by the availability of fine fuels, that additional ignition more frequent role in the development of freshwater by humans is not thought to have had a major impact and brackish water marsh habitats (Myers and Ewell

40 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 1990; Noss 2013), though there has been little effort surrounding upland habitat, will have no benefits for to identify long-term fire trends in the lower Delta. the turtles. Some of these turtle species are primarily A fairly well-documented history exists of recent upland species and use the Delta rarely or not at all. anthropogenic burning in the Delta marshes from the Most of those that do are threatened above all by loss 19th through the first half of the 20th century. But it’s of suitable upland (non-flooded) habitat for egg laying harder to imagine how these islands of wet marshes, and hatchlings. A great number are lost while trying fragmented and isolated by deep swamps or rivers, to lay eggs on the grassy highway medians that run would have ignited naturally with much frequency. through the Delta (Nelson 2003). The decline of the Delta’s once abundant canebrake habitats raises other questions about the possible While there are many uncertainties associated with interaction of fire and periodic flooding and the loss region-specific impacts of climate change, we may be of connectivity to longleaf pine ecosystems, particu- able to assess the relative vulnerability of the Delta and larly along the riverine corridors of the northern Delta surrounding region based on previous climate fluc- and the lower Alabama River (Noss 2013). tuations. The Gulf of Mexico’s stabilizing influence on the climate of the area is often cited as one of the Heterogeneity within and between upland and wetland reasons for the persistence of Miocene, Pliocene, and landscape communities is promoted and mediated by Pleistocene floral and faunal elements that have disap- fire, which makes continuity between communities peared from much of the country (Graham 1999b; and across broad landscapes particularly important. Noss et al. 2015). Evidence exists that this Delta may Because of this fire dependency, a self-maintaining also be somewhat buffered from sea level rise. Unlike landscape would require tens of thousands of acres the sediment-limited estuarine systems across much of relatively unbroken, fire-maintained forest or of the Gulf Coast, this Delta’s primary floodplain has a savanna, with connectivity to surrounding habitats sediment surplus to bank on as sea level rises. As such, where fire return intervals may be longer. Because of it appears to be better equipped than most estuaries to fragmentation, sources of ignition must be artificially handle moderate levels of sea level rise, even at levels introduced into ecosystems and the fires managed higher than in the past century. Less clear, however, on a regular basis. Even with artificial prescribed is the rate or degree of sea level rise that will begin to fire regimes, relatively large unbroken landscapes overwhelm the potential increase in trap efficiency. are needed to manage fire intensity and frequency. Loss of connectivity between longleaf uplands and Sea level rise, even at existing moderate levels, is surrounding broadleaf and wetland communities has much more likely to impact anthropogenic features resulted in greatly reduced fire frequencies in those in and around Delta, which have no built-in adapta- areas. These “ecotonal” marginal areas are integral to tions to sea level rise. Ancient and historic features, the existence of many, if not most, of the listed engen- such as shell middens, the Mississippian mounds, dered, threatened, and special concern species (e.g., colonial and Civil War sites, and even the modern Red Hills Salamander, Reticulated Flatwoods Sala- city of Mobile, continue to subside and face severe mander, American chaffseed, pot of gold lily, Wher- erosive events as sea level continues to rise. Efforts ry’s pitcher plant). to conserve and preserve these features will have to take this challenge into account. Also more likely to Vulnerabilities and Challenges be impacted are the unusually rich natural communities on the margins of the Delta and its feeder streams The movement of wildlife from floodplain to uplands and uplands. Blackwater streams and basins and their and back again has already been compromised by surrounding floodplain communities, for example, residential and commercial areas, roads, and other might normally rise with sea level to occupy slightly developments. Many endangered or rare species are higher elevations. But residential and urban develop- highly dependent on continued connectivity between ment is concentrated along the blackwater streams, so uplands and wetlands (e.g., indigo snakes, black bear, there are fewer and fewer areas where these commu- freshwater turtles, numerous bird species). The 3 nities can go. For this reason, remaining undeveloped counties surrounding the Delta support what is likely uplands surrounding these blackwater streams should the greatest turtle diversity in the world. But protect- be prioritized for conservation. ing the Delta alone, without giving attention to the

Biotic Landscape - Chapter 6 41

7. Ecoregional Basis for Biodiversity Richness and Endemism R. Scot Duncan, Birmingham Southern University

The greater Mobile-Tensaw River area is one of the lower Alabama and Tombigbee river areas have the most biologically rich and ecologically diverse weathered and sliced into these sediments to create regions of its size in the Southeast, and possibly in a topographically complex landscape, despite being all of temperate North America. Within this area is a dubbed a “plain” (Lacefield 2013). Any change in dazzling range of ecological conditions, from sunny elevation, aspect, and soil type alters the sunlight, fire-dominated woodlands to blackwater swamp groundwater, and nutrients available for plant growth. forests to steep shady ravines (Duncan 2013). This Subsequently, tremendous ecosystem diversity is diverse collection of highly productive ecosystems sustained across the Atlantic Plain and represented in sustains such a profusion of species that some refer to the greater Mobile-Tensaw River area (Duncan 2013). it as “America’s Amazon.” There is no single reason for why this region is so rich. Instead, the influences of The third biogeographic realm shaping the greater 3 forces have shaped this region’s biological diversity: Mobile-Tensaw River area is the waters of the Mobile the intermingling of 3 distinct biogeographic realms, River Basin. The Mobile River Basin drains a major- a benevolent climate, and the unstoppable forces of ity of Alabama, and minor portions of Mississippi, ecological change. This chapter explores how these Tennessee, and Georgia (Mettee et al. 1996). Due to forces created the greater Mobile-Tensaw River area the interaction between the Southeast’s benevolent and its diverse landscape. climate and the southern Appalachian Mountains, the basin has been the origin point for hundreds of Biogeographic Realms freshwater species, especially mussels, snails, fishes, and crayfishes. Consequently, the Mobile River Basin The biogeographic realms shaping the greater is a global hot spot for aquatic biodiversity (Bosc- Mobile-Tensaw River area are the Gulf of Mexico, the hung and Mayden 2004; Williams et al. 2008; Duncan Atlantic Coastal Plain, and the waters of the Mobile 2013). Many of its species, including basin endemics River Basin. The Gulf is a warm ocean basin fed by and more widespread species, inhabit the waters of a continuous stream of warm, clear waters from the the greater Mobile-Tensaw River area (Boschung and Caribbean Sea. Along the northern coast these waters Mayden 2004; Schuster and Taylor 2004; Williams et are infused with a broth of nutrient- and sediment- al. 2008). laden freshwater spilling off of the rainiest region of temperate North America (Gore 1992; Oey et al. Climate 2005). The resulting mixture supports a flourishing array of marine ecosystems, many of whose species The region’s generous climate is the second force migrate into the Mobile River Delta to breed or mature shaping the biodiversity of the greater Mobile-Tensaw (Wallace 1997). River area. The region receives plentiful rainfall and sunlight relative to most other regions of North The Atlantic Plain physiographic region, in which America (Chen and Gerber 1990; Chaney 2007). This the greater Mobile-Tensaw River area is embedded, combination of moisture, light, and heat sustains high extends from New England to the Yucatan Penin- rates of primary productivity in terrestrial and aquatic sula (USGS 2010). The region is composed of ancient systems, and sustains globally significant floral and marine sediments exposed as sea levels gradually faunal diversity. The only temperate region on earth receded from a highstand peaking 145 million years with comparable biological productivity and biodiver- ago (Lacefield 2013). Across this plain are sedimen- sity is Southeast Asia (Duncan 2013). tary bands, each of which is composed of semi- consolidated or unconsolidated materials distinct in Frequent lightning along the Gulf Coast (Chen and their structure, chemistry, and erodibility (Osborne Gerber 1990) is another climatic feature promot- et al. 1989; Lacefield 2013). The surface waters of ing biological diversity in the greater Mobile-Tensaw

Biotic Landscape - Chapter 7 43 River area. Lightning-generated fires regularly purge Fluctuating sea levels during the Pleistocene have ecosystems of shrubs and trees that would otherwise been implicated in promoting the evolution of new usurp the light needed by the region’s great diversity fish species along the coastal plain (Near et al. 2003). of wildflowers and grasses. The most celebrated pyrogenic ecosystem is the longleaf pine woodland, but fire Subregions Within the Greater Mobile- played a crucial role in many of the greater Mobile- Tensaw River Area Tensaw River area’s ecosystems (Duncan 2013). Three factors, as described above—the intermingling of 3 biogeographic realms, a benign climate, and the Ecological Change pervasiveness of ecological change—have created The third force shaping the greater Mobile-Tensaw subregions within the greater Mobile-Tensaw River River area is ecological change. Landscapes subjected area with distinct ecologies and unique amalgama- to a moderate degree of change—whether on a regu- tions of species. An effective tool for classifying such lar or irregular basis—often sustain a greater diver- subregions throughout North America was developed sity of species than more stable landscapes (Krebs by the US Environmental Protection Agency and other 2009). Examples abound across varied time scales. entities in an effort to classify lands based on physiog- As anglers can attest, daily tides facilitate the mingling raphy, geology, soils, flora, fauna, and other variables of both freshwater and marine species in the lower (Commission for Environmental Cooperation 1997). Delta. Seasonal changes in freshwater, sediments, and Known as the ecoregion approach, lands are classified nutrients flushing through the greater Mobile-Tensaw in a 4-tiered hierarchy. River area cause the expansion and retreat of the Delta’s marshland (Smith 1988). Many aquatic species Level IV ecoregions are the most specific tier of clas- correspondingly migrate in and out of the Delta, sification (Griffith et al. 2001a) and the remainder of including crustaceans, fishes, and even a population this chapter explores their occurrence in the greater of the West Indian manatee (Pabody et al. 2009). High Mobile-Tensaw River area. Generally, ecoregions are above these waters, the proliferation of spring plant described in terms of their historical condition (5000 growth feeds a profusion of insects, which is fuel for to 200 years ago), instead of their current condition, birds arriving from the tropics. The onset of winter which is often radically different due to anthropogenic dormancy in the greater Mobile-Tensaw River area’s forces on the landscape. The importance and effect of forests forces a nearly complete turnover in the bird these latter influences are discussed briefly at the end. species present between summer and winter. 1) Gulf Barrier Islands and Coastal Marshes Changes over the intervals of years and decades also Ecoregion sustain high levels of biological diversity. The original The lower Delta is a labyrinth of shallow bays, chan- fire-return interval for the longleaf pine woodlands nels, marshes, and low-lying forests all belonging to and other fire-dependent ecosystems of the greater the Gulf Barrier Islands and Coastal Marshes Ecore- Mobile-Tensaw River area is not known for certain, gion (classification number 75k). Herbaceous growth, but was likely 1 to 3 years (Frost 2006). Hurricanes including grasses, iris, lilies, rushes, arums, and other periodically blow down trees en masse in forests and non-woody plants crowd the shallow waters and adja- woodlands, thereby creating space for opportunis- cent low-lying lands. Trees tolerant of waterlogged tic species to temporarily thrive and increasing the soils survive at slightly higher elevations. This ecore- chance of intense wildfires (Liu et al. 2007). gion is found intermittently from Louisiana through Geologic and climate change over the millennia the Florida Panhandle, and in Alabama it includes salt has also enriched the greater Mobile-Tensaw River marshes and barrier island ecosystems. area’s biological diversity. During the Pleistocene This ecoregion includes the most dynamic of the (2.6 million years ago to 11.7 thousand years ago) greater Mobile-Tensaw River area’s ecosystems, given there were dramatic fluctuations in sea level and that the abiotic factors controlling it vary daily and climate (Lacefield 2013). During cold glacial periods, seasonally in dramatic fashion, and consequently northern species colonized the Southeast, and relict species diversity is quite high. Each day the rising tide populations persist in places sheltered from wildfire pushes back against the riverwater to bring salt-laden and the summer sun (Duncan 2013; Lacefield 2013). waters and marine animals into the lower Delta. The

44 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Alabama coast receives one high and one low tide At slightly higher elevations throughout this ecore- each day and water levels change hourly. The magni- gion are floodplain forests. During winter and spring, tude of tidal change within a daily tidal cycle can range they are flooded with muddy river water for weeks from 67 cm to 3 cm (26 to 1.2 inches) at the mouth or months. Tree species composition varies depend- of the Delta, depending on the lunar phase (NOAA ing on the depth and duration of flooding, but all are 2015). Winds and extremes in atmospheric pressure lush and diverse because water is always available and can augment or dampen these extremes (Smith 1988). nutrient-rich silt is deposited each year (Sharitz and Due to elevation gradients within the lower Delta, Mitsch 1993). Many trees support epiphytic orchids, some areas are rarely inundated, others are rarely ferns, and bromeliads. Aquatic wildlife is abundant, exposed, while many areas fall somewhere in between and diversity of fishes, reptiles, and amphibians is (Smith 1988). Consequently, variation in elevation, world renowned (Duncan 2013). Terrestrial mammals tidal magnitude, and exposure to saltwater govern the are abundant when water levels are low, and birds composition of plant communities in the lower Delta. are plentiful in migration and during the summer. Some areas are dominated by tall grasses, others by The region’s productivity nurtured the rise of Bottle floating plants, and still others only by submerged Creek, one of the most important pre-Columbian vegetation (Duncan 2013). southeastern Native American cities (Brown 2003a). With notable exceptions endemism to the upper and There are also dramatic seasonal changes in the lower lower Delta seems to be minimal. Most species in Delta. From winter through spring, high volumes these ecosystems are widespread, due in part to the of cold nutrient- and silt-laden freshwater pour periodic mixing of aquatic and semi-aquatic species through the Delta’s channels. Where fresh silt accu- along the coastal plain whenever sea levels dropped mulates, new marshland appears in the late spring during the Pleistocene’s interglacial periods. when temperatures warm, the flooding subsides, and plant growth peaks (Finch 1999a; Duncan 2013). 3) Southern Pine Plains and Hills Ecoregion From late spring into late fall discharge from the basin The uplands bordering the Delta host radically differ- wanes (with the exception of the occasional tropical ent ecosystems that collectively are part of the South- cyclone), and marine waters have more sway in the ern Pine Plains and Hills (65f) Ecoregion. Sediments lower Delta. During this time juvenile crabs, shrimps, here are clays and well-drained sands, weathered fishes, and other saltwater species use the lower Delta remains of marine sediments deposited in the past as a nursery. Higher salinities brought by high tides 23 million years (Osborne et al. 1989; Griffith et al. and waves generated by the daily sea breeze cause 2001b). The resulting topography is predominantly seasonal dieback of freshwater marshes in vulnerable broad hills and level plains, but small streams create areas (Finch 1999a; Duncan 2013). narrow riparian zones and ravines (Duncan 2013).

2) Floodplains and Low Terraces Ecoregion Prior to European-African settlement, this land- Upper delta ecosystems are the heart of the greater scape was dominated by the longleaf pine woodland, Mobile-Tensaw River area, including its river chan- a savanna ecosystem with scattered pines. Abundant nels, oxbow lakes, ponds, swamps, and floodplain light striking the ground, combined with frequent forests (Griffith et al. 2001b). This is the Floodplains rain, sustained hundreds of herbaceous plant species, and Low Terraces Ecoregion (the two Level IV ecore- and subsequently supported a great diversity of verte- gions comprising the upper portions of the Delta, brates, particularly amphibians and reptiles (Frost Southeastern Floodplains and Low Terraces [65p] and 2006; Means 2006; Finch et al. 2012). This savanna- Floodplains and Low Terraces [75i], are treated here woodland ecosystem is the most biologically diverse together). Trees are the dominant vegetation due to ecosystem in North America outside of the wet trop- being beyond the reach of the salty tides. Historically, ics (Finch et al. 2012). bald cypress populated many of the shallower areas where seasonally low water levels allowed seedlings The longleaf woodlands were sustained by the to establish. But logging and upstream dams vastly frequent recurrence of low-intensity wildfires that reduced the extent of cypress swamps. In their place killed fire-intolerant broadleaf trees and shrubs are stands of fast-growing swamp and water tupelo continuously attempting invasion from nearby ripar- (Finch 1999b; Griffith et al. 2001b; Duncan 2013). ian ecosystems (Frost 2006; Duncan 2013). Within this ecosystem, plant community composition varied with

Biotic Landscape - Chapter 7 45 subtle changes in drainage patterns (Peet 2006). More The Red Hill’s original ecosystems included pyro- dramatic topographic changes created other ecosys- genic broadleaf and longleaf pine woodlands on the tems, including the seepage bogs famed for a diversity hilltops, ravine forests on the steep slopes, and ripar- of carnivorous plants (Peet 2006; Duncan 2013). ian forests along the small and large streams of the region (Mount 1975; Griffith et al. 2001b). Because the Endemism to this ecoregion is high, but given its ravines are sheltered from the sun and receive ground- great expanse across the Southeast, most species water from the slopes above, they remain moist and have historic distributions extending beyond the cool throughout the summer, thus providing refuge greater Mobile-Tensaw River area. But given the near- from wildfires and the harsh summer sun for many complete loss of this ecosystem to land use change heat- and fire-intolerant plant species more abundant (Frost 2006), every surviving population of longleaf- at northern latitudes. Some are relict populations that associate species has great conservation value. It is established in the Pleistocene. Ironically, they grow worth mentioning that the width of the Mobile River alongside representatives of tropical plant families Delta, and even that of the swamps and floodplain that adapted to the warm temperate zone climate, but forests of the Tombigbee and Alabama rivers to the also cannot tolerate wildfire (Duncan 2013). north, were range expansion barriers to many small vertebrates. More than a few taxa are found only to Anthropogenic Change the west or east of these river systems (Mount 1975). While the forces that created the ecosystems of the 4) Buhrstone/Lime Hills Ecoregion greater Mobile-Tensaw River area are grand and enduring, they alone cannot sustain the region’s biodi- At the very northern margin of the greater Mobile- versity in the face of anthropogenic change. Much has Tensaw River area is a region known as the Red Hills. been lost already. An earthen causeway interrupts the This area of steep hills and deep ravines belongs to the natural exchange of fresh and salt water in the lower Buhrstone/Lime Hills (65q) Ecoregion. Like much of Delta. Cities and farms far to the north send pollut- the greater Mobile-Tensaw River area, these ecosys- ants into the waters feeding the Delta. Dams upstream tems have received little scientific scrutiny and new interrupt the migrations of fishes and distort the natu- discoveries should be expected. The hills exist due to ral hydrology. The overharvest of plants and animals Eocene and Oligocene marine deposits of claystone, has weakened or extirpated many populations. Many sandstone, and limestone that are more resistant to exotic invasive species have permanently established erosion than the softer sediments of the surrounding and have reshaped the greater Mobile-Tensaw River region (Osborne et al. 1989; Lacefield 2013). In addi- area’s ecology. Two centuries of land use change have tion, a salt dome known as the Hatchetigbee Dome has reduced the greater Mobile-Tensaw River area’s natu- added to the high relief of the region and contributed ral upland ecosystems to an assortment of slivers and mineral springs to the landscape (Lacefield 2013). small patches. Sea level rise over the next century The unique soils and extreme topography of the will cause retraction of marshland in the lower Delta. Red Hills relative to the surrounding plains sustains Fortunately, remedies and mitigations exist, but only species usually found farther north and above the Fall for a public willing to protect the greater Mobile- Line, at the extreme inland limits of the Coastal Plain Tensaw River area from these threats. (Mount 1975; Boschung and Mayden 2004). Due to its uniqueness and geographic isolation, this ecoregion may harbor more endemic species than any other within the greater Mobile-Tensaw River area. The best known example of this endemism is the ecoregion’s flagship species, the Red Hills Salamander.

46 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 8. Vegetation Communities L. J. Davenport, Samford University

This chapter provides a glimpse of the vegetation Bartram canoed to the head of the Delta, the conflu- communities of the greater Mobile-Tensaw River ence of the Tombigbee and Alabama rivers and the area, first through review of the historical observa- Cutoff. He ascended the Tombigbee River and, “just tions of William Bartram and Chareles T. Mohr, then within its capes,” entered “a large lagoon, or capa- through more contemporary works. It concludes with cious bay of still water,” many acres in extent and brief discussions of the Delta today, and how to access dominated by American lotus (Nelumbo lutea). He it. described fully its leaves, flowers, and fruits, even the sweet taste of its seeds. William Bartram’s Lyrical Descriptions William Bartram, in his famous Travels (1791) through Finally, Bartram stopped at a bluff and, opposite it, the southern British colonies of eastern North Amer- “a district of swamp or low land, the richest I ever ica, arrived in Mobile in July 1775 after a 2-year trek saw. . . . [A]s for the trees I shall forbear to describe from Savannah, Georgia. Eager to experience and them, because it would appear incredible” that the explore the Delta that he had hurriedly crossed, he bald cypress (Taxodium distichum), green ash (Fraxi- immediately returned east in a “trading boat” to nus pennsylvanica), American sycamore (Platanus “Taensa” (now Lower Bryant’s Landing in northern occidentalis), Eastern cottonwood (Populus deltoides), Baldwin County, Alabama), then procured a “light sweetgum (Liquidambar styraciflua), “and others, are canoe” to travel upriver. There he found river cane by far the tallest, straitest and every way the most enor- (Arundinaria gigantea) growing “to a great height and mous that I have seen or heard of.” He was equally thickness” and discovered a new species of evening impressed by the sizes of the canes, “thirty or forty primrose (Oenothera grandiflora), “perhaps the most feet [9 to 12 m] high, and as thick as a man’s arm.” pompous and brilliant herbaceous plant yet known to exist.” Over 100 years later—after the seeds that Descriptions of Charles T. Mohr Bartram collected that day and sent to England had More than a century after Bartram’s short visit—and spread widely across Europe—this primrose formed after 40 years’ study—Mobile pharmacist and bota- the basis for Hugo de Vries’ critical studies of plant nist Charles T. Mohr described the vegetation of the mutations (Cleland 1935; Davenport 2011). Delta in much greater (though less poetic) detail in his life’s work, Plant Life of Alabama (Mohr 1901). I Continuing upriver, Bartram paddled through “high cannot improve upon Mohr’s work. So for the rest forests and rich swamps” with “the Canes, and Cypress of this treatment, I will use his words to describe the trees of an astonishing magnitude, as were the trees vegetation communities of the Delta, while provid- of other tribes.” Then leaving the river channel “and ing modern nomenclature (Kral et al. 2011; Alabama penetrating the awful shades,” he passed through Plant Atlas 2015) and additions as needed. “stately columns of the Magnolia grandiflora. . . . What a sylvan scene is here! . . . I recline on the verdant bank, 1) Cypress Swamps and view the beauties of the groves”: red buckeye Mohr was one of the last botanists to visit and describe Aesculus pavia A. parviflora ( ), bottlebrush buckeye ( ), unaltered cypress swamps in the Delta, what he Prunus caroliniana Carolina laurel cherry ( ), oakleaf termed “paludial arboreal associations” or “cypress Hydrangea quercifolia Ilex hydrangea ( ), dahoon holly ( brakes.” Mohr (1901) described them thusly: cassine), pyramid magnolia (Magnolia pyramidata, which he described here as a new species), odorless The bottom lands of the Mobile River and the wax-myrtle (Morella inodora), Carolina buckthorn islands in the delta . . . are covered with a high (Frangula caroliniana), two-winged silverbell (Hale- forest of deciduous trees, common to them sia diptera), crossvine (Bignonia capreolata), coral and the lowlands of the same character along honeysuckle (Lonicera sempervirens), and Eastern the Tombigbee and Alabama rivers. . . . Where gum bully (Sideroxylon lanuginosum).

Biotic Landscape - Chapter 8 47 the banks are almost perpetually submerged water level [Figures 8-1 and 8-2]. Black gum they are covered with cypress. This largest of [Nyssa biflora], water oak [Quercus nigra], water the Atlantic forest trees was formerly found in hickory [Carya aquatica], green ash [Fraxinus the upper part of the river delta in great perfec- pennsylvanica], more rarely Southern red oak tion. The mighty trunks rise to a total height [Quercus falcata], [American] elm [Ulmus amer- of from 100 to 120 feet [30 to 37 m] and over, icana), [Eastern] cottonwood [Populus deltoi- with a diameter, measured above the buttresses des], and overcup oak [Quercus lyrata] . . . form which expand the bases, of from 3 to over 5 the high forest overshadowing the smaller trees, feet [0.9 to 1.5 m]. . . . The assemblage of these of which the most conspicuous are planer tree monarchs of the forest in the compact cypress [Planera aquatica], red maple [Acer rubrum], brake, surrounded by the peculiar cone-shaped [Eastern hop-hornbeam (Ostrya virginiana)], excrescences (cypress knees) rising from their [and several species of hawthorn (Crataegus)]. roots 1 to 2 feet [0.3 to 0.6 m] and more above the dark unruffled surface of the water, presents In addition, Mohr (1901) noted American snowbell a feature in the arboreal flora . . . at once strange (Styrax americanus), devilwood (Cartrema ameri- and imposing. In these brakes the tupelo gum cana), possum-haw (Ilex decidua), Georgia holly (Ilex (Nyssa aquatica) is the only associate of the longipes), and parsley hawthorn (Crataegus marshal- cypress, which it rivals in size, and the Carolina lii) “form the shrubby undergrowth. The handsome swamp ash (Fraxinus caroliniana) is the only [two-wing] silverbells, the dahoon holly, and the tree of small size thriving in the gloomy shade swamp dogwood (Cornus foemina) occupy the dried beneath these trees. outskirts of these swamps.” In the depths of these “drier” swamps, Mohr (1901) 2) Wet Hardwood Sites found that cabbage palm (Sabal palmetto) “reaches its Mohr (1901) went on to describe the plant associa- perfection, the trunk[s] rising from 2 to 3 feet [0.6 to tions on slightly higher ground: 0.9 m] above the ground, the fan-shaped leaves with their stalks 8 to 10 feet [2.4 to 3 m] long. Black willow In the mire of the swamps above the level of long- (Salix nigra) and cottonwood cover the recent allu- continued overflows, a variety of hardwood vium.” And deeper in these swamps are sensitive fern trees mingle with the cypress and finally super- (Onoclea sensibilis), royal fern (Osmunda spectabilis), sede it on ground slightly above the ordinary Virginia chain fern (Anchistia virginica), green arrow-arum (Peltandra virginica), and swamp spider-lily (Hymeno- callis occidentalis).

In the extremely rich and diverse habitats of these swamps, Mohr (1901) found that the shallow pools were filled with awl-fruit sedge (Carex stipata), common fox sedge (Carex vulpinoidea), rice cut-grass (Leersia oryzoi- des), white cut-grass (Leersia virginia), dense-flower smart- weed (Persicaria glabra), and lizard’s-tail (Saururus cernuus). Eastern mosquito fern (Azolla caroliniana), the liverwort Riccia, and water Figure 8-1. Cypress-gum-pine border, Byrnes Lake Boat Landing north of Spanish Fort. (Photo credit: Lawrence Davenport.)

48 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area spangles (Salvinia minima) float on the water’s surface.

3) Grass Beds In another section of his introduction to Plant Life of Alabama, Mohr (1901) described the “limnaean” vegetation of the Delta:

The still waters of the estuary of [the] Mobile River and of the larger streams emptying into the upper part of the bay, fresh, except at long intervals, when it is slightly brackish, harbor a number of submerged species, forming in their Figure 8-2. Hardwood pine border, Byrnes Lake Boat Landing north of Spanish Fort. (Photo credit: Lawrence Davenport.) dense mass subaquatic meadows….

He listed the following species as constituting these “meadows” or grass beds: horned pondweed (Zannichellia pal ustris), wigeon-grass (Ruppia maritima), coon’s-tail (Ceratophyllum demersum), curly pond-weed (Potamoge- ton crispus), small pondweed (Potamogeton pusillus ssp. pusillus), water-thread pondweed (Potamogeton diversi- folius), long-leaf pondweed (Potamogeton nodosus), clasp- ing-leaf pondweed (Potamo- geton perfoliatus), Eurasian water milfoil (Myriophyl- lum spicatum), leafy blad- Figure 8-3. Marsh with Mobile skyline, Five Rivers Delta Resource Center, Spanish derwort (Utricularia foliosa), Fort. (Photo credit: Lawrence Davenport.) long-beak buttercup (Ranun- culus longirostris), grassleaf arrowhead (Sagittaria 4) Open River Marshes graminea), American lotus, American eelgrass (Vallis- In his portrait of the Delta, Mohr (1901) also painted neria americana), and Canadian waterweed (Elodea its open river marshes (Figures 8-3, 8-4, 8-5): canadensis). To Mohr’s list I would add sago pondweed (Stukenia pectinata) and turtle grass (Thalassia The islands in the lower part of the delta and the testudinum), while emphasizing the dominance of low banks of the streams in the tidewater region American lotus in some areas. See Raines (2015) for a are covered with deep and extensive open recent description of the latter. marshes, the soft silt of which, rich in humus,

Biotic Landscape - Chapter 8 49 Figure 8-4. Marsh and cypress-gum edge, Historic Blake- Figure 8-5. Boardwalk along Tensaw River marsh edge, ley State Park, Spanish Fort. (Photo credit: Lawrence Historic Blakeley State Park, Spanish Fort. (Photo credit: Davenport.) Lawrence Davenport.)

supports an association of paludial plants (halo- Caribbean-based Southern cattail (Typha domingen- phytes), which take root firmly in the soil, their sis), which is now well established in certain Delta mostly strong, interlacing rhizomes forming marshes. a dense sod, resisting the action of waves and winds. Reed-like grasses, large rushes, and tall Mohr further noted basic successional trends in these umbelliferous plants are the most conspicuous marshes, with wild rice, soft-stem bulrush, and broad- features of this association. leaf cattail being “among the first [plant species] to gain a firm hold on the muddy shoals, constantly formed Here he included common reed (Phragmites austra- by the deposits of silt with which the turbid waters lis), salt-meadow cordgrass (Spartina patens), big are charged.” He also added the following grami- cordgrass (Spartina cynosuroides), annual wild rice noids to his growing list: red-root flatsedge (Cype- (Zizania aquatica), water millet (Zizaniopsis miliacea), rus erythrorhizos), straw-color flatsedge (Cyperus switchgrass (Panicum virgatum), maiden-cane (Pani- strigosus), rusty flatsedge Cyperus( odoratus), sallow cum hemitomon), broadleaf cattail (Typha latifolia), sedge (Carex lurida), green-white sedge (Carex albo- soft-stem bulrush (Schoenoplectus tabernaemontani), lutescens), broad-wing sedge (Carex alata), Southern river bulrush (Bulboschoenus fluviatilis), saw grass cutgrass (Leersia hexandra), purple bluestem (Andro- (Cladium jamaicense), jointed flatsedge Cyperus( pogon glaucopsis), angle-stem beaksedge (Rhyncho- articulatus), spotted water hemlock (Cicuta macu- spora caduca), and short-bristle horned beaksedge lata), stiff cowbane (Oxypolis rigidior), and water (Rhynchospora corniculata). Plus he added the follow- parsnip (Sium suave). To Mohr’s list I would add the ing more broad-leaved herbs and shrubs: saltmarsh

50 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area morning glory (Ipomoea sagittata), broadleaf Indian- plantain (Arnoglossum ovatum), Virginia saltmarsh- mallow (Kosteletskya virginica; Figure 8-6), small- head doll’s-daisy (Boltonia diffusa), giant ironweed (Vernonia gigantea ssp. giantea), bull-tongue arrowhead (Sagittaria lancifolia ssp. media), giant arrowhead (Sagittaria montevidensis), saltmarsh loosestrife (Lythrum lineare), pickerel-weed (Pontederia cordata; Figure 8-7), pale dock (Rumex altissimus), creeping burhead (Echinodorus cordifolius), broadleaf arrowhead (Sagittaria latifolia), and Southern seaside gold- enrod (Solidago sempervirens var. mexicana).

Mohr (1901) ended his lengthy discussion of the Delta’s open river marshes by noting, “The dark waters of ditches and shallow pools at the outskirts of these marshes are filled with the floating stems of” floating primrose-willow Ludwigia( peploides spp. glabrescens) “and bordered by” floating marsh- pennywort (Hydrocotyle ranunculoides), whorled marsh-pennywort (Hydrocotyle verticillata), cursed buttercup (Ranunculus sceleratus), threadleaf mock bishopweed (Ptilimnium capillaceum), and sticky joint-vetch (Aeschynomene viscidula). Other plants noted were marsh spikerush (Eleocharis palustris), three-rib arrowgrass (Triglochin striata), needle-pod rush (Juncus scirpoides), many-head rush (Juncus polycephalus), hairy-pod cowpea (Vigna luteola), and climbing hempweed (Mikania scandens). Pea-tree Figure 8-7. Pickerel-weed, Pontederia cordata, Bayfront (Sesbania herbacea) and bagpod (Sesbania vesicaria) Park, Daphne. (Photo credit: Lawrence Davenport.)

“occupy almost alone the alluvial banks bordering the swamps, covered with the debris left behind after every overflow.”

5) Dry Edges and Beaches Mohr (1901) also described the littoral zone, extending along the eastern shore of Mobile Bay to the beaches of the Fort Morgan Peninsula and Dauphin Island. Surpris- ingly, he offered few details of the Bay and its dry edges, except for the sand-stabilizing presence of Brazilian bay- hops (Ipomoea pes-caprae Figure 8-6. Virginia salt-marsh mallow, Kosteletskya virginica, Bayfront Park, ssp. brasiliensis). Other vines Daphne. (Photo credit: Lawrence Davenport.) and low-lying shrubs are

Biotic Landscape - Chapter 8 51 death—which came, sadly, just 2 weeks before the publication of Plant Life of Alabama— Roland M. Harper was named botanist for the Geological Survey of Alabama. During his long (1905-1966) and fruitful association with the Survey, Harper published over 600 letters, scientific articles, and books. (For a synopsis and analysis of Harper’s works, see Davenport and Hubbs 1995). While several of these (Harper 1913, 1920, 1943) deal with Alabama’s forests and plant associations, none offers the details provided by Mohr. Figure 8-8. Beach morning-glory, Ipomoea imperati, on dry sand spit, Village Point Park Preserve, Daphne. (Photo credit: Lawrence Davenport.) Harper’s first treatment (Harper 1913) is admittedly trumpet-creeper (Bignonia capreolata), beach morning superficial or “broad brush,” based on 3 train rides glory (Ipomoea imperati; Figure 8-8), marsh false along the Louisville & Nashville Railroad across the bindweed (Calystegia sepium ssp. limnophila), rough lower end of the Delta. He listed only the main tree cocklebur (Xanthium spinosa), common partridge species seen from the train car windows: bald cypress, pea (Chamaecrista fasciculata), and dwarfed Chinese black willow, cottonwood, American elm, sweetbay, Tallowtrees (Triadica sebifera). red bay (Persea pubescens), sweetgum, red maple, black gum, tupelo gum, and green ash.

6) Shell Hammocks Harper’s second treatment (Harper 1920), based on Finally, Mohr (1901) noted shell hammocks near the the same railroad trips, contains a few more details, sea and along the Delta’s “narrow tortuous marine especially among the shrubs, vines, and herbs. His channels.” These “heaps of bivalve shells, frequently third treatment (Harper 1943), again devoted to trees, many yards in length and from 6 to 15 feet [1.8 to includes details from a 1927 trip from Mount Vernon, 4.6 m] and over in height,” are “the accumulation of in north Mobile County, near the Delta’s upper end. refuse from the food supply which served a race of Here he divides those trees into larger and smaller, men unknown to history.” Dominant plants on Delta upper and lower, corresponding largely with Mohr’s shell mounds are live oak (Quercus virginiana), South- designations of wet hardwood sites and “cypress ern magnolia (Magnolia grandiflora), pignut hickory brakes.” (Carya glabra), Carolina buckthorn (Frangula caroliniana), red buckeye (Aesculus pavia), and Carolina E. Lucy Braun, the dominant plant ecologist of her wolfberry (Lycium carolinianum). Growth of the last generation, used even broader strokes. In her master- 2 species may have been encouraged by Mohr’s “race piece, Deciduous Forests of the Eastern United States, of men unknown to history,” with the first being used Braun (1950) made no specific reference to the Delta, as a fish poison and the second one for food (Raines even though she relied heavily on the works of Mohr 2014). and Harper in describing the forest vegetation of the Southeast. The same is true of Christensen (1988); Later Studies while providing detailed accounts of the vegetation zones of the Southeastern Coastal Plain, he used no Later workers have not improved on the work of examples from the Mobile-Tensaw Delta. And in the Mohr, especially in his detailed knowledge of Delta most recent treatment, Sharitz and Mitsch (1993) pull plant associations and dynamics. Four years after his

52 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area details and plant lists from many solid sources. But again, they make no specific reference to the Delta.

The Delta Today Today, the Delta remains an unbroken cypress-tupelo swamp, as described so enthusiastically by Bartram. And its edge habitats and marshes still support the plant communities depicted by Mohr. But a number of changes must be noted.

Although some champions remain in isolated places, the huge trees celebrated by Bartram are gone, logged out during the past 150 years. The same is true of the equally celebrated cane brakes. While cane itself remains in the Southeast, the number and size of individuals is much diminished, as is the importance of cane brakes as a distinct ecosystem (Platt and Brantley 1997).

The completion of the Causeway across the lower Delta in 1926, cutting east-west across the north- south flow of water, has modified salinity regimes, such that some vegetation zones have been enhanced and others diminished. Additionally, many grass beds along Mobile Bay’s eastern shore have suffered heavy silt loads from construction projects (Raines 2014).

Access The vegetation zones of today’s Delta can be read- Figure 8-9. Water’s edge, Byrnes Lake Boat Landing north ily visited by water and land. Public and private boat of Spanish Fort. (Photo credit: Lawrence Davenport.) landings allow access to both the eastern and western portions, with the Five Rivers Delta Resource Center (near Spanish Fort) in the middle (Figure 8-9). Also highly recommended are the boardwalk/trail from Bayfront Park in Daphne (Figures 8-10, 8-11, 8-12) and the E. O. Wilson Boardwalk at Historic Blake- ley State Park (Figure 8-13).

Figure 8-10. The Jackson Oak, Village Point Park Preserve, Daphne. (Photo credit: Lawrence Davenport.)

Biotic Landscape - Chapter 8 53 Figure 8-11. Warning sign, Village Point Park Preserve, Daphne. (Photo credit: Lawrence Davenport.)

Figure 8-12. American alligator, Alligator mississippiensis, viewed from boardwalk, Bayfront Park, Daphne. (Photo credit: Lawrence Davenport.)

54 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figure 8-13. E. O. Wilson Boardwalk, Historic Blakeley State Park, Spanish Fort. (Photo credit: Lawrence Davenport.)

Biotic Landscape - Chapter 8 55

9. Exploring the Unknown—The Insects of the Mobile-Tensaw River Delta John W. McCreadie and Peter H. Adler, University of South Alabama

Who will inherit the earth? In many ways, evolution introduce the insect biodiversity of the Delta, we has answered this question: the insects. In a world provide preliminary accounts of a small selection of rich with life, 60% of all known species are insects, insect groups. currently numbering more than 1 million (Adler and Foottit 2009). The reigning kings of biodiversity, What Is Known of Insects in the beetles, account for well over one third of all insect Delta? species. Even within the beetles, a single group, the Little has been published on the insects of the Delta. weevils, includes 3 to 4 times as many species as all This is both exciting and daunting, since almost every- species of birds and mammals combined. thing we learn about the insects in the Delta is new. The Mobile-Tensaw River Delta (hereafter, as the Apart from a US Department of Agriculture (USDA) locals call it, “the Delta”) is a relatively pristine, roughly report on the control of tent caterpillars in the 1970s 1000 km2 (390 mi2) area of subtropical wetlands about (Abrahamson et al. 1982), the first paper published on 60 km (37 mi) long and 16 km (10 mi) wide, stretching insects in the Delta concerned a curious insect known from the confluence of the Alabama and Tombigbee as a water scorpion (Figure 9-1; Ihle and McCreadie rivers south to the head of Mobile Bay. The Delta is 2003). This stealth predator hides in the vegetation one of the nation’s largest river deltas. A staggering of calm waters, waiting for prey to wander by. Water 15% of the contiguous nation’s water runs through scorpions with barnacles attached to them attest to it. Much of the area remains in a natural state, which the intermingling of freshwater and saltwater near the is remarkable given that the Southeast has relatively little land under federal or state protection (Loomis and Echo- hawk 1999).

The Delta is suspected of being a hotbed of insect biodiversity— “suspected” because we actually know little about the insects of the Delta. Consisting of bottomland hardwood forests interlaced with large rivers, streams, canals, bayous, and marshes, the Delta is challenging to traverse; movement often is possible only by boat. Such an inaccessible habitat has protected its denizens and shrouded many of the smaller creatures, such as the insects, in mystery.

In our investigations of the insects of the Delta, we initiated an inventory and established an arthropod museum to house reference specimens. To Figure 9-1. The water scorpion Ranatra with a barnacle attached to its thorax. (Photo credit: Laura T. Miller, West Virginia Deptartment of Agriculture.)

Biotic Landscape - Chapter 9 57 southern border of the Delta. We also have published eventually examine the effects of landscape features a study on larval dragonflies (McCreadie et al. 2005), on insect biodiversity and community structure. Each several new distribution records, and a couple of new site had 1 Malaise trap initially (for incidence data), and species descriptions. No doubt a number of insects later 4 traps (25 m [82 ft] apart) to permit estimations collected from the Delta are spread across collections of relative abundance of each species per site. This and museums in North America. However, museum arrangement prevented complete loss of data from a specimens typically are encountered only by system- site if individual traps were destroyed by storms. We atists working on specific taxonomic groups, and the also established 20 sites for sampling aquatic insects information is not easily tracked by nonspecialists. along a north-south axis across the Delta, each in a Hence, we are in a better position to discuss what we habitat with unique aquatic vegetation (e.g., submer- do not know about the insects of the Delta than what gent vegetation, debris) and microhabitat (e.g., slow we do know. lotic, fast lotic, bayou). At each site, five 5-min bank- side samples were obtained using a mesh net (Ihle and The Total Insect Bio-inventory Project McCreadie 2003; McCreadie et al. 2005). of the Mobile-Tensaw Delta Search methods also include ad hoc examinations of Nearly all that is known about the insects of the Delta specific habitats, such as animal burrows, which rely has been gathered under the umbrella of the Total on an investigator’s knowledge of the group of inter- Insect Bio-inventory Project of the Mobile-Tensaw est. Although these methods typically lack repeatable Delta. This project was initiated and developed by J.W. selection protocols, and data are not easily quanti- McCreadie and P.H. Adler in 2000, largely with funds fiable, they can add significantly to the number of provided by the Alabama Center for Estuarine Stud- species discovered. ies. The Total Insect Bio-inventory Project’s primary long-term goal is to produce a robust inventory of To process collected material, personnel at the Univer- the insect fauna in the Mobile-Tensaw Delta over the sity of South Alabama place fully labeled samples course of a 20- to 30-year survey. In our endeavor to in vials of 80% ethanol and send them to Clem- catalog as many species as possible, we have enlisted son University, where material is sorted to family by the help of more than 60 collaborators from around students with appropriate taxonomic training. Insects the world. Because the Southeast has less protected such as calyptrate flies are dried chemically with land than most other areas in the United States and is facing the potential battering effects of climate change, the need for a bio-inventory of the relatively pristine Delta wetlands is particularly urgent.

Sampling and Collecting Our interests cover the insects of both aquatic and terrestrial environments. Accordingly, collecting for the Total Insect Bio-inventory Project has taken two basic forms: Malaise traps (Figure 9-2) for terrestrial insects and large hand-held nets for aquatic species.

We have found Malaise traps to be an efficient means of collecting insects in the Delta. Four collecting sites in the lower half of the Delta were established in unique topographical or vegetational areas to a) maximize the Figure 9-2. Malaise trap, the primary means of collecting insects in the number of species collected and b) Delta. (Photo credit: John W. McCreadie, University of South Alabama.)

58 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area hexamethyldisilazane (Brown 1993), which also preserves DNA (deoxyribo- nucleic acid; Koch et al. 1998), should contemporary or future workers wish to perform molecular analyses. Dried specimens are pinned appropriately and a collection label affixed to the pin. Other groups of insects, such as Doli- chopodidae, Psychodidae, Simuliidae, and Tipulidae, are retained in ethanol vials. To accommodate representative specimens collected in the Delta, the University of South Alabama Arthro- pod Depository was established in 2001.

Insects Most of the insects identified from the Delta reflect our own interests, such as the dragonflies (Odonata) and true flies (Diptera); our choice of collect- Figure 9-3. Larva of the blue dasher dragonfly (Pachydiplax longipennis). (Photo credit: John W. McCreadie, University of South Alabama.) ing methods; and, most importantly, the availability of expert taxonomic collaborators. More than 50,000 insects have been collected species—represented by a single speci- sorted to the family level, and thousands await identifi- men—was Nasiaeschna pentacantha (Rambur), the cation by a willing specialist. We have collected groups Cyrano darner. Subsequent to our regimented larval for which there are no active specialists to identify collections, we added 4 more species—Enallagma the specimens. For some groups, therefore, we have pollutum (Hagen), Ischnura kellicotti (Willliamson), long lists of families, genera, and species, but for other Ischnura prognata (Hagen), and Arigomphus pallidus groups we have no identified specimens. Given these (Rambur)—to the list of odonates in the Delta, bring- constraints, insects in 108 families, 366 genera, and ing the total to 20 species. more than 800 species have been identified, entered into a database, and curated in the University of South 2) Diptera—Flies Alabama Arthropod Depository. The true flies of the order Diptera are a highly diverse group. They represent about 10% of the planet’s 1) Odonata—Damselflies and Dragonflies biodiversity, making these small and often overlooked One charismatic order of insects is the Odonata, which insects one of the most species-rich groups of organ- includes nearly 5700 species in the world, over 460 isms in the world (Brown 2005; Adler and Foottit 2009; in North America, and 170 in Alabama. The aquatic Courtney et al. 2009). In the Delta, we have collected larvae (Figure 9-3) and terrestrial adults of all species more than 185 genera and 400 species of flies, at least are highly specialized predators. Monthly samples 6% of which are new species. Many additional species taken for 1 year from Delta streams and rivers more are state records for Alabama and often represent the than 10 m (33 ft) wide yielded 1151 larvae in 4 families, northernmost or southernmost distribution records. with 16 species, or 9.2% of all odonate species known Thus, the Delta appears to be a “hot spot” for fly from Alabama (McCreadie et al. 2005). The number biodiversity. We feature 10 families of flies to provide of species decreased with proximity to Mobile Bay, an indication of the extent of the Delta’s biodiversity. possibly because of increasing salinity. The most common species was Erythemis simplicicollis (Say), the eastern pondhawk, representing 34% of all collected specimens, followed by Enallagma signata (Hagen), the orange bluet, at 21%. The most infrequently

Biotic Landscape - Chapter 9 59 Anthomyiidae (Figure 9-4) The nearly 2000 anthomyiid flies of the world are diverse in their habits. Some species, the so-called “root maggots,” develop in roots and stems, whereas others live in flowers and leaves, some feed on kelp along the seashore, some are fungivores, and still others are scavengers, parasites, and commensals in the nests and burrows of animals. Thus far, 21 spec- imans have been submitted for identification. Eleven genera and 12 species have been collected, among which are 1 new species and 11 state records, including the southernmost record for Eustalomyia vittipes (Zetterstedt), a species also found in the Old World.

Another anthomyiid in the Delta is Figure 9-4. A species of the genus Anthomyia (family Anthomyiidae). (Photo Delia platura (Meigen), the seedcorn credit: Steve A. Marshall, University of Guelph.) maggot. As a seed feeder, this cosmo- politan species can be a sporadic pest of agricultural crops, such as corn and soybeans.

Calliphoridae (Figure 9-5) The blow flies, best known as ubiquitous colonizers of carrion and excre- ment, also include parasites of birds, mammals, and earthworms. They number more than 1520 species in the world. The conspicuous, shiny blue and green flies that appear almost immediately around dead animals have great value as forensic indicators, providing insights into time since death. Five species were found among 48 specimens in our Delta collections, of which 73% were Lucilia coerulei- viridis (Macquart), the green bottle fly. The sex ratio of green bottle flies in our samples was strongly skewed: 33 Figure 9-5. Green bottle flies (family Calliphoridae) depositing eggs on a females to 2 males. dead snake. (Photo credit: Steve A. Marshall, University of Guelph.)

60 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Ceratopogonidae (Figure 9-6) Biting midges, no-see-ums, punkies, wings-with-teeth — the numerous vernacular names for these blood- sucking flies allude to their tiny size and the annoying biting habits of the female flies, primarily in the genus Culicoides, which attack humans. The family contains an enormous number of species, with about 6000 described and many more remaining to be discovered. The females of the majority of species feed on insect blood. To date, 53 species in 15 genera have been collected in the Delta, including 3 new species and 2 state records. Another 3 genera and 33 species have been collected from Baldwin and Mobile counties adjacent to the Delta. One new species, Brachypogon laneae, a member of a group that feeds on the Figure 9-6. A biting midge of the genus Palpomyia (family Ceratopogoni- blood of other flies, already has been dae) that preys on other insects. (Photo credit: Steve A. Marshall, University of Guelph.) described from the Delta (Swanson and Grogan 2011).

Chironomidae (Figure 9-7) The nonbiting midges are among the most common and ubiquitous flies in the world, totaling more than 7200 described species. The larvae develop in aquatic habitats, such as streams, rivers, ponds, and lakes, and in organi- cally rich terrestrial habitats, such as soil. Most species feed on detritus and small organisms, but some are predaceous. Chironomids are frequently used in water quality assessments. Some species are vulnerable to degraded aquatic environments, but others are remarkably tolerant, including the bloodworms—species with hemoglobin—that can achieve enormous populations in poorly oxygen- ated habitats. The adults are small, Figure 9-7. A male nonbiting midge (family Chironomidae). (Photo credit: delicate, mosquito-like flies often Steve A. Marshall, University of Guelph.) found dead in light fixtures. They take water and sugary substances, such as insect honeydew. A meager sample of chironomids from the Delta yielded 46 species, of which at least 13% are new to science and 46% are state records.

Biotic Landscape - Chapter 9 61 The Chironomidae represent one of the richest dipteran families in the Delta, and continued identification will drive the species list far higher.

Dolichopodidae (Figure 9-8) Members of the family Dolichopodi- dae—long-legged flies—are typically metallic with long legs and almost comically oversized male genita- lia. Most adults are predaceous and usually inhabit humid terrestrial environments. Larvae are found in a wide range of habitats, such as tree holes, sap flows, streams, and plant stems. The family is quite large, with more than 7300 species known in the world. Our collections of Dolichopodidae from the Delta have yielded more than 90 species, of which 14 are new Figure 9-8. A typical metallic long-legged fly (family Dolichopodidae). to science. Only 9 species of long- (Photo credit: Steve A. Marshall, University of Guelph.) legged flies were known in the state of Alabama before 2000 (Pollet et al. 2004).

Muscidae (Figure 9-9) The family Muscidae is far more diverse than suggested by its two most notorious synanthropic members, the house fly (Musca domestica L.) and the stable fly (Stomoxys calcitrans L.). The world’s 5200-plus species represent a wide variety of larval habitats, including carrion, decaying vegetation, soil, bird nests, and freshwater. Adults of some species have evolved piercing- cutting mouthparts to take vertebrate blood. Among a sample of 97 adults collected in Malaise traps in the Delta, 10 genera and 17 species were identified, including 2 probable new species. The most frequently collected muscid, representing one quarter of the iden- Figure 9-9. Flies in the same family as the house fly (family Muscidae). tified specimens, was the widespread (Photo credit: Steve A. Marshall, University of Guelph.) Coenosia lata (Walker), followed by Neodexiopsis major (Malloch; 14%).

62 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Psychodidae (Figure 9-10) The tiny moth flies are familiar to most people as denizens of bathrooms and communal showers where they breed in accumulated organic matter in drains. The annoying bathroom species belies a far richer taxonomic and ecological diversity of moth flies associated with a variety of wet habitats. More than 3000 species have been described for the world. Adults are largely nocturnal. Collections in the Delta have yielded 18 species in 10 genera, including 2 new species and 2 state records. More than 20 additional species have been found in adjacent areas of Baldwin and Mobile counties, suggesting the aquatic environment of the Delta supports a substantial psychodid fauna. Figure 9-10. A moth fly (family Psychodidae). (Photo credit: Steve A. Mar- Simuliidae (Figure 9-11) shall, University of Guelph.) The pesky gnats known as black flies or buffalo gnats are small, compact flies that conjure the image of lillipu- tian American bison. The females of about 90% of North American species take a blood meal; only birds and mammals serve as hosts. Black flies breed exclusively in flowing freshwater and are best represented in moun- tainous areas, where the elevational gradient offers a variety of riverine habitats. Given the elevational monot- ony of the Delta, we would expect few species. Our Malaise trap samples bear this out, yielding only 3 species (about 1%) of the 255 simuliid species known from North America (Adler et al. 2004). One species feeds on the blood of mammals, and the other 2, including the notorious turkey gnat, Simulium meridionale (Riley), take the blood of birds. Remarkably, one Figure 9-11. A black fly (family Simuliidae) that feeds on the blood of mammals. (Photo credit: Steve A. Marshall, University of Guelph.) of these species is new to science—an early season, Coastal Plain endemic that was chromosomally investigated by one of our graduate students (Glea- son 2012). An additional 5 species

Biotic Landscape - Chapter 9 63 have been collected as larvae from streams and rivers flowing into the Delta in Baldwin and Mobile counties.

Tachinidae (Figure 9-12) Flies in the family Tachinidae constitute the second most species-rich group of Diptera, approaching 10,000 described species in the world and more than 1340 in North America. The larvae of all species of tachinids are parasitoids of insects, with scattered examples of species attacking a spider, centipedes, and scorpions. A preliminary sorting of our Malaise trap collections revealed 19 genera, 29 species, and 7 state records, including the northernmost record for Ormia dominicana (Townsend). This pale brownish-orange tachinid is a para- Figure 9-12. A member of the large family of parasitoid flies (family Tachini- sitoid of Orocharis luteolira (Walker), dae). (Photo credit: Steve A. Marshall, University of Guelph.) the false jumping bush cricket; the fly locates its host acoustically based on the mating song (Lehmann 2003).

Tipulidae (Figure 9-13) Delicate, long-legged flies of the family Tipulidae sensu lato—the crane flies—are one of the most species-rich families of insects on Planet Earth (although some specialists recognize the superfamily Tipuloidae and split the Tipulidae into 4 families). More than 1610 species of crane flies are found in North America, representing about 10.5% of the world’s total (de Jong et al. 2008). They are well represented in our Delta samples, with 31 species in 16 genera so far recorded. Our most commonly collected species was Erioptera caliptera (Say), the larvae of which live in damp soil.

Figure 9-13. A typical crane fly (family Tipulidae). (Photo credit: Steve A. Marshall, University of Guelph.)

64 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Summary—More than 800 Species and for the contributions to this effort by the following Counting scientists: D. Ackland (Anthomyiidae), J. A. Cammack With a modest total of 800-plus species recorded thus (Calliphoridae), G. Curler (Psychodidae), J. Epler far, our goal to inventory the insects of the Delta contin- (Chironomidae), W. L. Grogan and D. A. Swanson ues. Lest the ultimate objective be misinterpreted, we (Ceratopogonidae), R. Hurley (deceased) and J. B. emphasize that the inventory is not an endpoint, but Runyon (Dolichopodidae), A. Pont (Muscidae), D. M. rather a starting point for understanding, protecting, Wood (Tachinidae), and C. W. Young (Tipulidae). We conserving, managing, and even harvesting the riches are indebted to S. A. Marshall for generously allowing of the Delta’s insect fauna. us to use his photographs of representative flies. We thank E. Benton and W. K. Reeves for their dedication Acknowledgments to sorting material to family level and coordinating identifications with many of the specialists. Large surveys of organisms, especially of arthropods, are possible only because of the specialized knowledge and enthusiasm of specialists. We are grateful

Biotic Landscape - Chapter 9 65

10. Vector-Borne Disease Systems Nathan Burkett-Cadena, Florida Medical Entomology Laboratory, University of Florida

Vector-borne diseases are afflictions caused by a the idea that mosquitoes could carry yellow fever, pathogen carried from one host to the next by another or other diseases, was not fathomed by the medical living organism. The agent of disease, often called the community, much less the average person. Not until pathogen, is adapted to thrive not only within the body more than 100 years later did scientists and physicians of its host, but also within the body of its carrier, called begin to understand the association between mosqui- the vector. The vector acquires the pathogen from the toes and human disease, and then only imperfectly. host, usually through the act of biting and blood feed- Like malaria, yellow fever was thought to be caused ing. The term “Vector-borne disease system” refers by “bad air” that emanated from rotting vegetation to all of the elements (both living and non-living) abounding in marshes and swamps. that influence the cycle of transmission of a vector- borne disease. It most often includes the pathogen, its Josiah Nott, a prominent physician in Mobile was one vectors, various hosts, the habitat where transmission of the first to notice how the spread of yellow fever occurs, and the weather and climate that influence the in a community coincided with the mosquito season habitat and biologies of hosts and vectors. (Nott 1848). Dr. Nott likened the spread of the disease to the behavior of the mosquito moving through the Vector-borne diseases can be caused by a wide vari- landscape. While Nott’s writings fell short of stating ety of organisms. Malaria, for example, is caused by that mosquitoes actually carry a pathogen, the philo- a protozoan (animal-like single-celled organism), sophical link was established between mosquitoes Lyme disease by a bacterium, yellow fever by a virus, and disease. and dog heartworm by a roundworm. Organisms that transmit vector-borne diseases are as diverse as pathogens and include mosquitoes, ticks, sand flies, kissing bugs, mites, lice, and fleas, among many others (Mullen and Durden 2009). In the greater Mobile- Tensaw River area, the most important vector-borne diseases of humans are viruses transmitted by mosquitoes.

Yellow Fever and the Greater Mobile-Tensaw River Area The greater Mobile-Tensaw River area, particularly the port of Mobile, occupies a special place in the history of vector-borne diseases. Originally located at Twenty-seven Mile Bluff, the city of Mobile was relocated in 1711 to its current location partly due to recurring epidemics of yellow fever. Figure 10-1. Historic marker at Old Church Street Cemetery, Mobile, AL, Unfortunately, the early Mobilians did commemorating the interment of early residents of the greater Mobile- not escape yellow fever by relocating Tensaw River area who fell victim to vector-borne diseases, particularly yellow fever. (Photo credit: Nathan Burkett-Cadena, University of Florida.) their city (Figure 10-1). At that time

Biotic Landscape - Chapter 10 67 This important first step fostered a scientific atmo- Cases of Eastern equine encephalitis occur through- sphere that eventually led to experiments in Havana, out the eastern United States, from Maine to south- Cuba, in 1901 that demonstrated how yellow fever ern Florida, most commonly in low-lying areas where was transmitted by mosquitoes. An attendant of those swamps and swamp-inhabiting mosquitoes are found. experiments was William Crawford Gorgas, another The mosquito species Culiseta melanura is the most prominent Mobilian, who would eventually become important vector of Eastern equine encephalitis Surgeon General of the US Army. After the conclu- virus, although it very rarely bites humans or other sion of the Yellow Fever Commission in Havana, mammals. Culiseta melanura predominantly bites Gorgas was charged with overseeing sanitation during songbirds, such as the familiar northern cardinal. construction of the Panama Canal (1904 to 1914). When Culiseta melanura bites an infected songbird, Having observed first-hand the experiments proving the virus in the bird’s blood passes to the mosquito. that mosquitoes transmit yellow fever, Gorgas used Inside the body of the mosquito the virus must pass this knowledge to implement the first widespread from the digestive tract into the body cavity, and then sanitary programs to protect humans from mosquito- to the salivary glands. During this process the virus borne diseases. By draining swamps, fumigating living encounters multiple transmission barriers that it must quarters, and destroying mosquito breeding sites, be adapted to pass, otherwise the mosquito will not Gorgas’s sanitarians protected laborers construct- transmit the virus when it bites another animal. ing the Panama Canal from yellow fever and malaria, diseases that had thwarted a French attempt to build Although birds are not particularly affected by Eastern the canal 2 decades earlier (Dolan and Silver 1968). equine encephalitis, the virus can be deadly to humans and horses. In symptomatic cases of Eastern equine Vector-borne Diseases and the Greater encephalitis the fatality rate is greater than 50%. Mobile-Tensaw River Area Today Since Culiseta melanura rarely bites humans or other mammals, other mosquito species, such as Coquillet- Today yellow fever and malaria, the scourges of Old tidia perturbans, which bite both birds and mammals, Mobile, are no longer found in the US. An effec- are important as vectors of the virus to humans and tive vaccine for the yellow fever virus was developed domestic animals (Figure 10-2). Species such as in 1937, and a widespread yellow fever vaccination Coquillettidia perturbans, which carry a pathogen program, coupled with the new science of mosquito from the reservoir host to humans or other animals control, led to eradication of yellow fever from the US that are adversely affected by a pathogen, are some- and many other parts of the developed world. Malaria times called “bridge vectors.” Many other mosquito- was also eradicated from the US, although much later borne viruses that affect humans, such as West Nile than yellow fever. No vaccine exists for malaria. Thus virus and St. Louis encephalitis, have transmission the National Malaria Eradication Program, which cycles similar to that of Eastern equine encephalitis, commenced operations in 1947, utilized application with birds as the natural host for the virus. The bird of DDT (dichlorodiphenyltrichloroethane) and the and mosquito species important in transmission draining or removal of mosquito breeding sites to usually differ from one virus to the next. eliminate malaria as a significant public health problem in the continental US by 1949. Although these 2) Tensaw Virus immensely important diseases are no longer a threat in Another mosquito-borne virus found in the greater Alabama, several other vector-borne diseases are still Mobile-Tensaw River area that warrants mention is found in the region, some of which have a profound Tensaw virus, named after the Tensaw River, which impact upon human health. flows through western Baldwin County before reaching Mobile Bay. Tensaw virus was discovered during 1) Eastern Equine Encephalitis extensive mosquito-borne virus investigations in Eastern equine encephalitis—sometimes known as Baldwin County during the 1950s and 1960s by teams “EEE” or “Triple E”—is probably the most important of scientists from the Centers for Disease Control vector-borne disease in the greater Mobile-Tensaw and Prevention (Stamm 1958; Sudia et al. 1968). In River area (Mullen and Hribar 1988). The disease is that era, very little was known about the transmission caused by a virus transmitted by certain mosquito cycles of mosquito-borne diseases. Many of today’s species that thrive in wooded swamps, such as those basic concepts concerning mosquito-borne virus that dominate much of the Mobile-Tensaw Delta.

68 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area equine encephalitis, the scientists discovered, characterized, and named viruses that were previously unknown.

Tensaw virus is not known to cause disease in humans, but it can be lethal in certain rodents (Coleman 1969). Foxes, rabbits, and some rats are probably important hosts of Tensaw virus (Sudia et al. 1968; Bigler et al. 1975), although relatively little is known about the ecology of this virus. The mosquito Anopheles crucians is considered the most important vector of Tensaw virus in the southern US (Calisher et al. 1986). This mosquito is abundant in swampy areas of the Deep South, and is one of the more common species in the greater Mobile-Tensaw River area. Anopheles crucians mainly bites Figure 10-2. Transmission cycle of a mosquito-borne disease: Eastern equine mammals, but will also bite herons and encephalitis. The mosquito species Culiseta melanura and certain wild song- other wading birds. birds, such as the northern cardinal, are important vectors and reservoirs of the virus in swampy habitats. Other mosquitoes, such as Coquillettidia perturbans, pick up the virus from infected birds and transmit to epidemic Tick as Vectors of Human hosts, such as humans and horses. Pathogens Several tick species that are vectors of human pathogens occur in the greater transmission were established during those stud- Mobile-Tensaw River area. These include the Gulf ies in southern Alabama, which were conducted in Coast tick (Amblyomma maculatum), the lone star tick response to outbreaks of Eastern equine encephali- (Amblyomma americanum), and the deer tick (also tis at farms near swampy lands bordering tributaries known as black-legged tick, Ixodes scapularis) (Figure of the Tensaw River. Thousands of mosquitoes were 10-3). These ticks transmit a variety of human diseases, captured, identified, and tested for mosquito-borne including Lyme disease (Borrelia burgdorferi), Rocky viruses. In addition to the virus that causes Eastern Mountain spotted fever (Rickettsia ricketsii), southern

Figure 10-3. Medically important tick species of the greater Mobile-Tensaw River area: A) Amblyomma americanum, lone star tick; B) Amblyomma maculatum, Gulf Coast tick; C) Ixodes scapularis, black-legged tick or deer tick. (Image credits: A) and B) courtesy of James Gathany, Centers for Disease Control and Prevention; C) courtesy of Nathan Burkett-Cadena, University of Florida.)

Biotic Landscape - Chapter 10 69 tick-associated rash illness (Borrelia lonestari), and The next time the tick, now a nymph, bites an animal American boutonneuse fever (Rickettsia parkeri). it has the potential of transmitting the pathogen to its Some of these tick-borne diseases—particularly Lyme new host, which could be a human, dog, deer, or other disease and Rocky Mountain spotted fever—have long animal. been associated with the northern and western US, and doctors in those areas are accustomed to diagnos- Nymphs are only about half the size of the adult tick ing and screening patients for the pathogens. Many (Figure 10-4), so they are more likely to be overlooked physicians in the southern US, however, are unac- during tick checks than adult ticks. Since ticks usually customed to diagnosing tick-borne diseases, so many need to be attached for more than 24 hours before cases of tick-borne diseases, such as Lyme disease, are infecting their host, the duration of tick attachment undiagnosed and unreported in the region (Clark et is an important factor determining risk of transmis- al. 2014). sion. This is one reason that infected nymphal ticks are more dangerous to humans than adult ticks, which Small mammals, especially rodents, are the most are larger and more easily found on the body. important wild reservoir hosts of tick-borne diseases in the US (Mullen and Durden 2009). Rodent species, such as the white-footed mouse, deer mouse, meadow Concluding Thoughts vole, and woodrat, are important hosts for the patho- Some vector-borne diseases such as dengue and gens and important hosts for the tiny tick larvae— West Nile virus are clearly associated with the “peri- often called seed ticks—that transmit the pathogens. domestic” landscape (Mullen and Durden 2009). This Seed ticks that feed on an infected wild animal can is primarily because mosquitoes that transmit these pick up pathogenic organisms from the animal’s diseases, such as Aedes aegypti and Culex quinque- blood. The blood-fed seed tick drops off the host fasciatus, proliferate in urban and suburban environ- animal and sheds its exoskeleton to grow to the next ments. The larvae of these mosquitoes are not found (larger) stage, called the “nymph”, another immature in swamps or ponds, but have adapted to man-made stage in the life cycle of ticks (Figure 10-4). As with objects, such as unkempt birdbaths, discarded tires, mosquito-borne diseases, the pathogenic organism old paint cans, flower pots, and other artifacts that replicates in the body of the blood-fed vector tick. can hold water and fallen leaves or other organic debris that are the food of the mosquito larva. Humans create larval habitat for the vectors of West Nile virus, dengue, and yellow fever, so these diseases are most common in populated areas. Many other serious vector-borne diseases, such as Eastern equine encephalitis and Lyme disease, are transmitted by vector species that do not prosper in populated areas, but instead thrive in wild habitats. As humans encroach upon, invade, and otherwise alter habitats where the vectors of arthropod-borne diseases are most abundant, we increase our risk of exposure to these pathogens (Weaver 2005).

Much research is still needed to fully document and understand the importance of vector-borne diseases in the greater Mobile-Tensaw River area. It is overwhelmingly likely that undiscovered vector-borne diseases are currently circu- lating in the wetlands of the region. Tensaw Figure 10-4. Life cycle of the lone star tick, Amblyomma americanum. virus, for example, was discovered in this

70 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area hot spot of biodiversity, during studies investigating the potential vectors and reservoirs of these patho- other mosquito-borne diseases. Advances in molec- gens are needed to help better understand this unique ular biology and techniques for detection of novel aspect of the greater Mobile-Tensaw River area. pathogens will certainly yield new species of vector- borne pathogens. Studies that focus on the ecology of

Biotic Landscape - Chapter 10 71

11. Freshwater Mollusks of the Greater Mobile- Tensaw River Area Michael M. Gangloff, Appalachian State University

The freshwater mollusk fauna of the greater Mobile- likely historical assemblage structure, identifies the Tensaw River area is among the least frequently key freshwater mollusk populations in this system, collected mollusk assemblages in Alabama and and notes threats to those populations. the southeastern United States. The vast majority of mollusk habitats in this region occur within the Geographical Scope principal rivers. Difficulties involved with sampling Mollusk habitats in the greater Mobile-Tensaw River mollusks from these deep, swift rivers make them area include the Alabama, Mobile, and Tombigbee unforgiving places to conduct mollusk surveys (Figure rivers upstream to the confluences of Big Flat Creek 11-1). Since most mollusk habitats in these large rivers with the Alabama River and Santa Bogue and Satilpa exist at depths that preclude effective hand or mask- creeks with the Tombigbee River. The upstream and-snorkel sampling, much of our knowledge about boundaries of this reach coincide with the locations of this system’s historical mollusk assemblages has been the Claiborne and Coffeeville locks and dams on the gleaned from dead shells collected on gravel bars or Alabama and Tombigbee rivers, respectively (Williams archeological middens. et al. 2008). Major tributaries of the Alabama River During the past 20 years the region has received more in this reach include Big Flat, Halls, Holly, Lime- attention from freshwater biologists employing scuba stone, Majors, Pigeon, Pine Log, Randons, Reedy, and to survey mussel and snail populations in the deeper Wallers creeks and Little River (Figure 11-2). Bassett, reaches of the lower Alabama, Mobile, and lower Bilbo, Jackson, Leatherleaf, Lewis, Limestone, Salt, Tombigbee rivers, and from surveys of tributaries in Santa Bogue, Satilpa, and Tauler creeks are major trib- Baldwin, Clarke, Mobile, Monroe, and Washington utaries to the Tombigbee River (Figures 11-3 through counties. Recent work has produced a more complete, 11-6). The Alabama and Tombigbee rivers join east if largely qualitative, picture of this region’s freshwater of Calvert, Alabama to form the Mobile River. The mollusk resources. This chapter describes the region’s Mobile River is characterized by a wide floodplain interwoven with distributary or slough channels (Williams et al. 2008). Named distributary channels of the Mobile River include the Middle and Tensaw rivers.

Historical Surveys Early freshwater mollusk biologists working in Alabama were dissuaded from working in much of the greater Mobile-Tensaw River area by the remote and inaccessible nature of its rivers. Much of what we know of the region’s mollusk fauna prior to the Figure 11-1. Alabama River main channel habitat looking upstream from Claiborne 1950s comes from specimens Landing, Monroe County, Alabama. (Photo credit: Michael M. Gangloff, Appala- recovered from ancient Native chian State University.)

Biotic Landscape - Chapter 11 73 American middens, as well as from material collected by early workers including H. H. Smith and R. L. Howard from small streams (McGregor and Dumas 2010; Peacock et al. 2014; see Author’s Note). Archeological excavations near the salt springs in southern Clarke County, a historically important natural and economic resource from prehistoric times until fairly recently, revealed a long-standing Native American presence in the region. McGregor Figure 11-2. Little River headwaters, Claude D. Kelly State Park. This stream forms and Dumas (2010) docu- the southern boundary of Monroe County and drains portions of the Citronelle mented a species-rich histori- Formation. Its tannin-stained waters have pH levels apparently too low to allow for shell formation by mollusks. No bivalves or gastropods are known from this large cal freshwater mussel fauna tributary of the lower Alabama River. (Photo credit: Michael M. Gangloff, Appala- (19 species, including 4 taxa chian State University.) currently listed as threatened or endangered by the US Fish and Wildlife Service) in the lower Tombigbee River, near the confluence of Salt Creek in southern Clarke County. Inter- estingly, the majority of individuals in middens appear to be species that are numerically dominant in recent surveys (McGregor and Garner 2006; Peacock et al. 2014).

Between 1964 and 1976, Royal Sutkiss at Tulane University collected mussels from sites across the Alabama River drainage. These collections include material from 11 sites primarily in the lower Alabama River, as well as from the Tensaw River, the Big Flat and Bear Creek tributaries, and Figure 11-3. Bassett Creek near Jackson, Alabama is a fast-moving, gravel-bottomed tributary to the Tombigbee River. Surveys in 2011 revealed populations of at least one oxbow lake (see 7 freshwater mussel taxa near the Clarke County Road 15 bridge crossing. This Author’s Note). Other collec- large but poorly surveyed stream likely supports populations of other unionid tions from this time period taxa. (Photo credit: Michael M. Gangloff, Appalachian State University.) are sporadic, but together with archeological material provide a glimpse of the region’s pre- impoundment mollusk fauna.

74 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Recent Surveys The era of recent mollusk surveys in the lower Tombigbee and Alabama river drainages began in the 1980s with review of numerous freshwater mussel and snail taxa for protection under the Endangered Species Act. In 1989 the US Fish and Wildlife Service added the Heavy Pigtoe (Pleurobema taitianum), Southern Combshell (Epioblasma penita), and Stirrupshell (Quadrula stapes) mussels to the Endangered Species List (USFWS 1989). In 1990 the USFWS listed the Inflated Heelsplitter mussel (Pota- milus inflatus) as threatened, and in 1991 the recently rediscovered Tulo- toma Snail (Tulotoma magnifica) was listed as endangered (USFWS 1990, 1991, 1993). These actions neces- Figure 11-4. Leatherwood Creek is more typical of low-gradient swampy sitated additional surveys for these channels in the Salt Springs region of Clarke County. This stream supports populations of Toxolasma texasensis, Pyganodon grandis, Glebula rotundata, taxa across their historical ranges in and Cincinnatia integra, a hydrobiid snail. (Photo credit: Michael M. Gangl- the Mobile Basin (Pierson 1991). off, Appalachian State University.) Initially, mollusk surveys in the lower Alabama River were conducted by USFWS and Alabama Depart- ment of Conservation and Natural Resources (ADCNR) personnel. But survey needs quickly exceeded existing personnel and fiscal resources, so biologists with other agencies, universities, utilities, and private stakeholders joined the effort. Following the rediscovery in 1982 of Tulotoma magnifica in the Coosa River near Wetumpka, Alabama Power Company and USFWS began surveys in shoal habitats in the Alabama, Cahaba, and Coosa rivers to assess remaining populations. These surveys later revealed that the distribution of Tulotoma magnifica populations extended downstream to the tailwaters of Claiborne Lock Figure 11-5. Limestone Creek on the Fred Stimpson Sanctuary is characterized by a highly incised and high-gradient, third-order channel with relatively and Dam (USFWS 2011). Although coarse gravel substrates. Elimia taitiana are extremely abundant in this no populations of Tulotoma Snail stream, but appear to be the only mollusks present. (Photo credit: Michael were found in the lower Alabama M. Gangloff, Appalachian State University.) River downstream of Claiborne, these and other surveys during the 1980s and 1990s provide additional

Biotic Landscape - Chapter 11 75 are highly constrained and predict- able. Water chemistry parameters, including dissolved oxygen, pH, and concentrations of Ca2+ and other cations, influence the shell building and waste excreting abilities of freshwater mollusks. Low pH and cation concentrations apparently exclude most native freshwater mollusks from blackwater streams draining poorly buffered strata (e.g., Citro- nelle Formation) in the southeastern quadrant of the greater Mobile- Tensaw River area. In contrast, larger streams draining portions of the Coastal Plain in northern Clarke and Monroe counties intersect limestone outcroppings, and are consequently well-buffered and hence more condu- Figure 11-6. Floodplain wetland or backwater habitat adjacent to the main cive to shell growth. Mussel assem- stem of the Tombigbee River on Fred Stimpson Sanctuary, Clarke County, blages typical of well-buffered Gulf Alabama. Such organic-matter rich and oxygen poor habitats may form Coastal tributary streams commonly when river meanders become cut off during natural channel migration or as include thin-shelled taxa (subfamily a result of anthropogenic channel manipulations. Typical mollusks occurring in these types of habitats include the freshwater mussels Anodonta subor- Lampsilinae) in the genera Lampsilis biculata, Glebula rotundata, Plectomerus dombeyanus, Pyganodon grandis, and Villosa spp. fingernail clams, and air-breathing snails. (Photo credit: Michael M. Gangloff, Appalachian State University.) Although unionid assemblages in large river systems in the Mobile

Basin may share numerous species with tributary data about freshwater mollusk assemblages in this assemblages, heavy-shelled taxa (subfamily Amblemi- reach (Pierson 1991; see Author’s Note). nae) are far more abundant in faster-water main During the last 25 years, freshwater mollusk surveys channel habitats. Main channel habitats are domi- in the region have targeted tributaries of the lower nated by species like Elliptio crassidens, Plectomerus Alabama and Tombigbee drainages (McGregor et al. dombeyanus, Quadrula asperata, and Regenaia ebena. 1999), and the lower main stem Alabama, Mobile, and In contrast, thinner-shelled taxa (subfamily Anodon- Tombigbee rivers (Hartfield and Garner 1998; Miller tinae) are generally the primary taxa found in off- 2000; McGregor and Garner 2006). Recent focused channel slough and oxbow habitats (although the mussel surveys have examined portions of Big Flat thick-shelled taxa P. dombeyanus and Glebula rotun- (Gangloff et al. 2009) and Bassett creeks (Gangloff data may also occur in these habitats) (Williams et al. 2011), as well as streams on the Stimpson Wildlife 2008; Haag 2012). Sanctuary in Clarke County, administered by Alabama Integrating results of recent surveys and studies of Division of State Lands (Gangloff et al., unpublished prehistoric middens reveals that as many as 74 species data, unreferenced). These surveys were conducted of freshwater pearly mussels (Bivalvia: Unioniformes: in association with research on small dams (Gangloff Unionidae and Margaritiferidae), clams (Sphaeriidae), et al. 2009, 2011), infrastructure maintenance (Gangl- and gastropods historically occurred in this portion off 2011), and biodiversity inventories (see Author’s of the Mobile Basin (Tables 11-1 and 11-2). This is a Note). substantial portion of the approximately 65 freshwa- Mollusk Assemblage Structure ter mussel and 125+ gastropod taxa known from the Mobile Basin (Garner et al. 2004; Williams et al. 2008; Because they occupy linear habitats (i.e., stream Johnson et al. 2012). drainage networks), freshwater mollusk distributions

76 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area X X X X X X X X X X X X X X X X X X X X Recent X X X X X X X X X X X X X X X X X X X X X X X X X X X Historical ? T T T T T B M M M M M M M M B, T B, T M, T M, T M, T M, T M, T M, T M, T M, T M, T B, M B, M B, M, T B, M, T Habitat I I I S S S S S S S S S S S S S S S S S U FT FE FE FT FT FC FC FC FC Status Pondmussel Southern Hickorynut Fragile Papershell Black Sandshell Washboard Threehorn Wartyback Yellow Sandshell Yellow Alabama Moccasinshell Common Name Asian Clam Swamp Fingernail Clam River Fingernail Clam Herrington Fingernail Clam Striated Fingernail Clam Long Fingernail Clam Threeridge Flat Floater Delicate Spike Elephantear Southern Combshell Southern Pigtoe Round Pearlshell Finelined Pocketbook Mucket Orangenacre Southern Pocketbook Southern Fatmucket Alabama Pearlshell Rayed Creekshell Rock Pocketbook Butterfly Alabama Spike Ligumia subrostrata Obovaria jacksonaia Leptodea fragilis Ligumia recta Megalonaias nervosa Obliquaria reflexa Lampsilis teres Medionidus acutissimus Scientific Name Corbicula fluminea Musculium partumeium Sphaerium fabale Sphaerium occidentale Sphaerium striatinum Musculium transversum Amblema plicata Anodonta suborbiculata Elliptio arctata Elliptio crassidens Epioblasma penita Fusconaia cerina Glebula rotundata Hamiota altilis Hamiota perovalis Lampsilis ornata Lampsilis straminea Margaritifera marrianae Anodontoides radiatus Arcidens confragosus Ellipsaria lineolata Elliptio arca Family Cyrinidae Pisidiidae Unionidae Margaritiferidae Systematic list of freshwater bivalve taxa collected, conservation status (S-Stable, FC-Federal species of Concern, FT-Federally Threatened, FE-Federally Threatened, Concern, FT-Federally species of status (S-Stable, FC-Federal conservation bivalve taxa collected, list of freshwater 11-1. Systematic Table Alabama, Mobile, M-Main stem (B-Backwater/slough, and habitats occupied concern, U-Unknown/non-assessed), SC-Special I-Introduced, EX-Extinct, Endangered, River area greater Mobile-Tensaw surveys in the freshwater mollusk recent (1990-2015) (1900-1990) and during historical T-Tributaries) rivers, and/or Tombigbee al. (1998), et Turgeon follows Taxonomy counties, Alabama). Washington and Clarke, tributaries in Baldwin, rivers and and Tombigbee Mobile, Alabama, stem (main (2012). and Lydeard al. (2008), and Campbell Williams et

Biotic Landscape - Chapter 11 77 X X X X X X X X X X X X X X X X X X 38 Recent X X X X X X X X X X X X X X X X X X X X X X X X X 52 Historical T T T T T M M M M M M M M M M M M B, T M, T M, T M, T M, T M, T M, T B, M B, M, T Habitat S S S S S S S S S S S S S S S S S S FE FT FE FE FT EX EX SC Status Rayed Kidneyshell Southern Mapleleaf Alabama Orb Monkeyface Gulf Mapleleaf Ridged Mapleleaf Stirrupshell Ebonyshell Southern Creekmussel Heavy Pigtoe Pigtoe True Inflated Heelsplitter Bleufer Giant Floater Pistolgrip Southern Clubshell Alabama Hickorynut Bankclimber Ovate Pigtoe Common Name Paper Pondshell Little Spectaclecase Southern Rainbow Lilliput Liliput Texas Fawnsfoot Pondhorn Ptychobranchus foremanianus Quadrula apiculata Quadrula asperata Quadrula metanevra Quadrula nobilis Quadrula rumphiana Quadrula stapes Reginaia ebena Strophitus subvexus Pleurobema taitianum Pleurobema verum Potamilus inflatus Potamilus purpuratus Pyganodon grandis Quadrula verrucosa Pleurobema decisum Obovaria unicolor Plectomerus dombeyanus Pleurobema perovatum Scientific Name Utterbackia imbecillis Villosa lienosa Villosa vibex Toxolasma parvum Toxolasma texasensis Toxolasma donaciformis Truncilla Uniomerus tetralasmus Unionidae, cont. Family Total Taxa Total Systematic list of freshwater bivalve taxa collected, conservation status (S-Stable, FC-Federal species of Concern, FT-Federally Threatened, FE-Federally Threatened, Concern, FT-Federally species of status (S-Stable, FC-Federal conservation bivalve taxa collected, list of freshwater 11-1. Systematic Table Alabama, Mobile, M-Main stem (B-Backwater/slough, and habitats occupied concern, U-Unknown/non-assessed), SC-Special I-Introduced, EX-Extinct, Endangered, River area greater Mobile-Tensaw surveys in the freshwater mollusk recent (1990-2015) (1900-1990) and during historical T-Tributaries) rivers, and/or Tombigbee al. (1998), et Turgeon follows Taxonomy counties, Alabama). Washington and Clarke, tributaries in Baldwin, rivers and and Tombigbee Mobile, Alabama, stem (main (2012), cont. and Lydeard al. (2008), and Campbell Williams et

78 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area X X X X X X X X X X 10 Recent X X X X X X X X X X X X X X X X X X X X X 21 Historical T T T T B B B B B M M M M M M M M B, T B, T B, T B, T M, T Habitat S S S S S S E S S S S S S S V V V U U FE FT FC Status Mimic Lymnaea Pewter Physa Rams-horn Two-ridged Bugle Sprite Marsh Rams-horn Gladiator Elimia Marsh Fossaria Unknown Somatogyrus Unknown Amnicola Salt Spring hydrobe Globe Siltsnail Midland Siltsnail Carib Fossaria Common Name Dented Elimia Caper Elimia Spotted Rocksnail HornsnailBrook Tulotoma Snail Tulotoma Round Mysterysnail File Campeloma Cylinder Campeloma Two-ridged Valvata Two-ridged Pseudosuccinea columella Physa heterostropha Helisoma anceps Micromenetus dilatatus Planorbella trivolvis Elimia hydeii Fossaria (Galba) humilis Somatogyrus sp. Amnicola sp. Pseudotryoniagrahamae Birgella subglobosus Cincinnatia integra Fossaria (Galba) cubensis Scientific name Scientific Elimia taitiana Elimia olivula Leptoxis picta Pleurocera vestita Tulotoma magnifica Tulotoma Vivparus intertextus Campeloma limum Campeloma regulare Valvata bicarinata Valvata Physidae Planorbidae Pleuroceridae Amnicolidae Cochliopidae Hydrobiidae Lymnaeidae Family Total Taxa Total Viviparidae Valvatidae Systematic list of freshwater gastropod taxa collected, conservation status (S-Stable, V-Vulnerable, FC-Federal species of Concern, FT-Federally Threatened, Threatened, of Concern, FT-Federally FC-Federal species status (S-Stable, V-Vulnerable, conservation gastropod taxa collected, list of freshwater 11-2. Systematic Table rivers, Mobile, and/or Tombigbee stem Alabama, M-Main occupied (B-Backwater/slough, and habitats E-Endangered), Endangered, U-Unknown, FE-Federally Mobile, area (main stem Alabama, River the greater Mobile-Tensaw mollusk surveys in freshwater and recent (1990-2015) historical (1900-1990) during T-Tributaries) al. (2013). follow Johnson et and status rankings Taxonomy counties, Alabama). Clarke, and Washington tributaries in Baldwin, rivers and and Tombigbee

Biotic Landscape - Chapter 11 79 Freshwater Bivalves 3) Pisidiidae Freshwater bivalves comprise the majority of the fresh- Fingernail clams (formerly Sphaeriidae) are very water mollusk species found in the greater Mobile- poorly surveyed throughout much of the southeast- Tensaw River area. Freshwater pearly mussels (Order ern US, and therefore their taxonomy, distribution, Unionoida) have the greatest number of species (50+ and ecological associations are poorly known. The taxa), but have also likely experienced some of the 6 records of sphaeriid taxa from the greater Mobile- most dramatic range shifts in this region. The other Tensaw River area were all collected during historical major group of freshwater bivalves, fingernail clams surveys (Table 11-1). There are no records of finger- (Family Pisidiidae), is poorly known from this region, nail clams subsequent to 1970, but populations of with few collections from Clarke, Mobile, and Wash- most fingernail clam taxa likely remain extant in the ington counties. region.

Most recently, the region has been invaded by the 4) Unionidae Asian clam (Corbicula fluminea). Populations of this Historically the lower Alabama, Mobile, and lower viviparous and highly tolerant bivalve are now found Tombigbee rivers likely supported as many as 50 throughout the main stem lower Alabama River and species of Unionidae. Although numerous species many of its larger, permanent tributaries (McGregor have apparently been extirpated from the lower et al. 1999; McGregor and Garner 2006). Estuarine Alabama and lower Tombigbee rivers as a result bivalves, including members of the families Dreisse- of hydropower and navigation projects, recent nidae (Mytilopsis leucophaeta) and Rangidae (Rangia surveys suggest as many as 30 species remain extant cuneata), occur in the lower reaches of the system in in the main stem Alabama River and its tributaries sections inundated by the salt wedge (Chadwick and (McGregor and Garner 2006; Gangloff et al. 2009). Feminella 2001). Unionid assemblages in the lower main stem Tombig- bee River are somewhat less diverse, with populations 1) Cyrinidae of 20 taxa reported during recent surveys. The freshwater bivalve family Cyrinidae has one representative in the study area, the introduced Asian Clam, Surveys by Royal Sutkiss from 1964 to 1976 revealed Corbicula fluminea. This species was likely introduced specimens of 24 freshwater mussel taxa in the main deliberately from Asia into streams in the Pacific stem Alabama River and in Big Flat Creek, includ- Northwest of North America in the early part of the ing records for 2 species currently listed as federally 20th century. It has subsequently colonized much of threatened (Hamiota perovalis and Medionidus acutis- North America, including much of the southeastern simus) (Williams et al. 2008; see Author’s Note). Spec- US. First collected from Alabama in the late 1960s, imens of federally endangered E. penita, Pleurobema the species has become one of the more common and decisum, and P. taitanum have been reported from widespread bivalves in the region (Jenkinson 1979; prehistoric middens near the Salt Creek saltworks, so see Author’s Note). Although numerous workers presumably these taxa were once extant in the nearby have speculated that Asian clams may impact native Tombigbee River (McGregor and Dumas 2010). No bivalves, there is little empirical evidence of competi- recent collections of these taxa have been made from tive exclusion or other direct impacts. the study area and they are presumed to be locally extirpated. 2) Margaritiferidae Tributary freshwater mussel assemblages in the greater The freshwater bivalve family Margaritiferidae is Mobile-Tensaw River area are typically quite different represented in the study region by a single species, from assemblages found in river habitats, although Margaritifera marrianae (Alabama Pearlshell), which some taxa—including Lampsilis straminea, L. teres, occurs in small tributaries of the Alabama and Escam- Potamilus purpuratus, and Quadrula rumphiana— bia rivers draining the Red Hills region of south commonly occur in both habitat types. In Big Flat Alabama (see Author’s Note). Populations are known and Bassett creeks, Quadrula asperata, Lampsilis stra- from the Big Flat and Limestone creek watersheds. minea, and Villosa lienosa are among the more numer- This species was added to the Endangered Species ous taxa, although these systems support at least 15 List by USFWS in 2012 (USFWS 2012a). other mussel species (Figure 11-7).

80 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Impounded reaches of the Alabama and Tombigbee rivers support species-poor mussel assemblages comprised of species that are tolerant of the slower flows, softer substrates and lower dissolved oxygen concentrations that typify these altered habitats. Although many unionids common in impoundments are thin- shelled taxa (e.g., Pyganodon grandis, Utterbackia imbecillis), some heavy- shelled taxa including Quadrula apiculata and Plectomerus dombeyanus are also common in impounded habitats (Figures 11-8 and 11-9).

5) Estuarine Bivalves Euryhaline bivalves (including Mytilopsis leucophaeta, Polymesoda Figure 11-7. Lampsilis straminea from Bassett Creek, Clarke County, Alabama. caroliniana, and Rangia cuneata) Specimens of 6 other unionid taxa were collected in this relatively large but are poorly sampled in the greater under surveyed Tombigbee River tributary. (Photo credit: Michael M. Gangl- Mobile-Tensaw River area. They are off, Appalachian State University.) likely present throughout the lower reaches of the Mobile River and its tributaries, though not reported to date in the Alabama or Tombig- bee rivers. However, as sea levels continue to rise and the floodplain of the Mobile River subsides as a result of upstream (impoundment-driven) sediment capture, these taxa will likely become more common (and may replace unionids) in deeper, main channel habitats expected to experience the greatest degree of salt wedge intrusion.

Freshwater Gastropods A total of 22 primarily freshwater gastropod taxa have been reported from the greater Mobile-Tensaw Figure 11-8. Quadrula apiculata collected from Donnelley Reservoir, an River area (Table 11-2). Of these, 21 impoundment on the Alabama River in Wilcox County. This species is common in impounded and free-flowing reaches of the Alabama and Tombigbee taxa were documented during histor- rivers in the greater Mobile-Tensaw River area. (Photo credit: Michael M. ical (pre-1995) surveys, whereas Gangloff, Appalachian State University.) recent surveys have revealed populations of only 10 taxa. This discrep- ancy is likely due to so few recent surveys targeting freshwater gastropods in this region. More focused surveys will probably reveal that this region still harbors a substantial

Biotic Landscape - Chapter 11 81 been ascribed to any known taxon. Careful genetic and morphological studies are needed to resolve the taxonomic affinities of these populations.

2) Pleuroceridae Compared to other parts of the Mobile Basin, the lower Alabama River area has relatively few pleurocerid species. Elimia taitiana appears to be the most widespread taxon in smaller tributary streams, including Limestone Creek (Figure 11-5). Pleurocera vesti- tum, a large silt-tolerant pleurocerid, occurs in slow- moving main channel and backwater habitats (Table 11-2). Elimia olivula, a species currently being considered for protection under the Endangered Species Act, occurs in the main stem Alabama River near Clai- borne, as does Leptoxis picta, a federally endangered pleurocerid that occurs in the tailwaters of Claiborne Lock and Dam and few other places on earth (Garner et al. 2004; see Author’s Note).

3) Viviparidae The Tulotoma Snail (Tulotoma magnifica), the only high-conservation priority viviparid snail extant in the greater Mobile-Tensaw River area, was initially described from material collected near Claiborne, Alabama. Presumed extinct until rediscovered in the Coosa River and several tributaries during the late 1980s, recent surveys detected a small population downstream of Claiborne Lock and Dam Figure 11-9. Plectomerus dombeyanus collected (Hershler et al. 1990; USFWS 2011). This discov- from Donnelley Reservoir, an impoundment on ery, along with detection of additional populations the Alabama River in Wilcox County. This species and evidence of population growth, led USFWS to is common in impounded reaches and backwater downlist T. magnifica from endangered to threat- habitats of the Alabama and Tombigbee rivers in the greater Mobile-Tensaw River area. (Photo ened. Other viviparid taxa in the lower Alabama River credit: Michael M. Gangloff, Appalachian State are include Campeloma limum and C. regularae (see University.) Author’s Note). The systematics and distribution of Campeloma spp. remain unclear and the genus needs genetic and morphological study. portion of its native freshwater gastropod fauna and may help clarify taxonomic issues associated with 4) Pulmonate Snails (Lymnaeidae, Physidae, several unresolved taxa. and Planorbidae) 1) Pebble Snails (Amnicolidae, Cochliopidae, Pulmonate snails lack gills and respire via pallial lungs. and Hydrobiidae) This allows them to tolerate low dissolved oxygen habitats, including ponds, sloughs, and marshes. The greater Mobile-Tensaw River area has a surpris- Because they are relatively tolerant of environmental ingly high diversity of hydrobiid snails, with repre- conditions that would be lethal to gilled snails (e.g., sentatives of 5 genera present. Of these, only Pseudot- Elimia, Leptoxis), few pulmonates are considered ryonia grahamae, the endemic Salt Springs Hydrobe, conservation priorities. However, there have been few appears to be a high conservation priority taxon recent collections of any members of these 3 fami- (Table 11-2). However, specimens of Amnicola and lies from the greater Mobile-Tensaw River area (see Somatogyrus collected from the region have not yet

82 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Author’s Note). Future researchers should make an mollusk populations in this region of the Gulf Coastal effort to document their occurrence and collect speci- Plain. Interestingly, recent surveys in Big Flat Creek, mens whenever possible. just downstream of Rikard’s Mill, revealed a species- rich mussel assemblage (Gangloff et al. 2009, 2011). Current Status and Threats to Mollusk This phenomenon appears to be relatively widespread Resources in southeastern Coastal Plain and Piedmont streams Comparisons between recent and historical survey and suggests that some structures (notably older data, combined with insights from studies of shells historic dams) may counterintuitively improve unio- in Native American middens, suggest that greater nid habitat in some streams (Gangloff 2013). Mobile-Tensaw River area mussel (and likely snail) Third, long-term stressors, including the effects of populations have changed over the past several centu- climate change and sea level rise, may reduce the ries. Although much of this change is likely due to downstream limits of freshwater mollusk habitat in widespread impoundment and dredging of large river the lower Mobile Basin. Although the effects of climate habitats, there is some evidence that the activities of change have been poorly studied in this system, their Native Americans may have altered mussel assem- consequences for freshwater resources likely pose the blages well before the arrival of Europeans. For exam- single greatest threats to biodiversity in this sparsely ple, Peacock et al. (2005) reported noticeable changes developed region of Alabama. to the composition of freshwater mussel assemblages in prehistoric middens, which they attributed to the Information gaps simultaneously represent the larg- advent of subsistence agriculture and possible changes est but most easily addressed threats to freshwater in water quality. mollusk diversity in the greater Mobile-Tensaw River area. Numerous systematic questions need to be Among many threats to native mollusk assemblages in resolved, including determination of the taxonomic the greater Mobile-Tensaw River drainage, the most affinities of several gastropod and unionid taxa (see pressing come from 3 sources. First, the construction, Author’s Note). Additionally, the US Fish and Wild- maintenance, and operation of large dams upstream life Service is currently reviewing the need to protect in the Mobile Basin for navigation, flood control, and several mollusk taxa found in the greater Mobile- power generation are chronic stressors to main stem Tensaw River area under the Endangered Species Act river and tributary mollusk populations (Williams et (Figure 11-10). Although the region’s larger streams al. 1992; USFWS 2000; Williams et al. 2008). Water have been well-covered by qualitative surveys (e.g., levels in the Alabama River downstream of Claiborne Hartfield and Garner 1998; McGregor and Garner Lock and Dam fluctuate by as much as 3 m (10 ft) daily 2006), smaller streams have been far less extensively and these changes have dramatic effects on mollusk surveyed and many streams or localities have only populations. Tributary confluences may be inundated been surveyed once during the past 45 years. It seems by rising waters and then rapidly dewatered during reasonable to assume that numerous additional popu- power generation events. Additionally, impounded lations and taxa remain undiscovered in the greater habitats are more likely to support populations of Mobile-Tensaw River area. Moreover, there have non-native unionids, like Anodonta suborbiculata and been few attempts to use quantitative or occupancy- Ligumia subrostrata, and may facilitate the stepping- based surveys to understand how local or landscape- stone advance of these taxa into native habitats. scale habitat parameters influence the diverse array of Second, land use at much finer scale including freshwater mollusk assemblages present in the greater management of upland and riparian forests, exurban Mobile-Tensaw River area. Finally, establishment of and infrastructure development, and the construction permanent monitoring sites, both in high diversity of small earthen impoundments in headwaters may mollusk habitats and in systems that support unique affect mollusk habitats and populations. However, taxa (e.g., Pseudotryonia grahamae), to assess tempo- evidence that these factors are impacting freshwater ral changes to population and assemblage structure mollusk populations is sparse and there have been no will help ensure the persistence of this region’s unique studies to the author’s knowledge that have quanti- mollusk resources. fied effects of land use on stream habitats or sensitive

Biotic Landscape - Chapter 11 83 Figure 11-10. Anodontoides radiatus from a tributary of the Alabama River in Wilcox County. This species occurs in small, sandy streams on the Coastal Plain from the Mississippi Drainage east to the Apalachicola Drainage in Florida and is currently being considered for federal protection by the US Fish and Wildlife Service. (Photo credit: Michael M. Gangloff, Appalachian State University.)

Author’s Note profile/Michael_Gangloff/contributions). The author Museum records for freshwater mollusks collected gratefully acknowledges the museums involved: the from localities within the greater Mobile-Tensaw River Auburn University Natural History Learning Center area, as described in this chapter, can be found in DOI: and Museum, the Mississippi Museum of Natural 10.13140/RG.2.1.1931.6241 at the author’s page on the Sciences, and the Florida Museum of Natural History ResearchGate website (https://www.researchgate.net/ at the University of Florida.

84 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 12. Crustaceans James A. Stoeckel and Brian S. Helms, Auburn University

This chapter reviews the current status of crustaceans often inhabit slow-moving surface waters, includ- in the greater Mobile-Tensaw River area, drivers and ing ditches, swamps, and sluggish streams, and can stressors impacting them, and key stewardship needs. be locally abundant (Penn 1956; Smith et al. 2011). Several species are largely confined to the lower Current Knowledge—Natural, Cultural, Alabama (P. bivittatus, P. clemmeri, P. evermanni, P. and Economic Importance lagniappe, P. lecontei, P. penni, and P. shermani). The lower Alabama River, lower Tombigbee River, Many crayfish found in the Mobile-Tensaw Delta and Mobile-Tensaw Delta host a diverse assemblage show moderate to strong propensity for burrowing, of freshwater crustaceans notable for their ecological including all 5 species of Cambarus. Fallicambarus is and economic importance. In the freshwater, non- also highly adapted to a subterranean existence, with a tidal portions of the system, the crustacean macro- vaulted carapace, spatula-like legs, and reduced eyes, invertebrate fauna is dominated by crayfish. As tidal antennae, and abdomen. While the Cambarus species influences and salinity increase in the Mobile-Tensaw can be found in a wide range of habitats, the 3 species Delta, fiddler crabs gradually replace crayfish as the of Fallicambarus found in the Mobile-Tensaw Delta dominant, semi-terrestrial burrowing crustacean and are often associated with pitcher-plant bogs, pine Blue Crabs (Callinectes sapidus) replace crayfish as the flatwoods, swamps, and seepage areas of tributaries dominant aquatic crustacean. Finally, as the Mobile- (Fitzpatrick 1987; Graydon 2009). Tensaw Delta empties into Mobile Bay, a wide range of crabs, prawns, and shrimps depend on estuaries as Some of the more intriguing crayfishes are the critical habitat for various life history stages. dwarf crayfishes represented by species in the genus Cambarellus. The 3 species found in the Delta are 1) Crayfish generally rare with highly restricted distributions Global crayfish diversity peaks in North America inhabiting oxbow lakes, heavily vegetated pools, with over 350 species (Taylor et al. 2007). Within ditches, and other sluggish aquatic habitats (Stites et North America, Alabama has the highest number of al. 2014). These animals are particularly small, with crayfish species of any state or province (86 species, the Least Crayfish being one of the smallest crayfishes plus several undescribed forms), presumably due in North America (less than 25 mm [1 in] as adult). to geologic diversity and isolation of major drainage Nearly nothing is known about their biology. basins (Ortmann 1905, 1931; Hobbs 1969; Schuster Crayfishes also have a rich cultural and culinary et al. 2008; Smith et al. 2011). Crayfishes are found in history in Alabama. The Red Swamp Crawfish a range of aquatic and terrestrial habitats, including (Procambarus clarkii) and the White River Crayfish streams, lakes, wetlands, and terrestrial burrows. They (P. zonangulus) are the 2 primary species cultured for play instrumental roles in community dynamics, with consumption (Smith et al. 2011). Although the native strong effects on other organisms, and can influence range of both species is unclear, it is generally agreed ecological function via herbivory, predation, organic that P. clarkii is native to at least far southern Alabama, matter breakdown, and sediment processing (Creed potentially north to the Fall Line. As of 2012, there 1994; Lodge et al. 1994; Momot 1995; Charlebois and were 12 commercial farms in Alabama, producing Lamberti 1996; Usio 2000; Helms and Creed 2005; slightly over $2.3 million in sales (USDA 2014). Aver- Twardochleb et al. 2013). age Alabama farm yields are around 1000 pounds In the Mobile Tensaw Delta, 6 genera are represented per acre (0.1 kg/m2) (Masser et al. 1997). Crawfish by 34 species (Table 12-1, Figure 12-1). Procambarus farms are often rice-crayfish-soybean polycultures, is the most diverse genus with 17 species. Procam- with crayfish inhabiting rice fields that are gradually barus spp. in the greater Mobile-Tensaw River area flooded during the warm spring months (Walls 2009).

Biotic Landscape - Chapter 12 85 Table 12-1. Crayfish found in the Mobile-Tensaw Delta (MD), lower Alabama River (AL), Lower Tombigbee (LT), and adjacent local watersheds. “AFS Status” is conservation status as assessed by the American Fisheries Society (Taylor et al. 2007) and “AL Priority” is conservation priority assigned by the Alabama Department of Conservation and Natural Resources.

Scientific Name Common Name AFS Status AL Priority MD LT AL Cambarellus diminutus Least Crayfish T P2 X Cambarellus lesliei Angular Dwarf Crayfish T P3 X X Cambarellus shufeldti Cajun Dwarf Crayfish T P2 X Cambarus diogenes Devil Crayfish CS P5 X X X Cambarus latimanus Variable Crayfish CS P4 X Cambarus ludovicianus Painted Devil Crayfish CS P3 X X Cambarus miltus Rusty Grave Digger T P3 X Cambarus striatus Ambiguous Crayfish CS P5 X Fallicambarus danielae Speckled Burrowing Crayfish T P3 X X Fallicambarus burrisi Burrowing Bog Crayfish CS P2 X Fallicambarus fodiens Digger Crayfish CS P5 X X Faxonella clypeata Ditch Fencing Crayfish CS P3 X X X Hobbseus prominens Prominence Riverlet Crayfish CS P2 X Orconectes chickasawae Chickasaw Crayfish CS P3 X X Orconectes holti Bimaculate Crayfish T P3 X X Orconectes jonesi Sucarnoochee River Crayfish T P3 X X Orconectes perfectus Complete Crayfish CS P5 X X Procambarus acutissimus Sharpnose Crayfish CS P5 X X Procambarus acutus White River Crayfish CS P5 X X X Procambarus bivittatus Ribbon Crayfish CS P3 X X X Procambarus clarkii Red Swamp Crayfish CS P5 X X X Procambarus clemmeri Cockscomb Crayfish CS P2 X Procambarus evermanni Panhandle Crayfish CS P2 X X Procambarus hybus Smoothnose Crayfish CS P2 X Procambarus lagniappe Lagniappe Crayfish T P2 X Procambarus lecontei Mobile Crayfish SC P2 X X Procambarus lophotus Mane Crayfish CS P3 X X Procambarus marthae Crisscross Crayfish SC P2 X Procambarus penni Pearl Blackwater Crayfish V P3 X X Procambarus shermani Gulf Crayfish CS P3 X X X Procambarus spiculifer White Tubercled Crayfish CS P5 X X X Procambarus versutus Sly Crayfish CS P5 X X X Procambarus vioscai paynei Payne’s Creek Crayfish C3 P3 X

86 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Animals are harvested from December through June (Masser et al. 1997).

2) Crabs Several species of fiddler crab are found in the Mobile Tensaw Delta. All are semi-terrestrial and breathe air. Some like Uca minax (Red Jointed Fiddler Crab) are found in freshwater and low salinity tidal marshes, whereas others like U. longisignalis (Gulf Mud Fiddler Crab) prefer moderate to high salinity habitats beyond the Delta. Males are easily recognized by an enlarged claw, which they wave back and forth to entice females into a nearby burrow for mating. Though not valued as a food item by humans, they are eagerly consumed by fishes, birds, and small mammals. Fiddlers also play an important role as ecosystem engineers, modifying their physical habitat through burrowing and feeding activities. Burrows are constructed for shelter, protection from predators, and reproduction. In large fiddler crab colonies, burrow density can reach 300/m2 (28/ft2), resulting in 120 to 160 cm3 of sediment/m2 (0.7 to 0.9 inch3 of sediment/ft2) excavated per day, thoroughly mixing the top 8 to 15 cm (3 to 6 inch) of sediment (McCraith et al. 2003). Burrowing activity can strongly influence transfer of energy and nutrients in marshlands, bringing increased organic material to the surface and fostering growth of marsh grasses, while speeding decomposi- tion of plant material via stimulation of microbial communities (Katz 1980; Montague 1980; Daiber 1982; Bertness 1985).

Blue Crabs (Callinectes sapidus) are also found in low to high salinity habitats but, unlike fiddler crabs, live primarily in water. Blue Crabs of both sexes are eagerly sought by commercial

Figure 12-1. Select crayfish from the Mobile-Ten- saw Delta, lower Alabama River, lower Tombig- bee, and adjacent watersheds. A) Cambarus miltus, B) Faxonella clypeata, C) Cambarellus diminutus, D) Cambarellus shufeldtii, E) Orconectes jonesi, F) Orconectes perfectus, G) Fallicambarus burrisi, H) Fallicambarus fodiens, I) Procambarus shermani, J) Procambarus marthae, K) Procambarus lagniappe, L) Procambarus versutus, M) Procambarus acutissimus, N) Procambarus clarkii. (Image credit: Guenter Schuster.)

Biotic Landscape - Chapter 12 87 and recreational fishermen. In Alabama, annual Key Drivers and Stressors harvest increased from less than 1 million pounds Due in part to their complex life cycles, crustaceans in (0.45 million kg) before 1940 to a peak of 4.2 million lb the greater Mobile-Tensaw River area are affected by (1.9 million kg) in 1984, before decreasing to an aver- many environmental drivers and stressors, which we age of 2 million lb (0.9 million kg) per year—with a review here. dockside value approximately $2 million—since 2000 (VanderKooy 2013; Audubon Nature Institute [date 1) Connectivity unknown]). One of the most important factors impacting crustaceans is status of habitat connectivity (Pringle 3) Shrimp 2003). A lack of connectivity (i.e., habitat fragmen- The young of many marine crustaceans are also found tation) is considered one of the leading causes of in the low salinity, estuarine habitats of the Delta and global species loss (Fischer and Lindenmayer 2007), upper Mobile Bay. Shrimp are of particular impor- particularly when associated with limited resource tance, with the Brown Shrimp (Penaeus aztecus) use or restricted gene flow (Fahrig 2003; Omerod designated as Official State Crustacean of Alabama in 2003). Many species of crayfish, particularly endemic March 2015. Brown Shrimp, White Shrimp (P. setifer- species, may have low dispersal rates, which increases ous), and Pink Shrimp (P. duorarum) are all harvested their susceptibility to loss of connectivity (Bubb et al. by the Alabama shrimp fleet. The Alabama shrimp fish- 2006; Helms et al. 2015). Connectivity between favor- ery began in the early 1900s and peaked in the 1980s able habitat patches is essential for population persis- with an estimated 2000 commercial shrimp licenses tence (Figure 12-2). Discontinuities such as culverts, issued. Commercial shrimping remains a valuable road crossings, and dams either physically restrict industry with 17.7 million lb (8 million kg) landed movement of individuals or create vast areas of inhos- in 2014 (dockside value greater than $57 million). pitable habitat, both of which can isolate popula- Recreational shrimping is also a popular hobby and tions, with detrimental effects (Hartfield 2010; Adams family tradition with greater than 1000 recreational 2013; Foster and Keller 2011). Further, connectivity fishing licenses issued to Alabama citizens every year between terrestrial and aquatic habitats is critical for (Alabama Gulf Seafood 2015). many burrowing crayfish species. For C. diogenes, a species that moves between open water and terrestrial

Figure 12-2. Connectivity between aquatic and semi-terrestrial habitats is critical for crayfishes. Many species burrow exclusively in terrestrial environments, either in floodplains (A) or isolated wetlands (B). These primary burrowers rely on subsurface groundwater and move from subterranean to terrestrial habitats and to some degree from terrestrial to surface water habitats depending on species and life stage. Some species burrow in close proximity to surface water (C), often with direct connections to surface water. These secondary burrowers can freely move between terrestrial, subterranean, and surface water habitats. Other species spend their entire life cycle in surface water (D). Thus, maintaining hydrologic connectivity and protecting isolated wetlands is a fundamental component to successful crayfish conservation.

88 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Marine shrimp (Peneai- dae, Penaeus spp.) are dependent on saltwater- brackish connections (Figure 12-4). Mating, spawning, and development of early planktonic larval stages occur in offshore marine habitat. Post-larvae utilize shoreward currents and a combination of light and salinity cues to migrate to brackish Figure 12-3. Ohio Shrimp (Macrobrachium ohione) male (left) and female (right). This estuaries that serve as species is now rare to absent in the Mobile-Tensaw Delta. (Photo credit: Ray Bauer.) vital nursery areas of all 3 species. Brown Shrimp

post-larvae typically migrate to estuaries in the spring, burrows, decreased crayfish burrow abundance in White Shrimp in the summer, and Pink Shrimp in the terrestrial floodplains is associated with increased winter. Growth and survival of post-larval shrimp are stream incision (Helms et al. 2013). For other burrow- severely limited if they fail to reach estuarine nurser- ers, particularly those that inhabit bogs, wet meadows, ies. Once in the nurseries, post-larvae metamorphose and swamps (e.g., Fallicambarus species), draining of into the juvenile stage, which feed and grow in the wet fields and similar environs is a major threat (Gray- estuaries. Late-stage juveniles then migrate back to don 2009; Stites et al. 2014), as these activities lower the open ocean to complete their life cycle (Matthews the local water table and increase the prevalence of et al. 1991; Howe et al. 1999). Availability of high qual- inhospitable habitats. ity estuarine habitat and strong connectivity between For other crustacean taxa, critical connectivity occurs Mobile Bay estuaries and the Gulf are vital for main- at the interfaces between fresh, brackish, and saltwa- taining healthy populations of these highly valued ter habitats. A little known crustacean group, formerly species. abundant in the Gulf of Mexico coastal drainages, is Blue Crabs and fiddler crabs are also dependent the freshwater prawn or River Shrimp (Paleomonidae, on connectivity between Mobile Bay estuaries and Macrobrachium spp.). Six species are known from the offshore marine environments (Figure 12-4). Blue US, all dependent on freshwater-brackish connec- Crab females migrate out of brackish estuaries and tions. Macrobrachium ohione (Ohio Shrimp, Figure into high salinity offshore waters to hatch their eggs. 12-3), once supported a valuable US fishery as late as Upon hatching, the free swimming zoea (larvae) feed the 1930s (Mermilliod 1976; Bowles et al 2000), but in offshore water before metamorphosing into mega- numbers have since plummeted, presumably due to lopae and migrating back to estuaries, where they damming of large river systems and loss of connectiv- settle out and metamorphose into juveniles. Settle- ity with brackish estuaries. River Shrimp adults live in ment in Mobile Bay occurs from June through Novem- freshwater, but females must migrate downstream to ber, peaking from July through mid-October (Morgan deliver larvae that require saline habitats for proper et al. 1996). Males typically spend their entire juve- development (Oliver and Bauer 2011; Oliver et al. nile and adult lives in estuaries, while adult females 2012). Juveniles then migrate upstream to freshwa- will migrate back to offshore waters (VanderKooy ter habitats (Figure 12-4), but this migration may be 2013). Fiddler crab females are triggered by a combi- impeded by dams (Olivier et al. 2013). Still present in nation of tide and moon phase to migrate from estu- the lower Mississippi and Atchafalaya river systems aries to nearshore areas where their eggs hatch. The of Louisiana, Ohio Shrimp were reported from the resultant planktonic larvae migrate to high salinity Alabama River as late as 1971 (Bowles et al. 2000), but offshore waters and then back to estuaries, where, are now extremely rare or absent from the Mobile- similar to Blue Crab, the megalopae stage settles and Tensaw Delta.

Biotic Landscape - Chapter 12 89 flowing over firm sand substrates, a habitat preference shared with some other Procambarus and all the Orco- nectes species in the area (McGregor et al. 1999). Also of particular importance in the Mobile-Tensaw Delta is the presence of submerged woody debris and aquatic vegetation that crayfishes use for spawning, feeding, and refuge. The importance of such habitat is evident in the preponderance of species (e.g., many Procambarus spp., Cambarellus spp.) generally found in slow streams, vegetated sloughs, vernal pools, and even roadside ditches (Hobbs 1981; Walls 2009). Depletion of aquatic vegetation and woody debris can drastically affect crayfish populations (Taylor et al. 2007). Similarly, vegetated brackish habitats, such as marshes and areas containing submerged aquatic vegetation, are very important for young- of-year crab and shrimp (Orth and van Montfrans 1990; Heck et al. 2001; Rozas et al. 2013).

3) Sedimentation One of the more pervasive threats to freshwater crayfishes is sedimentation from sources such as land clearing, road runoff, and new construction. About 15% of stream miles assessed in the US are considered threatened or impaired by suspended sediments (USEPA 2013). Sedimentation can increase emigration, clog gills, reduce growth, and alter the interactions of Figure 12-4. Importance of linkages between freshwater, brackish, and saltwater habitats for crabs and shrimp as illustrated by life history-stage aquatic organisms, as well as bury criti- migrations. cal habitats (Newcombe and MacDon- ald 1991; Wood and Armitage 1997).

Fortunately, none of the watersheds in the lower metamorphoses into juvenile crabs in brackish water Alabama and Tombigbee rivers and Mobile-Tensaw estuaries. Delta are listed by the US Environmental Protection Agency (USEPA) as significantly impaired from sedi- 2) Habitat Type ment (USEPA 2015). Crayfishes in general depend on a variety of habitat types, ranging from gravel, cobble, and boulder to 4) Dissolved Oxygen sand and detritus substrates. Many crayfishes in the As evidenced by their preferred habitats and known greater Mobile-Tensaw River area are associated with natural history, the crayfish inhabiting the Mobile- sandy stream bottoms. For example, the Lagniappe Tensaw Delta generally are relatively tolerant of low Crayfish (P. lagniappe), appears to require cool water dissolved oxygen. Sensitivity to low dissolved oxygen is

90 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area species specific, with lower-lethal limits ranging from 7) Chemical Pollutants 0.4 to 2.6 mg/L (0.4 to 2.6 parts per million) depend- Crustaceans are susceptible to a range of metals, ing on the species. Distributions are typically driven plasticizers, herbicides, and pesticides (Krebs et al. by sublethal concentrations that are much higher 1974; Millikin and Williams 1984; Oehlmann et al. than lower-lethal concentrations (Nyström 2002). 2009), with metal toxicity frequently increasing with Since many of the lowland streams in the Mobile- decreasing salinity (Hall and Anderson 1995). Non- Tensaw Delta naturally have low dissolved oxygen, lethal effects of metals (e.g., copper) and herbicides the animals inhabiting these systems have adapted to (e.g., metolachor) on crustaceans include reduced these conditions. In fact, crayfish can respire out of chemosensory function and subsequent fitness (Wolf water as long as their gills—which do not collapse in and Moore 2002; Olsen 2011; Blinova and Cherkashin open air like those of most aquatic animals—retain 2012). Because they are important food items for vari- some moisture (McMahon 2001). If aquatic dissolved ous predators (Grimes et al. 1989), crustaceans can oxygen concentrations fall below certain thresholds, accumulate and transfer toxins such as PCBs (poly- many species (e.g., P. clarkii) will climb to the surface chlorinated biphenyls) from contaminated sediments via woody debris or banks, perching sideways to take to aquatic, terrestrial, and avian food webs. advantage of atmospheric oxygen (Nyström 2002). Oil and natural gas pollution comes from a combi- Blue Crabs and shrimp are highly mobile and typically nation of natural seeps (Farrington and McDowell move out of an area when dissolved oxygen levels 2013), localized oil spills, and large industrial acci- decline to dangerous levels. This phenomenon, long dents such as the Deepwater Horizon spill in 2010. recognized in Mobile Bay (Loesch 1960; May 1973; Although coastal oil spills garner significant publicity, Tatum 1982), is termed a “jubilee,” as large numbers recent evidence suggests that the major proportion of of crabs and shrimp migrate to the shores to escape petroleum spills occur inland (Yoshioka and Carpen- low-oxygen water. Jubilees attract large crowds of ter 2002). With increasing demand for petroleum, as people and result in popular celebrations in Mobile well as new and existing pipelines in Alabama, inland Bay (e.g., see City of Fairhope [date unknown]). oil pollution is potentially a concern for crustaceans However, low dissolved oxygen events can occasion- living in the freshwater/riverine portions of the Mobile- ally result in massive mortality, especially if crabs are Tensaw Delta. Crude oil and biodiesel chemicals are trapped in cages when the events occur (May 1973; sensory pollutants to crayfish that bind and otherwise VanderKooy 2013). Fiddler crab adults are primar- damage chemoreceptors, thus decreasing their abil- ily terrestrial, breathing air from the atmosphere, and ity to forage, mate, and avoid predators (Jurcak et al. thus are less affected by dissolved oxygen than Blue 2015). Oil pollution is also a potential problem for life Crabs and shrimp. stages of other crustacean species that are dependent on estuarine and saltwater portions of Mobile Bay. 5) Temperature and Salinity Oil can have a variety of negative effects ranging from Thermal tolerance of Blue Crabs is dependent on salin- physical clogging and morphological damage to gills ity, with crabs becoming less tolerant of temperature (Perry and VanderKooy 2015) to decreased survival extremes as salinity decreases (Tagatz 1969; Holland and reproductive effects (Anderson 2010). Planktonic 1971; for summary, see Perry and VanderKooy 2015). crab larvae drifting near the surface are particularly The optimal range for Blue Crabs has been estimated susceptible to hydrocarbon pollutants due to photo- at 0 to 27 parts per thousand salinity and 10 to 35oC induced toxicity. When exposed to polycyclic hydro- (50 to 95oF) (Copeland and Bechtel 1974). carbons in the presence of solar ultraviolet radiation, planktonic crab larvae exhibit much higher mortality 6) Freshwater Inflow than when exposed in the absence of ultraviolet radia- Freshwater inflow appears to be the most important tion (Peachey 2005; Alloy et al. 2015). Burrowing crus- driver of Blue Crab population dynamics in most, if taceans may be affected by oil residues buried beneath not all, of the northern Gulf of Mexico watersheds, salt marsh sediments decades after a spill (Culbertson including the greater Mobile-Tensaw River area et al. 2007). (VanderKooy 2013). Both juvenile abundance and commercial landings of adults tend to be high in wet years, but decrease in dry years.

Biotic Landscape - Chapter 12 91 Resource Conservation Implications 2013). Maintenance of salinity gradients is also para- For many of the crayfish in the greater Mobile-Tensaw mount. Alterations in freshwater inflow/saltwater River area, there is a general lack of species-specific life intrusion patterns, as a result of water withdrawal history, distribution, habitat requirements, and other for agricultural and industrial uses and global climate ecological information. Thus hindered, management change, can potentially have devastating effects. Large recommendations are often not targeted, but instead groundwater withdrawals have been reported in the are based on generalities or incomplete data. Addi- coastal zones of Baldwin and Mobile counties (Twil- tional survey and taxonomic work is also needed for ley et al. 2001). Such changes in supply, coupled with many crayfish taxa. For burrowing species, particu- changes in rainfall patterns (e.g., drought), are viewed larly Fallicambarus species, maintenance of the native as a major threat to the sustainability of the Blue Crab vegetation structure and hydrologic regime of coastal fishery of Alabama (Perry and VanderKooy 2015). plain swamps, wetlands, and pitcher plant bogs is crit- Most people are generally familiar with crabs, shrimp, ical, including the use of prescribed fire. Once critical and crayfish. However, many are shocked to learn that biological and distributional limits are recognized for there is more than one “type” of each of these animals, all pertinent species, better informed guidelines can and even more surprised upon learning about the be implemented for resource conservation. levels of Alabama’s aquatic biodiversity. Public knowl- The protection and maintenance of natural flows, edge of the natural heritage associated with our native seasonally flooded areas, groundwater fluctuations, crustaceans is critical to their conservation. Convey- and general hydrologic connectivity are of considering the ecological, commercial, and cultural value of able importance to the conservation of crustaceans in these animals is the essential first step towards effec- the greater Mobile-Tensaw River area. This is pertinent tive management and conservation. in terms of future construction of dams, impoundments, road crossings, and bridges (e.g., Rozas et al.

92 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 13. Fishes Dennis R. DeVries and Russell A. Wright, Auburn University

This chapter describes the fishes of the greater Mobile-Tensaw River area, including the diverse habitats available, impacts of salinity to species makeup, threats to fish diversity and populations, and resource conservation implications.

The Landscape for Fishes Aquatic habitats in the greater Mobile-Tensaw River area (hereafter “the Delta”) are diverse and complex, ranging from large-riverine habitat upstream, to complex braided river channel, bayou, creek, and marsh habitat downstream. Approximately 10 km (6 mi) below the confluence of the Alabama and Tombigbee rivers, the Mobile River divides into two main channels, the Mobile River Figure 13-1. Byrne’s Boat Ramp—a typical backwater habitat in the greater and the Tensaw River. Between them Mobile-Tensaw River area at the mouth of a blackwater tributary to the main channel. (Photo credit: Dennis DeVries and Russell Wright). are the braided streams, swamps, marshes, and lakes of the Delta (Figure 13-1). The upper Delta is dominated by vegetation typical of lower coastal plain swamps (Figures 13-2, 13-3). Submerged vegetation—important as fish habitat—is dynamic, and density and coverage can vary widely through time.

Backwater areas can have highly variable productivity and physical/ chemical conditions, such as low dissolved oxygen and low pH, which can be stressful to fishes. Water chemistry and clarity in the main channels are strongly influenced by river discharge, limiting growth and development of submerged vegetation. Further downstream in the lower Delta, terrestrial and marsh vegetation shift to grasses, sedges, Figure 13-2. Cypress root habitat—a complex freshwater swamp habitat cre- and shrubs (Figures 13-4 and 13-5). ated by bald cypress knees. (Photo credit: Dennis DeVries and Russell Wright). Aquatic plants are a mixture of native

Biotic Landscape - Chapter 13 93 species, such as American eelgrass and widgeongrass, and non-native invasive species such as Eurasian watermilfoil, hydrilla, water hyacinth, and common salvinia (Table 13-1). This complex vegetative mosaic, as well as physical/chemical characteristics of the water (e.g., depth, flow, clarity, pH, dissolved oxygen), supports the diversity of fishes in the Delta.

Fishes in the Greater Mobile-Tensaw River Area The Alabama River drainage report- edly supports 184 native fishes, including 33 endemics (Boschung and Mayden 2004; Freeman et al. 2005). While the overall biota of the Figure 13-3. Cypress root habitat—a lowland swamp and marsh habitat. Note Delta has been relatively less stud- the high water marks on the cypress trunks. During high water events, this ied than other large river systems lowland forest is flooded, creating important habitat for many aquatic species. (Photo credit: Dennis DeVries and Russell Wright). (McCreadie et al. 2005), several studies of its fish assemblages do exist, most notably two recent books on the fishes of Alabama (Mettee et al. 1996; Boschung and Mayden 2004). According to Mettee et al. (1996), the Delta and freshwater tributaries to Mobile Bay contain 131 species, 30 of which are marine. While marine species may be restricted to estuarine areas for much of the year, they can make large-scale movements into freshwater; similarly, freshwater species can make large-scale movements into water with higher salinities. Clearly, this area possesses an extremely diverse and dynamic fish species assemblage (Table 13-1).

Fishes of the Delta can be broadly classified into 3 groups, as described below: freshwater and estuarine Figure 13-4. Lower delta habitat—extensive shallow water marshes with species, marine migrants, and anad- both emergent and submersed vegetation are the primary habitat type in romous and catadromous species. the lower Delta just above the causeway, at the head of Mobile Bay. (Photo credit: Dennis DeVries and Russell Wright).

94 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 1) Freshwater and Estuarine Species Freshwater fishes, those found in inland freshwater systems, are resident throughout the Delta. The particular species of freshwater fish present in an area varies with ambi- ent salinity levels and specific habitat. Species with low tolerance to elevated salinity (e.g., minnows) are limited to strictly freshwater areas, whereas those with broad salinity tolerance (e.g., many of the Lepomis sunfishes) can be found throughout the region. Similarly, tolerance to reduced dissolved oxygen can provide an advantage for species in backwater or isolated water habitats that often experience oxygen depres- sion. Fishes such as bowfin and gars Figure 13-5. D’Olive Bay—emergent grasses are present at the lowermost (Lepisosteidae) that are capable of end of the Delta, at the head of Mobile Bay. This is typical habitat in this por- aerial breathing are common in back- tion of the estuary. Interstate 10 is visible in the background. (Photo credit: Dennis DeVries and Russell Wright). waters. Large-river species such as paddlefish, Alabama sturgeon, southeastern blue sucker, smallmouth buffalo, blue catfish, and flathead catfish inhabit the main channels, as well as the reservoirs and oxbows of the river and Delta. Some of these fishes undertake extensive spawning and feeding migrations within the river and Delta systems. For example, paddlefish tend to migrate to the oxbows of the river and backwaters of the Delta as feeding habitat in summer, to Mobile Bay in the winter, and back up the river in the spring to spawn (Mettee et al. 2009). Construc- tion of dams on the lower Alabama and lower Tombigbee rivers has altered critical habitat and blocked migration routes, changes that have contributed to decline of some large- river forms and imperilment of the Figure 13-6. Millers Ferry Lock and Dam—showing the gated spillways to the Alabama sturgeon (Parauka 2004) left and the navigational lock chamber to the right. This dam (at river km 214) and Claiborne Lock and Dam (at river km 117), the two lowermost dams (Figure 13-6). on the Alabama River, are significant barriers to movement of migratory fishes in the Mobile and Alabama river system. (Photo credit: Dennis DeVries and Russell Wright).

Biotic Landscape - Chapter 13 95 northern Gulf of Mexico and along Table 13-1. List of common names and scientific names of organisms referenced the Atlantic Coast from Florida to in the text. Massachusetts. Largemouth bass are among the warmwater game- Type of Organism Scientific Name Common Name fish species most sought after by Plants Vallisneria Americana American eelgrass US freshwater anglers (US Depart- Salvinia minima common salvia ment of the Interior, US Fish and Myriophyllum spicatum Eurasian watermilfoil Wildlife Service, and US Depart- Hydrilla verticillata Hydrilla ment of Commerce, US Census Eichhornia crasspes water hyacinth Bureau 2011). This freshwater Ruppia maritima widgeongrass species is common in the reser- Taxodium distichum bald cypress voirs of the lower Alabama River, and also abundant throughout the Invertebrates Pomacea maculata Amazonian apple snail Delta. Largemouth bass beyond Corbicula fluminea Asiatic clam the early fry stage can tolerate salin- Callinectes sapidus Blue crab ity over 10 part per thousand (ppt), Daphnia lumholtzi subtropical water flea while eggs and larvae are killed by Fishes Alosa alabamae Alabama shad salinity exceeding 4 ppt (Tebo and Scaphirhynchus suttkusi Alabama sturgeon McCoy 1964). Since salinity rarely Anguilla rostrate American freshwater eel exceeds 4 ppt in the Delta during Strongylura marina Atlantic needlefish the spring spawning season, juvenile largemouth can recruit past Ictalurus furcatus blue catfish this vulnerable stage except during Lepomis macrochirus bluegill periods of extreme drought, when Amia calva bowfin salinity can exceed 4 ppt. Cyprinus carpio common carp Largemouth bass in the lower Pylodictis olivaris flathead catfish Delta exhibit adaptations to life Ctenopharyngodon idella grass carp in this environment. Rather than Brevoortia patronus Gulf menhaden growing as large as those in strictly Acipenser oxyrhynchus desotoi Gulf sturgeon freshwater environments, Delta Micropterus salmoides largemouth bass largemouth bass tend to store high Polyodon spatula paddlefish amounts of fat, consistent with Sciaenops ocellatus red drum a strategy to withstand periods of high salinity and temperature Ictiobus bubalus smallmouth buffalo during late summer (Glover et al. Cycleptus meridionalis southeastern blue sucker 2013). These coastal largemouth Paralicthys lethostigma southern flounder bass populations tend to repro- Leiostomus xanthurus spot duce about a year earlier in life Cynoscion nebulosus spotted seatrout than inland populations and have Morone saxatilis striped bass a reduced life span. Estuarine resi- Mugil cephalus striped mullet dent fishes include primarily small- bodied species, such as killifishes (Fundulidae) and silversides, well adapted to withstand the extremes Of the freshwater fishes native to the Delta, large- of living at the edge of marshes where the physical/ mouth bass is the most extensively studied. Large- chemical environment can vary dramatically. mouth bass are native to eastern North American freshwater systems from southern Canada to north- 2) Marine Migrants ern Mexico (Figure 13-7). Populations are common in oligohaline portions of estuaries throughout the Marine migrants include species that use the estuary primarily as a nursery area and species that move in

96 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area and out of the Delta to feed on abundant prey as salinity varies seasonally. Species such as southern flounder, spot, and Gulf menhaden spawn in coastal marine waters and their eggs and larvae are carried into the estuary (Figure 13-8). Juveniles may spend months to a few years in the estuaries before migrating out to the bays and near coastal waters. These larval and juvenile fishes colonize the entire lower Delta, often thriving in waters with no measurable salinity. Other marine migrants only move into the Delta and up the Mobile River when salinity peaks during late summer. Movement of spotted seatrout and red drum into the Delta support active fisheries (Figure 13-9). Other fishes may be found many kilo- Figure 13-7. Large largemouth bass—this example of a largemouth bass from meters upstream, including Atlan- the Mobile-Tensaw Delta weighs more than 2.3 kg (5 lb). Largemouth bass tic needlefish, striped mullet, and support a tremendous recreational fishery, but rarely reach this size in the several species of sharks and rays. In Delta. (Photo credit: Dennis DeVries and Russell Wright). fact, prior to construction of dams on the lower Alabama River, striped mullet were common in the Cahaba River where they contributed to local harvest and recreational fishing (Boschung and Mayden 2004).

3) Anadromous and Catadromous Species The 3rd group of fishes that inhabits this diverse ecosystem comprises the anadromous and catadromous species. Alabama shad, Gulf sturgeon, and striped bass are examples of anadromous species that migrate up the river to spawn. The only catadromous species is the Ameri- can freshwater eel, which spends the majority of its life in freshwater and migrates to the ocean to spawn. All of these highly migratory fishes have Figure 13-8. Southern flounder—collected in the Delta. This species has a experienced population declines in complex life history. Spawning in offshore coastal waters, their eggs and the Mobile drainage since construc- larvae are then carried into the coastal estuary. Juveniles can live in very low tion of dams on the lower Alabama salinities and will remain in the estuary for months to years. (Photo credit: Dennis DeVries and Russell Wright). and lower Tombigbee rivers.

Biotic Landscape - Chapter 13 97 exceeds their physiological tolerances, while allowing marine species to move further inland.

Threats to Fishes in the Lower Alabama River and Mobile-Tensaw River Delta The Delta faces a number of threats that directly influence the diversity of fishes found there. Increasing human population is a key factor that simultaneously influences many other effects. Between 1990 and 2007, Baldwin County experienced a 75% increase in its population; in Mobile County there was a 7% increase. These rates of increase are likely to continue through at least 2025 (Mobile Bay National Estuary Figure 13-9. Two red drums—sometimes called “redfish”, collected from the Program 2008). Delta. (Photo credit: Dennis DeVries and Russell Wright). 1) Human Development and Salinity Habitat Loss Development associated with human population The Delta’s diversity of fish species is due in large part increase poses substantial threats to fish populations. to the mixing in this area of freshwater, estuarine, Although occurring nearly 100 years ago, construction and marine habitats. Whereas species in inland lakes in 1926 to 1927 of a causeway across the downstream are restricted to freshwater, and species in the open end of the Delta, at the northern end of Mobile Bay, gulf or ocean are restricted to high salinity, fishes in has had a variety of impacts on the Delta, some only the Delta can choose from a variety of salinities and a recently identified. Most obvious is altered water flow variety of habitats, both of which can vary seasonally and altered water connectivity between the Delta and and across years. Mobile Bay. Rather than flowing as a sheet between The mixing of marine and fresh water in the Delta the two as in the past, water is now funneled through creates a system that is spatially and temporally four 400-m (1300-ft) wide culverts, which has altered dynamic with respect to salinity. Salt concentration is energy and nutrient transfer between these areas a critical factor limiting survival and growth of fishes. (Goecker et al. 2009) and possibly led to increased In spatial terms, salinity levels tend to be highest in sedimentation north of the causeway (Valentine et al. downstream areas closest to Mobile Bay; however, 2004; Valentine and Sklenar 2006; see also Isphording salinity also tends to be elevated further north on et al. 1996 for an alternative perspective). A further the west (Mobile River) side of the Delta due to the implication of this reduced connectivity is the poten- clockwise gyre of water flow in Mobile Bay, an effect tially reduced nursery function of the Delta (Rozas et compounded by the deepened and maintained ship- al. 2013). Given the amount of time the causeway has ping channel. Because salt water is denser than fresh been in place, fishes have clearly adapted to its pres- water, salinity also tends to be greater at depth, form- ence. But it is important to be aware of these effects, ing saline wedges along the bottom that move north- particularly as human population density and devel- ward counter to the flow of the river. Salinity also opment continue to increase. exhibits strong seasonal variations. As river discharge Another impact of increased human development is decreases during late summer, salinity in the lower change in land use and land cover in and surround- Delta increases. Elevated salinity can limit distribu- ing the Delta (Ellis et al. 2011). Among many changes tions of freshwater species, as the concentration in land cover through time, impervious surfaces have

98 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area increased, natural habitats lost or degraded, and as the threats to fishes may be less direct, making forests reduced. eventual outcomes difficult to correlate and predict. Invasive species significant to fishes in the area include Additional habitat losses have taken place in river- a number of plants (Eurasian watermilfoil, hydrilla, ine reaches upstream of the Delta due to dredging water hyacinth, common salvinia), invertebrates (the and relocation of dredge spoils. Recent collections Amazonian apple snail, Asiatic clam, a subtropical by Haley and Johnston (2014) indicate changes from water flea), and fishes (common carp, grass carp). historical baseline data, suggesting that loss of habi- Invasive plants most certainly influence habitat tat is affecting the presence of species in riverine areas available for fishes (see, for example, Crowder and (Haley 2012; Haley and Johnston 2014). They also Cooper 1982 on freshwater systems). In some cases, presented evidence for homogenization of the fish the impacts of invasive invertebrates and other fishes, assemblage across the greater Mobile-Tensaw River such as common carp, occurred long ago and are thus area in recent collections. well established. Other invasives have arrived relatively recently and their effects have not yet been quantified 2) Mercury Pollution (e.g., DeVries et al. 2006; Martin et al. 2012). Mercury deposition in the Delta has been documented in several fish species, as well as in sediments, 5) Climate Change and has been traced primarily to atmospheric input Finally, at the broadest level, climate change is a major (Bonzongo and Lyons 2004; Warner et al. 2005). driving force that is likely to exert a strong influence Consumption advisories are regularly issued for many on fishes in the rivers and Delta. Climate change fish species in the region due to mercury (Alabama models predict not only increased temperatures for Department of Public Health 2015). Methylmercury coastal and estuarine areas, such as the Delta, but also concentrations in fishes have been shown to differ a suite of other effects, such as increased rates of sea across the length of the Delta between piscivore level rise; increased intensity and frequency of coastal species (e.g., largemouth bass and southern flounder), storms and hurricanes; and changes in freshwater and between predators and prey (e.g., bluegill and inputs, sedimentation, nutrient input, and resulting Blue Crab; Farmer et al. 2010). estuarine productivity (Scavia et al. 2002; Nelson et al. 2013; Rybczyk et al. 2013). Increased temperatures 3) Tropical Storms will directly affect the metabolic rates of ectotherms. Although tropical storms are natural phenomena, But the indirect effects of changes in salinity patterns they can still represent a threat to fishes in the study (due to changes in freshwater inflows), precipitation, area. Several large hurricanes have impacted this area and storm frequency are more complex and difficult in recent years, including hurricanes Frederic in 1979, to predict. In addition, effects of climate change may Ivan in 2004, and Katrina in 2005. Their impacts on enhance the ability of invasive species to enter the fishes have not been well studied, given the difficulty in system, further complicating predictions (Rahel and sampling after such storms. However, data on Hurri- Olden 2008). Given the complexity of these systems, cane Ivan demonstrated major changes in water levels we may never accurately predict these system-wide and salinity during landfall, followed by high commu- effects. If we are to begin generating testable predic- nity metabolism in the afternoon and low dissolved tions, additional research is needed to identify those oxygen levels the following morning (Park et al. 2007), factors most important to system dynamics. all factors thought to have strong impacts on fishes. Isphording (1994; cited in Baya et al. 1998) estimated Resource Conservation Implications that Hurricane Frederic removed nearly 300 million Conservation and management of the tremendous metric tons (660 billion lb) of sediment from the Delta diversity of fishes of the Delta involves common and bay, providing a “natural cleansing” to the area. sense, as well as some significant changes in human behavior. For example, rapid development in Mobile 4) Invasive Species and Baldwin counties highlights the need to protect The impacts of a growing suite of invasive species the riparian zones in the watershed as a whole, and represent an important potential threat to fishes in the specifically in local watersheds that drain directly into study area that must be addressed. Those impacts may the Delta. Sediment has been recognized as one of the be less evident than for threats described previously

Biotic Landscape - Chapter 13 99 sufficient and well-timed freshwater input, salinity will encroach inland, compromising production and survival of sensitive species.

Increasing the connectivity of the upper portions of the Mobile Basin watershed with the Delta and with the Gulf of Mexico is important for restoration of the most imperiled species. Fish populations that have suffered the greatest decline in this system are those whose historic migration patterns have been impeded or blocked entirely by the presence of upstream dams. Provid- ing greater connection to critical habitats by facilitating fish move- Figure 13-10. Excessive sediment from erosion is considered among the most ment, using either existing lock-and- important non-point source pollutants in Alabama. This is an example of a failed attempt to control erosion at a construction site in Baldwin County. dam structures or by developing new Note the collapsed silt fences. (Photo credit: Dennis DeVries and Russell passage technologies, could help Wright). restore these once abundant populations and increase the flow of nutrients and materials in the ecosystem.

While we cannot alter large-scale effects of climate change at the regional level (e.g., sea level rise), it is possible to monitor changes in habitat and adopt better management approaches. Increased salinity further upstream, greater storm impacts, and the potential for greater colonization by tropical species are potential threats to native species. Routine monitoring to detect the presence and determine the spread of current and new invasive species is important, whether they are climate influenced or not.

Clearly the fishes of the greater Mobile-Tensaw River area are impor- Figure 13-11. Sunset on the Mobile-Tensaw River Delta. (Photo credit: Dennis tant to the ecological function of the DeVries and Russell Wright). system. Many of these populations most significant nonpoint source pollutants impact- are important commercial and recre- ing aquatic environments (Figure 13-10). In the face ational species that support economically significant of growing demands on water upstream, it is not only industries. We must take the correct management and important to maintain the quality of water entering conservation steps to preserve this diverse commu- the rivers and the Delta, but the quantity and timing nity of fishes for future generations (Figure 13-11). of critical flows must also be maintained. Without

100 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 14. Reptiles and Amphibians Jen Williams, National Park Service; JJ Apodaca, Warren Wilson College; and Craig Guyer, Auburn University

This chapter describes the conservation status of in indiscriminant killing—and their importance in reptiles and amphibians in the greater Mobile-Tensaw natural systems is underappreciated. It is unfortunate River area. We follow the lead of the US Fish and that public support for species conservation often Wildlife Service by using “species” broadly to include depends on their perceived aesthetic quality or value both species and subspecies (USFWS 2015a). As an to humans. Despite reptiles serving as important example, for Coal Skinks (Plestidon anthracinus), we components in the food web as both predator and count 2 “species”: Northern Coal Skinks (P. a. anthra- prey, as well as their ecosystem services in the form cinus) and Southern Coal Skinks (P. a. pluvialis). of rodent and insect control, they do not garner levels This approach takes full advantage of the knowledge of public support enjoyed by other taxonomic groups, gleaned from conservation genetics. such as birds or mammals.

Reptile Conservation Ironically, those who do value reptiles can sometimes be the same ones contributing to their decline. Over- Alabama contains roughly one third of the reptile collection and wildlife trafficking of herpetofauna species found in the contiguous 48 states (ADCNR significantly impact population levels. Penalties for 2014a). This reptile richness is a result of Alabama’s wildlife smuggling often do not outweigh the mone- subtropical climate, diverse geography, and extensive tary profits and other benefits received from commit- riverine system (Mittermeier et al. 2015). Lack of glaci- ting those crimes. Threats to reptiles of the greater ation during the Pleistocene also contributed to the Mobile-Tensaw River area are not substantially differ- proliferation of Alabama’s flora and fauna (ADCNR ent from those faced by reptiles worldwide. However, 2014a). Over 90% of Alabama’s lizard species, 75% of the level of habitat loss here, coupled with high species its snake species, and nearly 70% of its turtle species diversity, makes the greater Mobile-Tensaw River area are documented, likely to occur, or once occurred an important conservation priority. in the greater Mobile-Tensaw River area (Table 6; Guyer et al. 2015 and Guyer et al., forthcoming). 1) Many Species of Conservation Concern One of those species, Southern Black-knobbed Map Turtle (Graptemys nigrinoda delticola), is endemic to Twenty-three percent of reptile species found in Alabama. the greater Mobile-Tensaw River area are classified as Species of Greatest Conservation Need by the Globally, 1 in 5 species of reptile is at risk of extinc- Alabama Department of Conservation and Natural tion, with freshwater ecosystems, such as those found Resources (2014a; Table 14-1) and approximately in the greater Mobile-Tensaw River area, having one 11% are listed as Endangered or Threatened under the of the highest proportions of at-risk species (Bohm federal Endangered Species Act. The latter includes et al. 2013). Top threats to reptile conservation are the Alabama Red-bellied Cooter (Pseudemys alabam- habitat loss and harvesting, with harvest of freshwa- ensis), officially designated as Alabama’s state reptile ter and marine reptiles of particular concern (Bohm in 1990. In addition, Eastern Diamondback Rattle- et al. 2013). Reptile conservation is further burdened snakes (Crotalus adamanteus), Southern Hognose by agriculture, aquaculture, urban development, and Snakes (Heterodon simus), and Alligator Snapping pollution (Bohm et al. 2013), along with exotic inva- Turtles (Macrochelys temminckii) are classified as sive species, pollution/contaminants, disease, parasit- Under Review under the Endangered Species Act. ism, and climate change (Gibbons et al. 2000). Southern Hognose Snakes have not been documented Public perception of reptiles poses yet another chal- in Alabama since 1970 and this species is possibly lenge to their conservation. These animals are often extirpated (Guyer et al., forthcoming). Despite the vehemently feared or disliked—frequently resulting release of Eastern Indigo Snakes (Drymarchon couperi;

Biotic Landscape - Chapter 14 101 Table 14-1. Reptile and amphibian species documented (D), likely to occur (L), or previously occurring (i.e., extirpated [E]) in the greater Mobile-Tensaw River area and their federal, state, and International Union for Conservation of Nature (IUCN) classifications (See Table 14-2).

Occurrence Federal State IUCN Class (See table (See Table (See Table (See Table Order Scientific Name - Species Common Name - Species caption) 14-2) 14-2) 14-2) REPTILES Anolis carolinensis carolinensis Northern Green Anole D P5 LC Lizards Aspidoscelis sexlineata sexlineata Eastern Six-lined Racerunner D P4 LC Ophisaurus ventralis Eastern Glass Lizard D P4 LC Ophisaurus attenuatus longicaudus Eastern Slender Glass Lizard D P2 LC Plestiodon anthracinus anthracinus Northern Coal Skink D P2 LC Plestiodon anthracinus pluvialis Southern Coal Skink L P2 LC Plestiodon egregius similis Northern Mole Skink D P3 LC Plestiodon inexpectatus Southeastern Five-lined Skink D P2 LC Plestiodon laticeps Broad-headed Skink D P5 LC Plestiodon fasciatus Common Five-lined Skink D P5 LC Sceloporus undulatus Eastern Fence Lizard D P5 LC Scincella lateralis Little Brown Skink D P5 LC REPTILES Agkistrodon contortrix Copperhead D P5 LC Snakes Agkistrodon piscivorus conanti Florida Cottonmouth D P5 LC Agkistrodon piscivorus piscivorus Continental Cottonmouth D P5 LC Carphophis amoenus helenae Midwestern Wormsnake L P5 LC Cemophora coccinea copei Northern Scarletsnake D P4 LC Coluber flagellum flagellum Eastern Coachwhip D P3, SP NE Coluber constrictor priapus Southern Black Racer D P4 LC Eastern Diamond-backed Crotalus adamanteus Rattlesnake D UR P2 LC Crotalus horridus Timber Rattlesnake D P5 LC Diadophis punctatus stictogenys Mississippi Ring-necked Snake D P5 LC Drymarchon couperi Eastern Indigo Snake E LT P1 LC Farancia erytrogramma Rainbow Snake D P1 LC Farancia abacura abacura Eastern Mudsnake L P4 LC Farancia abacura abacura reinwardtii Western Mudsnake D P4 LC Heterodon simus Southern Hog-nosed Snake E UR P1 VU Heterodon platirhinos Eastern Hog-nosed Snake D P3 LC Lampropeltis getula Eastern Kingsnake D P2 LC Lampropeltis holbrooki Speckled Kingsnake D P2 NE Lampropeltis elapsoides Scarlet Kingsnake D P4 NE Lampropeltis calligaster rhombomaculata Mole Kingsnake D P3 LC Micrurus fulvius Eastern Coralsnake D P1 LC Nerodia erythrogaster Plain-bellied Watersnake D p5 LC Nerodia cyclopion Mississippi Green Watersnake D P4 LC Nerodia floridana Florida Green Watersnake D P3 LC Nerodia clarkii Saltmarsh Watersnake D P2 LC Nerodia taxispilota Brown Watersnake D P4 LC Nerodia rhombifer Diamond-backed Watersnake D P4 LC

102 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Table 14-1. Reptile and amphibian species documented (D), likely to occur (L), or previously occurring (i.e., extirpated [E]) in the greater Mobile-Tensaw River area and their federal, state, and International Union for Conservation of Nature (IUCN) classifications (See Table 14-2), cont.

Occurrence Federal Class (See table (See Table State (See IUCN (See Order Scientific Name - Species Common Name - Species caption) 14-2) Table 14-2) Table 14-2) REPTILES Nerodia fasciata fasciata Banded Watersnake D P5 LC Snakes Nerodia fasciata confluens Broad-banded Watersnake D P5 LC (cont.) (Nerodia sipedon pleuralis Midland Watersnake D P5 LC Opheodrys aestivus aestivus Northern Rough Greensnake D P5 LC Pantherophis guttatus Red Cornsnake D P3 LC Pantherophis spiloides Gray Ratsnake D P5 NE Pituophis melanoleucus lodingi Black Pinesnake D LT P1 LC Pituophis melanoleucus mugitus Florida Pinesnake L P2 LC Regina rigida sinicola Gulf Crayfish Snake D P5 LC Rhadinaea flavilata Pine Woods Littersnake D P3 LC Storeria occipitomaculata occitomaculata Northern Red-bellied Snake D P5 LC Storeria dekayi wrightorum Midland Brownsnake D P5 LC Storeria dekayi limnetes Marsh Brownsnake D P5 LC Sistrurus miliarius barbouri Dusky Pygmy Rattlesnake D P3 LC Tantilla coronata Southeastern Crowned Snake D P4 LC Thamnophis sirtalis sirtalis Eastern Gartersnake D P5 LC Thamnophis sauritus sauritus Common Ribbonsnake D P4 LC Virginia striatula Rough Earthsnake D P5 LC Virginia valeriae valeriae Eastern Smooth Earthsnake D P5 LC Virginia valeriae elegans Western Smooth Earthsnake D P3 LC REPTILES Apalone mutica calvata Gulf Coast Smooth Softshell L P3 LC TURTLES Apalone ferox Florida Softshell L P3 LC Apalone spinifera aspera Gulf Coast Spiny Softshell D P4 LC Caretta caretta Loggerhead Sea Turtle D LT P1 E Chelonia mydas Green Sea Turtle D LT P1 E Chelydra serpentina Snapping Turtle D P4 LC Chrysemys dorsalis Southern Painted Turtle D P5 NE Deirochelys reticularia Chicken Turtle L P3 NE Dermochelys coriacea Leatherback Sea Turtle D LE P1 LC Gopherus polyphemus Gopher Tortoise D LT, C1 P2 VU Graptemys nigrinoda nigrinoda Northern Black-knobbed Turtle D P3, SP LC Graptemys nigrinoda delticola Southern Black-knobbed Turtle D P3, SP LC Graptemys ouachitensis ouachitensis Ouachita Map Turtle D P3 LC Graptemys pulchra Alabama Map Turtle D P3, SP NT Kinosternon subrubrum subrubrum Southeastern Mud Turtle D P5 LC Kinosternon subrubrum hippocrepis Mississippi Mud Turtle D P5 LC Lepidochelys kempii Kemp’s Ridley Sea Turtle D LE P1 VU Macrochelys temminckii Alligator Snapping Turtle D UR P3, SP VU Pseudemys alabamensis Alabama Red-bellied Cooter D LE P1 E Pseudemys concinna concinna Eastern River Cooter D P5 LC

Biotic Landscape - Chapter 14 103 Table 14-1. Reptile and amphibian species documented (D), likely to occur (L), or previously occurring (i.e., extirpated [E]) in the greater Mobile-Tensaw River area and their federal, state, and International Union for Conservation of Nature (IUCN) classifications (See Table 14-2), cont.

Occurrence Federal State IUCN Class (see table (See Table See Table (See Table Order Scientific Name - Species Common Name - Species caption) 14-2) 14-2) 14-2) REPTILES Turtles Pseudemys concinna floridana Coastal Plain Cooter D P5 LC (cont.) Sternotherus minor peltifer Stripe-necked Musk Turtle D P4 LC Sternotherus odoratus Eastern Musk Turtle D P4 LC Terrapene bauri Florida Box Turtle D P4 NE Terrapene carolina Eastern Box Turtle D P4 VU Trachemys scripta scripta Yellow-bellied Slider D P5 LC Trachemys scripta elegans Red-eared Slider D P5 LC REPTILES Crocodilians Alligator mississippiensis American Alligator D LT(SA) P5 AMPHIBIANS Reticulated flatwoods Salamanders Ambystoma bishopi salamander E LE P1 VU Ambystoma maculatum Spotted Salamander D P4 LC Ambystoma opacum Marbled Salamander D P4 LC Ambystoma talpoideum Mole Salamander D P4 LC Ambystoma texanum Small-mouthed Salamander D P2 LC Ambystoma tigrinum Tiger Salamander D P2 LC Amphiuma means Two-toed Amphiuma D P4 LC Amphiuma pholeter One-toed Amphiuma D P2 NT Amphiuma tridactylum Three-toed Amphiuma D P4 LC Desmognathus auriculatus Southern Dusky Salamander D P1 LC Desmognathus conanti Spotted Dusky Salamander D P5 LC Desmognathus monticola Seal Salamander D P5 LC Eurycea cf. chamberlani (panhandle form) Unnamed Species D P3 DD Eurycea cirrigera Southern Two-lined Salamander D P5 LC Eurycea guttolineata Three-lined Salamander D P5 LC Eurycea longicauda Long-tailed Salamander D P5 LC Eurycea quadridigitata Dwarf Salamander D P3 LC Hemidactylium scutatum Four-toed Salamander D P4 LC Necturus cf. beyeri Gulf Coast Waterdog D P3 DD Necturus lodingi Unnamed Waterdog D P3 DD Notophthalmus viridescens louisianensis Red-spotted Newt D P5 LC Phaeognathus hubrichti Red Hills Salamander D LT P1 LC Plethodon grobmani Southeastern Slimy Salamander D P5 LC Plethodon mississippi Mississippi Slimy Salamander D P5 LC Plethodon websteri Webster’s Salamander D P5 LC Pseudotriton montanus flavissimus Gulf Coast Mud Salamander D P3 LC Pseudotriton ruber vioscai Southern Red Salamander D P4 LC Siren intermedia intermedia Eastern Lesser Siren D P5 LC Siren intermedia nettingi Western Lesser Siren D P3 LC Siren lacertina parda Greater Siren D P3 LC

104 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Table 14-1. Reptile and amphibian species documented (D), likely to occur (L), or previously occurring (i.e., extirpated [E]) in the greater Mobile-Tensaw River area and their federal, state, and International Union for Conservation of Nature (IUCN) classifications (See Table 14-2), cont.

Occurrence Federal State IUCN Class (see table (See Table (See Table (See Table Order Scientific Name - Species Common Name - Species caption) 14-2) 14-2) 14-2) AMPHIBIANS Acris crepitans Northern Cricket Frog D P5 LC Frogs Acris gryllus Southern Cricket Frog D P5 LC Anaxyrus quercicus Oak Toad D P3 LC Anaxyrus fowleri Fowler’s Toad D P5 LC Anaxyrus terrestris Southern Toad D P5 LC Gastrophryne carolinensis Eastern Narrow-mouthed Toad D P5 LC Hyla avivoca Bird-voiced Treefrog D P5 LC Hyla chrysoscelis Cope’s Gray Treefrog D P5 LC Hyla cinerea Green Treefrog D P5 LC Hyla femoralis Pine Woods Treefrog D P5 LC Hyla gratiosa Barking Treefrog D P4 LC Hyla squirella Squirrel Treefrog D P5 LC Lithobates capito Gopher Frog D P1 NT Lithobates catesbeianus Bullfrog D P5 LC Lithobates clamitans Green Frog/ Bronze Frog D P5 LC Lithobates grylio Pig Frog D P4 LC Lithobates heckscheri River Frog D P1 LC Lithobates palustris Pickerel Frog D P4 LC Lithobates sevosus Mississippi Gopher Frog E P1 CE Lithobates spheocephalus Leopard Frog D P5 LC Pseudacris crucifer Spring Peeper D P5 LC Pseudacris feriarum Upland Chorus Frog D P5 LC Pseudacris nigrita Southern Chorus Frog D P5 LC Pseudacris ornata Ornate Chorus Frog D P3 LC Scaphiopus holbrooki Eastern Spadefoot D P4 LC

Biotic Landscape - Chapter 14 105 Table 14-2. Federal, State, and International Union for Conservation of Nature (IUCN) classification definitions for reptiles and amphibians.

Level Rank Definition Federal (USFWS LE Listed Endangered—a species in danger of extinction throughout all or a significant portion of its range. 2012b, 2015a) LT Listed Threatened—a species likely to become endangered within the foreseeable future throughout all or a significant portion of its range. (SA = due to similarity of appearance.) C Candidate—a species under consideration for official listing for which there is sufficient information to support listing. UR Under Review—species that have been petitioned for listing and for which a 90-day finding has not been published or for which a 90-day substantial has been published but a 12-month finding have not yet been published in the Federal Register. Also includes species that are being reviewed through the candidate process, but the Candidate Notice of Review has not yet been signed. State (ADCNR 2014a, P1 Priority 1/Highest Conservation Concern—species critically imperiled and at risk of extinction/extirpation 2014b and 2015) because of extreme rarity, restricted distribution, decreasing population trend/population viability problems, and specialized habitat needs/habitat vulnerability. Immediate research and/or conservation action required. P2 Priority 2/High Conservation Concern—species imperiled because of 3 of 4 of the following: rarity; very limited, disjunct, or peripheral distribution; decreasing population trend/population viability problems; specialized habitat needs/habitat vulnerability. Timely research and/or conservation action needed. P3 Priority 3/Moderate Conservation Concern—species with conservation problems because of insufficient data or because of 2 of 4 of the following: small populations; limited, disjunct, or peripheral distribution; decreasing population trend/ population viability problems; specialized habitat needs/habitat vulnerability due to natural/ human-caused factors. Research and/or conservation action recommended. P4 Priority 4/Low Conservation Concern—species that are secure, yet conservation concerns exist because of 1 of 4 of the following: relative abundance; limited, disjunct, or peripheral distribution; decreasing population trend/population viability problems; specialized habitat needs/increasing habitat vulnerability due to natural/ human-caused factors. Research on specific problem suggested. P5 Priority 5/Lowest Conservation Concern—species that are demonstrably secure, with size of population stable/ increasing, geographical distribution stable/expanding, population trend/population viability stable/increasing, relatively limited habitat vulnerability due to natural/ human caused factors, or an unusual visitor to the state. No specific monitoring or conservation action needed. SP Species Protected—species protected by Regulation 220-2-.92 (Nongame Species Regulations), which prohibits the take or attempt to take, capture, kill, possession, sale, trade or offer for anything of monetary value of nongame wildlife species or any parts thereof listed in the regulation, unless allowed by permit. IUCN (2012 and CE Critically Endangered—a taxon that meets any of the criteria for Critically Endangered and is facing an 2015) extremely high risk of extinction in the wild. DD Data Deficient—a taxon is data deficient when there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status. E Endangered—a taxon that meets any of the criteria for Endangered and is facing a very high risk of extinction in the wild. LC Least Concern—a taxon that has been evaluated against the criteria and does not qualify for Critically Endangered, Endangered, Vulnerable, or Near Threatened. Widespread and abundant taxa are included in this category. NE Not Evaluated—a taxon that has not yet been evaluated against the criteria. NT Nearly Threatened—a taxa that does not qualify for Critically Endangered, Endangered, or Vulnerable now, but is close to qualifying for or is likely to qualify for a threatened category in the near future. VU Vulnerable—a taxon meets any of the criteria for Vulnerable and that is facing a high risk of extinction in the wild.

106 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Federally Threatened) at 9 locations in Alabama in the conservation. Alternative ways to promote Eastern 1970s and 1980s (Speake and Smith 1987; Speake et al. Diamondback Rattlesnake conservation to the public 1987), recent surveys produced no confirmed sight- would be to display captive-raised individuals. While ings of this species (Hart 2002) and it also is believed Opp is not in the greater Mobile-Tensaw River area, its to be extirpated in the state. However, in 2010 and close proximity makes it likely that many of the Eastern 2011 Eastern Indigo Snakes were released in Cone- Diamondback Rattlesnakes over the years have come cuh National Forest in Covington County, a county from the study area. Simple conservation measures, that neighbors Mobile County to the east. The hope such as limiting or prohibiting harvest of this species is that a breeding population will become established from the wild, can go a long way to conserve Eastern in the area (Godwin et al. 2011). The greater Mobile- Diamondback Rattlesnakes in the greater Mobile- Tensaw River area contains unique and high quality Tensaw River area and throughout the state. habitat in the Perdido River Longleaf Hills Tract (a state-owned preserve) in Baldwin County that could 3) Turtles well serve both Eastern Indigo Snakes and Southern Twenty years ago, Lydeard and Mayden (1995) wrote Hognose Snakes for reintroduction efforts (Guyer et a paper exalting the biodiversity of Alabama, giving al., forthcoming). particular attention to turtles. Alabama boasts remarkable turtle diversity (Table 14-1). It claims over half of 2) Rattlesnakes US freshwater turtle species (Lydeard and Mayden The longest-running rattlesnake roundup (1960 to 1995) and, in global analyses, ranks second in turtle present) occurs annually in the City of Opp, Alabama species richness (Buhlmann et al. 2009; Mittermeier (City of Opp 2015), where members of the public 2015). provide the snakes used each year at the festival. In 2015, 88 Eastern Diamondback Rattlesnakes were Lydeard and Mayden (1995) noted that insufficient on display (Elofson 2015). The rodeo is a 2-day event protections were in place to conserve the biota of that includes handling and racing Eastern Diamond- Alabama and urged “recognition, protection, and back Rattlesnakes and selling Eastern Diamondback recovery of critical habitat, particularly for hot spots Rattlesnake meat at concession stands. No snakes are such as the aquatic systems of Alabama.” The State killed at the event. The city’s Planning Director said of Alabama provided a conservation mechanism for the rodeo generated considerable revenue from the turtles with enactment in 2012 of a prohibition on the more than 20,000 people in attendance—more than commercial harvest of wild freshwater turtles, their triple the current population of Opp—during the eggs, and turtle parts on both public and private lands. 2012 festivities (Johnson and Wallace-King 2012). For Turtle harvest in Alabama and other states in the US decades, gassing of burrows was the principal method spiked in recent years due to the market for turtle used to collect large numbers of Eastern Diamond- meat in Asia, where native turtle populations have back Rattlesnakes for purposes such as rattlesnake plummeted due to unsustainable harvest. Alabama rodeos. This practice was made unlawful in 2009, but Department of Conservation and Natural Resources Guyer et al. (2015) fear decades of use may have left an Administrative Code Chapter 220-2, however, allows “environmental legacy . . . across southern Alabama,” up to 2 turtles per day to be taken from the wild for referring to the time that will be required to recover personal use. from this overharvest. Administrative Code Chapter 220-2 also allows turtle Eastern Diamondback Rattlesnakes are a Priority 2 farming, provided the proper permits are acquired species (Table 14-1), of high conservation concern in and turtles from the wild are not used in propagation. the State of Alabama, and, as such, their take should be Currently, there are 6 permitted turtle farmers in the prohibited along with Priority 1 species and all other state (Mark Sasser, biologist for the Alabama Divi- Priority 2 species. However, they are not afforded the sion of Wildlife and Freshwater Fisheries, personal same protection due to political resistance and atti- communication, unreferenced), with snapping turtles, tudes of the general public (Guyer et al. forthcoming). sliders, and softshells being common exports. In fact, In fact, unlimited harvest is allowed. unlimited harvest is allowed.

The City of Opp Rattlesnake Rodeo is endorsed as a family-friendly event that promotes rattlesnake

Biotic Landscape - Chapter 14 107 4) American Alligators The State of Alabama contains at least 73 species of In addition to these steps taken for turtle conserva- amphibians, with a handful of additional possibili- tion, Alabama showed leadership in conservation of ties (roughly one third of US amphibian diversity). American Alligators (Alligator mississippiensis). When The greater Mobile-Tensaw River area contains an this species was at the brink of extinction earlier in the astounding 53 species (excluding 2 extirpated species; 20th century, Alabama was the first state to put in place Table 14-1). While other regions in North America protective measures. From 1938 until 2006, nearly may boast higher total numbers of amphibian species 70 years, alligator hunting was banned in the state due to narrow endemics (e.g., the Southern Appa- (Felsher 2014). Today sustainable alligator hunting is lachians), the greater Mobile-Tensaw River area allowed, which boosts Alabama’s economy through includes sites with the highest total amphibian rich- the sale of licenses and other associated hunting costs. ness on the continent (Apodaca 2010; Jenkins et al. 2015). 5) Changing Public Attitudes This incredible richness is due to a coalescence of th The 20 century saw a shift in public attitudes toward historical influences and more recent habitat avail- predators, from viewing them as vermin to an appre- ability. Environmental niche modeling has revealed ciation of their value as components of ecosystems, that the climate in this area is ideal for a high number particularly their role as managers of prey populations. of amphibians (Apodaca 2010), and predicted climate Herpetologists have hoped for similar shifts in atti- changes do not appear to substantially alter the tudes toward reptiles and such changes have already projected species composition (Apodaca, unpub- begun. For example, the State of Georgia used to host lished data, unreferenced). Additionally, the Alabama 3 rattlesnake rodeos, but the Fitzgerald Rattlesnake River drains more than 110,000 km2 (42,000 mi2)— Roundup was rebranded as the Fitzgerald Chicken including everything from 1st-order streams to expan- Festival and the rattlesnake roundup in Claxton, sive floodplains—providing both the various habitats which ran for more than 4 decades, became instead required to host such a vast number of native amphib- a wildlife festival in 2012. If the words of Lydeard ians and a barrier large enough to promote vicariance. and Mayden are further heeded, rigorous conserva- In fact, the greater Mobile-Tensaw River area lies tion efforts in the greater Mobile-Tensaw River area within the most prominent “suture zone,” a hot spot can be similarly effective in protecting species like the for phylogeographic breaks, for amphibians in North Eastern Diamondback Rattlesnake, restoring species America (Rissler and Smith 2010). At least 3 pairs of such as Eastern Indigo Snakes and Southern Hognose amphibian sister species likely speciated entirely due Snakes, and further spotlighting Alabama as a leader to the Alabama River system. in reptile conservation. For several other species, the river represents the east- Amphibian Conservation ern or western fronts of their range (Soltis et al. 2006). Just as the greater Mobile-Tensaw River area is of high From a conservation standpoint, this historical anom- value for reptile conservation, so it is for amphibian aly offers the possibility to preserve a high amount of conservation. Given that amphibians are one of the amphibian diversity in a relatively small geographic most threatened vertebrate groups in the world and area. Given that several amphibian species have signif- are declining at an alarming rate (Stuart et al. 2004; icant phylogeographic breaks at or near the greater IUCN 2008), identifying and protecting areas of high Mobile-Tensaw River area, it is also an opportunity species richness and endemism is the most efficient to preserve a significant amount of genetic diversity, approach to stemming the tide of global amphibian thus giving them the necessary evolutionary capac- declines (Margules and Pressey 2000; Sarkar et al. ity to respond to ongoing and future threats, such as 2006; Apodaca 2010). The southeastern United States chytrid fungi (Batrachochytrium dendrobatidis and B. provides the best opportunity to pursue this approach salamandrivorans) and global climate change. in North America. The region is a world hot spot for amphibians (Duellman 1999; Apodaca 2010; Jenkins 1) Red Hills Salamander 2015), with over 140 described species, various evolu- The greater Mobile-Tensaw River area contains several tionarily significant lineages (Rissler and Smith 2010), globally imperiled and declining species, including the and 23 genera, including a high number of endemics. Federally Threatened Red Hills Salamander (Phaeog- nathus hubrichti) and 7 state-listed species (Table

108 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 14-1). Perhaps the most charismatic member of this exposes them to the perils of small populations and group is the highly imperiled Red Hills Salamander. eventually the extinction vortex (Frankham 2010). This unique species, the only member of its genus, is Accordingly, any successful amphibian conservation endemic to a narrow band within 6 counties in south plan for the greater Mobile-Tensaw River area should Alabama. In fact, it is the only terrestrial vertebrate consider conservation corridors, as well as large completely endemic to the State of Alabama. In 2000, portions of ideal habitat. the Red Hills Salamander was officially designated as Alabama’s state amphibian. The majority of known 4) Economic and Cultural Value of populations of Red Hills Salamanders reside near the Amphibians eastern bank of the Alabama River. Consequently, It is easy to overlook the economic and cultural value preservation efforts in the study area provide the best of amphibians. However, in many parts of the south- chance to conserve habitat for the long-term protec- eastern US, and certainly in many ecosystems within tion of this species (Apodaca 2010). the greater Mobile-Tensaw River area, amphibians are the most abundant vertebrates (Burton and Likens 2) Mississippi Gopher Frogs and Reticulated 1975; Jaeger 1979; Hairston 1987; Welsh and Droege Flatwoods Salamanders 2001). This abundance makes them exceptionally Protected habitats in the greater Mobile-Tensaw River important components of the ecosystem and corner- area would also provide an excellent opportunity to stones of many food webs and communities (Hairston reintroduce 2 of the most endangered amphibians 1987; Welsh and Droege 2001). This value is especially in the world—Mississippi Gopher Frogs (Lithobates true in wetland and floodplain communities, which sevosus) and Reticulated Flatwoods Salamanders play a disproportionately important role in the health (Ambystoma bishopi)—both of which are known from and functioning of ecosystems (Gibbons et al. 2006). only a handful of localities in the wild and are both historically found in the greater Mobile-Tensaw River Economically and culturally, amphibians play a signif- area. Mississippi Gopher Frogs are currently found icant role in fully functioning wetland ecosystems, at only one pond in southern Mississippi, and in any particularly among those ecosystem services consid- given year may have less than 100 surviving adults ered most valuable: biodiversity support, water qual- (Richter et al. 2003). Reticulated Flatwoods Salaman- ity improvement, and carbon management (Zedler ders are found in only a small portion of their original and Kercher 2005; Hocking and Babbit 2014). The range and the number of relatively healthy populations economic value of such services throughout the is likely in the single digits. Both of these amphibian greater Mobile-Tensaw River area undoubtedly species have active captive breeding programs, and reaches into the tens of millions of dollars per year. would be excellent candidates for reintroduction activities in the greater Mobile-Tensaw River area. Thwarting Extinction Such efforts are vitally important to the continued The US has witnessed the extinction of numerous survival of these species. species, lineages, and unique species assemblages over the past centuries. Without drastic actions, 3) Threats to Amphibians extinction rates may climb sharply at the end of this The greater Mobile-Tensaw River area is certainly century (Pimm et al. 1995; Mace et al. 2005). Alabama not immune to the myriad threats to biodiversity in has the dubious distinction of leading the continental the Southeast. Similar to the threats facing reptiles in US in number of extinctions (Stein 2002), largely due the greater Mobile-Tensaw River area, chief threats to human-caused impacts to its vast network of rivers to amphibians here include habitat loss, conversion, and accompanying habitats. Additional protection and fragmentation. The 3 aforementioned imperiled in the greater Mobile-Tensaw River area would be a amphibian species are glaring examples of the effects tremendous first step in a quest to thwart the continu- that habitat loss can have on the region’s amphib- ing loss of biodiversity in the southeastern United ians. The limited dispersal ability of herpetofauna States, and would serve as a vital component of global previously mentioned makes many species especially efforts to stem amphibian biodiversity loss worldwide. susceptible to habitat loss and fragmentation, which

Biotic Landscape - Chapter 14 109

15. Avifauna of the Lower Alabama River and Mobile-Tensaw Delta Joel A. Borden and C. Smoot Major, University of South Alabama

Birds are the most abundant and diverse terrestrial and rivers with high degrees of variation in intensity vertebrate group on Earth. At over 10,000 species, and periodicity (US Army Corps of Engineers 2015). and counting, they comprise a third of all terrestrial Variations such as these have been linked to patterns vertebrate species (Pough et al. 2013). Alabama has in primary productivity in other dynamic river delta over 400 species of birds that are either full-time resi- systems (Junk et al. 1989; Høberg et al. 2002; Tockner dents or part-time migrants (Alabama Ornithological and Stanford 2002; Lindholm et al. 2007). Society 2015). This chapter reviews the avifauna of the greater Mobile-Tensaw River area, beginning with As part of the Eastern Gulf Coastal Plain ecoregion, a review of species present, those at risk, and those the greater Mobile-Tensaw River area is a portion of who migrate through, and closing with a discussion of 1 of 6 “biodiversity hot spots” in the United States conservation implications. (Chaplin et al. 2000). Many taxa endemic to this ecoregion are restricted to cypress-dominated and Significance to Avifauna of the pine-dominated wetlands and uplands, which char- Greater Mobile-Tensaw River Area acterize much of lower Alabama. Pine-dominated wetlands and uplands, which are now reduced to less The greater Mobile-Tensaw River area, including than 5% of their original coverage, have become two the Mobile-Tensaw Delta, supports high biodiversity of the most endangered ecosystems in North America (over 200 species) and provides critical habitat for (Noss and Peters 1995). Thus, in keeping with state- avian populations across the Eastern Gulf Coastal wide declines in biodiversity, the Eastern Gulf Coastal Plain and much of the Eastern Deciduous Forest Plain is also considered to be a “species endangerment ecoregions. This area, from the Red Hills in Monroe hot spot” (Flather et al. 1998). County to the vast expanse of braided streams, bottomland hardwood forests, and coastal marshes that straddle Baldwin and Mobile counties, is an often harsh and ever-changing environment for plants and animals that call this watershed home.

Habitat diversity, environmental patchiness, and natural- and human- induced disturbance define this highly dynamic ecosystem. The ecosystem plays a crucial role as an avian breeding ground, plus serves as a migratory stopover for refuge and refueling and as a major flyway of the Central Gulf Coastal Region (Orme et al. 2005). In particular, the annual flood cycle is a key component and ecological driver of biodiversity in the region. Flooding is common; it occurs primarily during winter and early summer prolonged rain events that inundate small streams Figure 15-1. Purple Gallinule hatchling. (Photo credit: Kathy Hicks).

Biotic Landscape - Chapter 15 111 subject to considerable disturbances, both human-induced (e.g., eutrophication, development, industrial pollution) and natural (e.g., tropical storms, drought, flooding events). In fact, the high biodiversity of this region seems largely driven by disturbance.

Species Overview The greater Mobile-Tensaw River area plays a major role in the function of this section of the Gulf Coast, which is distinguished by some of the world’s most productive estuarine and coastal marine communities, providing habitat for waterfowl, wading birds, migra- Figure 15-2. Flight of Blue-Winged Teal. (Photo credit: Kathy Hicks). tory songbirds, birds of prey, and both game and non-game species. Notable species within the avian community are the marsh (wetland grasses) birds, such as the Clapper, King, Virginia, Black, and Yellow Rails; Common and Purple Gallinules (Figure 15-1); the American Coot; Red-winged Black- bird and Boat-tailed Grackle. These species require large expanses of marsh that provide cover for predator avoidance and nesting, and habitat for prey (Dunn and Alderfer 2011).

Huge annual flights of waterfowl also utilize marsh and forested backwaters of the greater Mobile-Tensaw River area during migration and as overwintering sanctuaries. These species include Green-winged and Blue-winged Teal (Figure 15-2), Mallard, Gadwall, American Wigeon, Northern Pintail, North- Figure 15-3. Male Wood Duck. (Photo credit: Kathy Hicks). ern Shoveler, Canvasback, Redhead, Ring-necked Duck, Wood Duck

(Figure 15-3), Greater and Lesser Scaup, Buffle- Moreover, the Mobile-Tensaw deltaic system is of head, and Hooded and Red-breasted Merganser evolutionary significance. As reviewed by Soltis et al. (Dunn and Alderfer 2011). (2006), the Delta and surrounding regions constitute a major phylogeographic break, which has acted as Perhaps, the most conspicuous group is the long- a significant barrier to gene flow resulting in distinct legged wading birds. These species lurk stealthily evolutionary lineages on either side of the Delta. This around the edges of river channels and across vast mud “hybridization” of community types, in part, explains flats looking for prey as varied as insects, fish, frogs, the high degree of endemic species and biodiver- snakes, hatchling alligators, and rodents. Included sity observed in southern Alabama. The Delta is also in this group are the Great Blue Heron, Great Egret,

112 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Snowy Egret, Tricolored Heron, Little Blue Heron, Reddish Egret (Figure 15-4), Black-crowned and Yellow- crowned Night-Heron, Green Heron (Figure 15-5), and Least Bittern (Gill 2007; Dunn and Alderfer 2011).

Another group reliant on this region are migratory songbirds that inhabit bottomland deciduous forests. Among these are warblers (Protho- notary [“swamp canary”] (Figure 15-6), Yellow-throated, Prairie, Northern Parula, Yellow-rumped, Black-and-white (Figure 15-7), American Redstart, Northern Water- thrush, Common Yellowthroat, and Kentucky, to name just a few); vireos (Red-eyed, White-eyed, Yellow- throated, and Blue-headed), wrens (Carolina, House, Winter, Sedge, and Figure 15-4. Reddish Egret. (Photo credit: Kathy Hicks). Marsh), Carolina Chickadee (Figure 15-8), Tufted Titmouse, and flycatchers (Eastern Phoebe, Eastern King- bird, Eastern Wood-Pewee, Acadian, Great-crested, and Least) (Dunn and Alderfer 2011). These species are ecologically important to the greater Mobile-Tensaw River area because they “repackage” energy, in the form of small arthropods, into a larger meal for raptors, small mammals, snakes, and other predators (Gill 2007; Pough et al. 2013).

Upland species inhabiting pine savannas and oak-hickory forests also play a crucial role in the ecological stability of this region (Gill 2007). The sparrows (Chipping, Savannah, Field, Grasshopper, Henslow’s, Song, White-crowned [Figure 15-9], and White-throated) disperse seeds and fall prey to small mammals, as well as Figure 15-5. Green Heron. (Photo credit: Kathy Hicks). to Cooper’s Hawks, Sharp-shinned Hawks, American Kestrels, and Merlins (Gill 2007; Dunn and Alderfer 2011). effect on its environment relative to its population size. If a keystone species is removed from its habitat, Primary cavity nesters—the woodpeckers—are criti- many other species would likely decline or disappear cal as “keystone species” within the greater Mobile- altogether (Pough et al. 2013). These diligent excava- Tensaw River area. The term keystone species is tors provide relatively safe and secure nesting oppor- defined as a species that has a disproportionately large tunities for not only their offspring, but other species’

Biotic Landscape - Chapter 15 113 Figure 15-6. Prothonotary Warbler. (Photo credit: Kathy Hicks).

Figure 15-7. Black-and-White Warbler. (Photo credit: Kathy Hicks).

114 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figure 15-8. Carolina Chickadee. (Photo credit: Kathy Figure 15-9. White-Crowned Sparrow. (Photo credit: Hicks). Kathy Hicks).

Figure 15-10. Pileated Woodpecker. (Photo credit: Kathy Hicks).

Biotic Landscape - Chapter 15 115 Kestrels, Tree Swallows, Carolina Chickadees, Eastern Screech-Owls, Prothonotary Warblers, and Brown- headed Nuthatches use abandoned woodpecker nests to raise their young. Many of these secondary cavity nesters are dependent on the 6 species (Northern Flicker, Pileated [Figure 15-10], Red-headed, Red-bellied, Downy, and Hairy) of woodpeckers for nesting sites as these are often the most secure sites within these habitats for raising their young (Gill 2007).

At the top of the “food chain” are the apex predators. These are also known as birds of prey or simply raptors. The greater Mobile-Tensaw River area has many species of raptors that play a significant role in reducing the size of Figure 15-11. Osprey. (Photo credit: Kathy Hicks). prey populations, as well as increasing the health of those species by removing weaker individuals. These include the Bald Eagle, Red-tailed hawk, Red- shouldered hawk, Northern Harrier, Osprey (Figure 15-11), Horned Owl, Barred Owl (Figure 15-12), Missis- sippi Kite, Swallow-tailed Kite (Figure 15-13), and Peregrine Falcon. These birds feed on an incredible assortment of prey, from insects like dragonflies to fishes, snakes, songbirds, waterfowl, and small mammals. As in all ecosystems, top predators are important as a “top down” control that prevents prey species from destroying habitat resources for other species (Steinmetz et al. 2003).

Species of Special Concern Of the 19 species listed under “High Conservation Concern” by Mirarchi Figure 15-12. Barred Owl. (Photo credit: Joel Borden). et al. (2004a), 17 utilize the Mobile- Tensaw Delta on a seasonal basis, if not year-round. Conservation biologists offspring as well (Martin et al. 2004). They also help use several parameters to designate a species’ current disperse fungal infection throughout forested habitats conservation status (Table 15-1). NatureServe (2015) which aids in softening dead standing timber (Jack- documents conservation status at 3 geographic scales: son and Jackson 2004). This softened wood is prized global (G), national (N), and state (S), with severity of by many other species as prime nesting habitat (Gill concern indicated from 1 to 5, G1 being the rarest to 2007). Species as varied as Wood Ducks, American G5 the most secure, for example. State-ranked species

116 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area are further characterized as breeding (B) or non-breeding (N). Lastly, the International Union for the Conserva- tion of Nature Red List rates species of “Least Concern” (LC) and “Near Threatened” (NT) (IUCN 2012; Table 15-2).

Nine species considered of “High Conservation Concern” in Alabama (Table 15-3) are highly dependent on wetland habitats: Least Bittern, Reddish Egret, American Black Duck, Swallow-tailed Kite, Northern Harrier, Black and Yellow Rails, White Ibis (Figure 15-14), and American Oyster- catcher. The other 9 species of high concern (Table 15-4) prefer upland habitats adjacent to swamps and bottomlands of the Mobile-Tensaw Delta. These include the southeastern Figure 15-13. Swallow-tailed Kite. (Photo credit: Kathy Hicks). subspecies of American Kestrel, Amer- ican Woodcock, Short-Eared Owl, Wood Thrush, Worm-Eating Warb- ler, Swainson’s Warbler, Kentucky

Table 15-1. Ranking species is conducted by NatureServe on Global (G rank), National (N rank), and a State rank (S rank) and are considered very reliable by most scientists, government agencies, and conservationists (NatureServe 2015).

Rank Ranking Criteria Definition for NatureServe GX Presumed Extinct (species)/Eliminated (ecological communities and systems) — Species not located despite intensive searches NX and virtually no likelihood of rediscovery. Ecological community or system eliminated throughout its range, with no restoration SX potential. GH Possibly Extinct (species)/ Eliminated (ecological communities and systems) — Known from only historical occurrences but still some NH hope of rediscovery. There is evidence that the species may be extinct or the ecosystem may be eliminated throughout its range, SH but not enough to state this with certainty. G! Critically Imperiled—At very high risk of extinction due to extreme rarity (often 5 or fewer populations), very steep declines, or N1 other factors. S1

G2 Imperiled—At high risk of extinction or elimination due to very restricted range, very few populations, steep declines, or other N2 factors. S2

G3 Vulnerable—At moderate risk of extinction or elimination due to a restricted range, relatively few populations, recent and wide- N3 spread declines, or other factors. S3

G4 Apparently Secure—Uncommon but not rare; some cause for long-term concern due to declines or other factors. N4 S4

G5 Secure—Common; widespread and abundant. N5 S5

Biotic Landscape - Chapter 15 117 Table 15-2. Species rankings for the IUCN red list (IUCN 2012).

Status Abbreviation Status Description For the ICUN Red Book List Extinct EX A taxon is Extinct when there is no reasonable doubt that the last individual has died. A taxon is presumed Extinct when exhaustive surveys in known and/or expected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record an individual. Surveys should be over a time frame appropriate to the taxon’s life cycles and life form. Extinct in the Wild EW A taxon is Extinct in the Wild when it is known only to survive in cultivation, in captivity or as a natural- ized population (or populations) well outside the past range. A taxon is presumed Extinct in the Wild when exhaustive surveys in known and/or expected habitat, at appropriate times (diurnal, seasonal, annual), throughout its historic range have failed to record an individual. Surveys should be over a time frame appropriate to the taxon’s life cycle and life form. Critically Endangered CR A taxon is Critically Endangered when the best available evidence indicates that it meets any of the criteria A to E for Critically Endangered, and it is therefore considered to be facing an extremely high risk of extinction in the wild. Endangered EN A taxon is Endangered when the best available evidence indicates that it meets any of the criteria A to E for Endangered, and it is therefore considered to be facing a very high risk of extinction in the wild. Vulnerable VU A taxon is Vulnerable when the best available evidence indicates that it meets any of the criteria A to E for Vulnerable, and it is therefore considered to be facing a high risk of extinction in the wild. Near Threatened NT A taxon is Near Threatened when it has been evaluated against the criteria but does not qualify for Criti- cally Endangered, Endangered or Vulnerable now, but is close to qualifying for or is likely to qualify for a threatened category in the near future. Least Concern LC A taxon is Least Concern when it has been evaluated against the criteria and does not qualify for Critically Endangered, Endangered, Vulnerable or Near Threatened. Widespread and abundant taxa are included in this category. Data Deficient DD A taxon is Data Deficient when there is inadequate information to make a direct, or indirect, assessment of its risk of extinction based on its distribution and/or population status. A taxon in this category may be well studied, and its biology well known, but appropriate data on abundance and/or distribution are lacking. Data Deficient is therefore not a category of threat. Listing of taxa in this category indicates that more information is required and acknowledges the possibility that future research will show that threatened classification is appropriate. It is important to make positive use of whatever data are available. In many cases great care should be exercised in choosing between DD and a threatened status. If the range of a taxon is suspected to be relatively circumscribed, if a considerable period of time has elapsed since the last record of the taxon, threatened status may well be justified. Not Evaluated NE A taxon is Not Evaluated when it is has not yet been evaluated against the criteria.

Table 15-3. High conservation state of Alabama concern Table 15-4. High conservation state of Alabama concern wetland species within the greater Mobile-Tensaw River for species inhabiting uplands adjacent to swamps and area. bottomlands of the greater Mobile-Tensaw River area.

Birds Natural Heritage Rank Red Book Birds Natural Heritage Rank Red Book Least Bittern G5, S2N, S4B LC American Kestrel G5, S3B, S5N LC Reddish Egret G4, S1B, S3N NT American Woodcock G5, S3B, S5N LC American Black Duck G5, S2B, S5N LC Short-Eared Owl G5, S2N LC Swallow-Tailed Kite G5, S2 LC Wood Thrush G5, S5 LC Northern Harrier G5, S3N LC Worm-Eating Warbler G5, S4B LC Black Rail G3-G4, S2N NT Swainson’s Warbler G4, S4B LC Yellow Rail G4, S2N LC Kentucky Warbler G5, S4B LC White Ibis G5, S2B, S3N LC Bachman’s Sparrow G3, S3 NT American Oystercatcher G5, S1 LC Seaside Sparrow G4, S2 LC

118 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Because of its long, unbroken tracts of bottomland forests, the greater Mobile-Tensaw River area acts as a flyway for a multitude of avian species. With places to hide, rest, and feed, this area is essential for species making the long, arduous journey from Central and South America through the southeastern US and on to the heart of North America to breed (Fitzpatrick 2004; Gill 2007).

Impacts of Birding The US Fish and Wildlife Service has estimated that birders contribute $85 billion in overall economic output per year, with $13 billion going toward state and federal income taxes (Gill 2007). With such a great diversity of bird species throughout the greater Figure 15-14. White Ibis. (Photo credit: Kathy Hicks). Mobile-Tensaw River area, birders are attracted here from all over the globe. Warbler, Bachman’s Sparrow, and Seaside Sparrow. Enthusiasts spend considerable amounts of money Additionally, 48 of 53 species (90%) listed under boosting economies in large cities and small commu- “Moderate Concern” in Alabama have been encoun- nities just to see a Bald Eagle, Swallow-tailed Kite, tered in the greater Mobile-Tensaw River area. Essen- Prothonotary Warbler, Great Egret, or Osprey for the tially half of all bird species found in Alabama utilize first time. Most birders are nature enthusiasts who are this region, making it a critical ecological corridor awed by the great diversity of our planet and wish to for avian population stability (Mirarchi et al. 2004; contribute to its maintenance for future generations. NatureServe 2015). Conservation Implications Neotropical Migrants While every state has witnessed species extinctions, such losses have not occurred uniformly across the Since the inception of the Alabama Breeding Bird nation. States exhibiting high numbers of extinctions Atlas, an estimated 36 out of 102 (35%) species that tend to boast high overall numbers of species and breed throughout the greater Mobile-Tensaw River fragile ecosystems with at-risk flora and fauna and/or area are Neotropical migrants, birds that breed in intense human impact. Unfortunately for Alabama, all North America but migrate to Central and South 3 of these scenarios apply. According to data collected America to overwinter (Alabama Ornithological Soci- by the nation’s Natural Heritage programs and The ety 2015). An additional 25 species use it as a major Nature Conservancy, Alabama is the most extinction- migration route on their journey north. Without the prone state of the mainland United States, with 96 varied and rich habitats of the greater Mobile-Tensaw documented species extinctions to date (Stein et al. River area, these species would surely be subject to 2000; NatureServe 2015). increased predation and starvation due to the lack of sufficient mature hardwood bottomland and upland Because of its ancient and complex geological terrain pine forests. spanning 3 major river basins, Alabama is home to more species of animals and plants than any other Migration Corridor state east of the Mississippi River. An estimated 10 to The greater Mobile-Tensaw River area also serves 20% of all species known to occur in Alabama are in as an uninterrupted corridor for the movement of peril due to ever-increasing pressures associated with species in and out of the Central Gulf Coastal Region. increased incidence of disturbance, both natural (e.g.,

Biotic Landscape - Chapter 15 119 hurricanes, flooding events) and human-induced generally lack enough space, food, nesting sites, or (e.g., eutrophication, development, industrialization, refuge from predators to support sustainable popula- oil spills) (Stein et al. 2000; Brawn et al. 2001). tions (Wilcove et al. 1998; Robinson et al. 1995).

We have already lost one bottomland hardwood One problem in particular for small woodland species inhabitant, Bachman’s Warbler (GH-Global Rank is the need for large areas of mature forest. Most of “Possibly Extinct,” SX-State Rank “Extinct,” our mature hardwood and pine forests have been LE-USFWS “Endangered,” CR-IUCN Red List relegated to thin strips that are mostly (if not entirely) “Critically Endangered”), to clearcutting of the tupelo edge habitat. These edge habitats, the physical bound- gum-cypress swamps and severe reduction of natural ary between one habitat type and another, tend to cane-brakes along river levees. And we likely have lost be hotter and dryer than suitable habitats in larger another, the Ivory-billed Woodpecker (G1, SX, LE, forests. Such environmental shifts increase time spent CR), the largest woodpecker in North America, which on eggs rather than foraging, thereby reducing paren- has certainly been extirpated or driven into extinc- tal energy stores. tion altogether. Along with these two losses, dozens more species have seen precipitous declines in both Predation also tends to be higher along edges. Not numbers and distribution (Anderson 1996; Mirarchi only are adults, their eggs, and offspring preyed upon et al. 2004a; Gill 2007). by mammalian, reptilian, and raptorial predators, but obligate brood parasites, especially Brown-headed Measurable reductions in biodiversity around the Cowbird and occasionally Yellow-billed Cuckoo, are globe have been attributed to many factors, includ- more common within edge habitats (Robinson et al. ing fragmentation of undeveloped land for farming, 1995). While these species do not eat eggs or hatch- logging, and suburban development, and invasive lings, they are responsible for dramatic declines in species (Murcia 1995; Dobson 2000; Stein et. al 2000; populations of vireos, warblers, flycatchers, and other Wittenberg and Cock 2001). Loss of habitat diversity Neotropical migrants that are forced to nest close to has also been implicated in species declines through- forest edges. Brood parasites do not build their own out the world (Dobson 2000; Stein et al. 2000). The nests. Instead they lay one of their eggs in an estab- disappearance and degradation of wetlands, as well lished vireo or warbler nest. The parasite’s egg hatches as changes in stream hydrology through dredging and first and its larger hatchling either pushes the remain- channelization, have also resulted in losses in diversity ing eggs and/or hatchlings out of the nest or demands and trophic structure (Russell et al. 2002; Ryan et al. more food from the host parents. Thus, parasitic 2002; Semlitsch 2003). behavior decreases resources available to the host’s offspring and greatly reduces survivorship (Robinson Changes in hydrodynamics, through channelization et al. 1995; Payne 1998; Davies 2000). and increased sedimentation caused by poor soil conservation, have altered natural flood cycles of Summary the Mobile-Tensaw Delta and surrounding uplands. Moreover, poorly managed private lands leased for Birds are an incredibly important component of the silviculture and a reduction in riparian zones have greater Mobile-Tensaw River area ecosystem. Many diminished available habitat necessary for spring of these species are imperiled due to habitat loss or migration and nesting that affect species of high and degradation through reduction in riparian zones, moderate concern. Neotropical migrant species fragmentation of large, contiguous forest tracts, and impacted by these conditions include, but are not reduction of wetlands from development. It is imper- limited to, Swainson’s Warbler, Cerulean Warbler, ative that this diverse and ecologically important area Kentucky Warbler, Worm-eating Warbler, Wood is managed for the use and enjoyment of future gener- Thrush, Prothonotary Warbler, Northern Parula, ations of scientists and nature enthusiasts alike. The and Louisiana Waterthrush (Ward 1989; Naiman and Mobile-Tensaw Delta has been called “a jewel in the Décamps 1997; Paetzold et al. 2005; Gill 2007). crown of North America” and “an American Amazon” by noted biologist, scholar, and native Alabamian, E. Habitat fragmentation has also been cited as a major O. Wilson (EO Wilson Biodiversity Foundation [date cause of population decline in wetland species, unknown]). We would do a great disservice to the upland pine savanna specialists, and large, contiguous generations yet to come should we fail to protect its mature forest nesters (Askins 1995). Small habitats beauty and diversity.

120 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 16. Mammals Troy L. Best, Auburn University

Modern research (Mirarchi 2004; Mirarchi et al. 1968; McBee and Baker 1982; White 1992; Best and 2004a, 2004b; Best and Dusi 2014) has recently Dusi 2014). Until the 1980s, nine-banded armadil- augmented standard listings and summaries about los were known only from southern Alabama, but by the mammals of southern Alabama by Howell (1909, the mid-1990s they had extended their range across 1921), Holliman (1963), and Mount (1984, 1986). At the state and well into Tennessee. Insects, especially present, the mammalian fauna of the greater Mobile- beetles and ants, are common items in their diet; they Tensaw River area contains 9 taxonomic orders repre- also eat millipedes, centipedes, earthworms, snails, sented by 22 families and about 58 species. This chap- eggs, amphibians, reptiles, birds, baby mammals, ter reviews mammals living in or extripated from the carrion, fruits, and berries. Although they consume greater Mobile-Tensaw River area. many harmful insects, nine-banded armadillos often are not welcome because of damage they cause to Overview vegetable gardens and flower beds. Of the 58 species of the greater Mobile-Tensaw River West Indian manatees are warm-season residents in area, most are rodents (20 species), carnivores (14), and around Mobile Bay. They have been observed as and bats (12) (Table 16-1). Some species have been far north as Claiborne Lock and Dam (Carmichael extirpated in Alabama (red wolf, cougar), are of ques- RH, et al., in litteris, unreferenced) on the Alabama tionable occurrence (jaguarundi), are known from the River, about 8 km (5 mi) upstream from Claiborne, region only by a few historical records (southeastern Monroe County. These large, streamlined aquatic pocket gopher, little brown myotis, marsh rabbit), or mammals have flippers instead of front legs, no hind are officially listed as endangered (West Indian mana- legs, and a flattened-spatulate tail (Husar 1978). Their tee, gray myotis, red wolf). closest relatives are elephants (Proboscidea) and Opossums, Manatees, and Armadillos Virginia opossums, West Indian manatees, and nine-banded armadillos are the only representatives of their taxonomic orders in the greater Mobile- Tensaw River area (Didelphimorphia, Sirenia, and Cingulata, respectively). Virginia opossums (Figure 16-1) occur statewide in all terrestrial habitats, but they favor floodplains and swamps. Hedges, piles of wood, fence rows, stream banks, and other habitats provide cover in suburban areas. These opportunistic omni- vores consume insects, carrion, fruits, garden vegetables, foods discarded by humans, and food left outside for pets. Virginia opossums are a common roadkill on our streets and highways (McManus 1974; Best and Dusi 2014), Figure 16-1. Virginia opossum (Didelphis virginiana) is a common species as are nine-banded armadillos (Wolfe in the greater Mobile-Tensaw River area of Alabama. (Photo credit: Troy L. Best.)

Biotic Landscape - Chapter 16 121 Table 16-1. Mammals of the greater Mobile-Tensaw River area of Alabama (Best and Dusi 2014) and a qualitative estimate of their current status in the region.

Order Family Species Current Status Didelphimorphia (American Opossums) Didelphidae (American Virginia opossum (Didelphis virginiana) Common Opossums) Sirenia (Dugongs, Manatees, and Sea Cows) Trichechidae (Manatees) West Indian manatee (Trichechus manatus) Endangered Cingulata (Armadillos) Dasypodidae (Armadillos) Nine banded armadillo (Dasypus novemcinctus) Common Rodentia (Rodents) Sciuridae (Squirrels) Eastern gray squirrel (Sciurus carolinensis) Common Eastern fox squirrel (Sciurus niger) Common Southern flying squirrel (Glaucomys volans) Common Eastern chipmunk (Tamias striatus) Uncommon Castoridae (Beavers) American beaver (Castor canadensis) Common Geomyidae (Pocket Gophers) Southeastern pocket gopher (Geomys pinetis) Rare Cricetidae (New World Rats Woodland vole (Microtus pinetorum) Uncommon and Mice, Voles, Hamsters, and Common muskrat (Ondatra zibethicus) Uncommon Relatives) Eastern woodrat (Neotoma floridana) Uncommon Golden mouse (Ochrotomys nuttalli) Uncommon Cotton deermouse (Peromyscus gossypinus) Common White-footed deermouse (Peromyscus leucopus) Uncommon Oldfield deermouse (Peromyscus polionotus) Common Eastern harvest mouse (Reithrodontomys Common humulis) Marsh oryzomys (Oryzomys palustris) Common Hispid cotton rat (Sigmodon hispidus) Common Muridae (Old World Rats and House mouse (Mus musculus) Common Mice) Brown rat (Rattus norvegicus) Common Roof rat (Rattus rattus) Uncommon Myocastoridae (Coypus) Coypu (Myocastor coypus) Uncommon Lagomorpha (Hares, Pikas, and Rabbits) Leporidae (Rabbits and Hares) Swamp rabbit (Sylvilagus aquaticus) Common Eastern cottontail (Sylvilagus floridanus) Common Marsh rabbit (Sylvilagus palustris) Rare Soricomorpha (Shrews, Moles, Desmans, and Soricidae (Shrews) Southern short tailed shrew (Blarina carolinensis) common Relatives) North American least shrew (Cryptotis parva) Common Southeastern shrew (Sorex longirostris) Common Talpidae (Moles) Eastern mole (Scalopus aquaticus) Common Carnivora (Carnivores) Family Felidae (Cats) Bobcat (Lynx rufus) Common Cougar (Puma concolor) Extirpated Jaguarundi (Puma yagouaroundi) Questionable Canidae (Wolves, Dogs, Foxes, Coyote (Canis latrans) Common and Jackals) Red wolf (Canis rufus) Extirpated Gray fox (Urocyon cinereoargenteus) Common Ursidae (Bears and Giant American black bear (Ursus americanus) Uncommon Pandas) Mustelidae (Weasels, Badgers, North American river otter (Lontra canadensis) Common and Otters) Long-tailed weasel (Mustela frenata) Uncommon American mink (Neovision vison) Common

122 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Table 16-1. Mammals of the greater Mobile-Tensaw River area of Alabama (Best and Dusi 2014) and a qualitative estimate of their current status in the region, cont.

Order Family Species Current Status Carnivora (Carnivores), cont. Mephitidae (Skunks and Stink Striped skunk (Mephitis mephitis) Common Badgers ) Eastern spotted skunk (Spilogale putorius) Uncommon Procyonidae (Raccoons, Ring- Raccoon (Procyon lotor) Common tails, and Coatis) Artiodactyla (Even-toed Ungulates) Suidae (Swine) Wild boar (Sus scrofa) Common Cervidae (Deer, Elk, Caribou, White-tailed deer (Odocoileus virginianus) Common and Moose)

hyraxes (Hyracoidea) (Best and Dusi 2014). They have Of the 4 species of squirrels, 3 (eastern gray squir- no predators, but scars from propellers and keels of rel, eastern fox squirrel, southern flying squirrel) are boats are evidence that humans may have a signifi- common in the greater Mobile-Tensaw River area, cant impact on populations (Hartman 1979). West but the eastern chipmunk is rare at this southern Indian manatees are nearly completely herbivorous edge of its geographic range (Hall 1981; Best and Dusi and usually forage on vegetation in shallow coastal 2014). American beavers (Castoridae) are common, freshwater and saltwater habitats (Husar 1978). but local populations may be reduced or removed by Causes of death associated with humans include colli- humans when they become a nuisance because they sions with boats; drowning in fishing nets; ingestion construct dams across streams and damage trees. The of fishing hooks, fishing lines, and wire; and entrap- family Geomyidae is represented only by the south- ment in locks, dams, and large pipes (Husar 1978; eastern pocket gopher, at the western edge of its Best and Dusi 2014). West Indian manatees are soli- geographic range (Merriam 1895; Harper 1952; Hall tary or occur in pairs (often a female and her young) 1981). Southeastern pocket gophers are associated or in small groups; larger groups may form near warm with rolling topography, sandy soils, longleaf pines springs in inland waterways during winter (Husar (Pinus palustris), and turkey oaks (Quercus laevis). 1978; Hartman 1979). In autumn, many of Alabama’s They also inhabit highway right-of-ways, parks, lawns, West Indian manatees migrate along coastal water- golf courses, orchards, cemeteries, and pasturelands. ways to warm water springs near Crystal River, Citrus Their burrowing mixes and aerates the soil, and their County, Florida (Carmichael RH, et al., in litteris, burrows provide shelter for a variety of organisms unreferenced). (Best and Dusi 2014).

Rodents The most diverse family of rodents in North America and in the greater Mobile-Tensaw River area is Cricet- Of the 6 families of rodents in the greater Mobile- idae. Each of the 10 species in this family occupies one Tensaw River area, two families (Muridae, Myocas- or more habitats in the area, from aquatic and marshy toridae) were introduced into North America. Within (golden mouse, common muskrat, marsh oryzomys), the family Muridae, native range of the house mouse to woodlands (woodland vole, eastern woodrat, is nearly all of Europe, Africa, and Asia; the brown rat cotton deermouse), to fallow agricultural fields and is native to central Asia; and the roof rat occurs natu- grasslands (white-footed deermouse, oldfield deer- rally in southeastern Asia (Best and Dusi 2014). These mouse, eastern harvest mouse, hispid cotton rat). species now occur throughout the world in close Although most are active at night, some also are active association with humans. Natural range of the family during daylight (woodland vole, common muskrat, Myocastoridae (coypu, also known as nutria) extends hispid cotton rat). Most species use underground from Brazil and Paraguay to the tip of South America burrows or nests in vegetation for protection from (Woods et al. 1992). Imported to the United States in environmental extremes, to escape from predators, to about 1900, the species escaped or was released inten- rest, and to rear young. In a study of small mammals in tionally in the 1930s, and increased dramatically after the upper Mobile-Tensaw Delta, Gay (2006) reported the fur market collapsed in the early 1980s (Best and that the cotton deermouse (37 individuals), marsh Dusi 2014).

Biotic Landscape - Chapter 16 123 oryzomys (25), and hispid cotton rat (1) were the only Mobile-Tensaw River area. All are common, but few rodents he encountered during intensive sampling in are observed due to their small size, concealment in spring and summer of 2004 and 2005. surface and subsurface habitats, and secretive nature. Southern short-tailed shrews occur in fallow agri- Eastern woodrats are somewhat exceptional for their cultural fields, roadsides, grasslands, brushy areas, elaborate denning sites (middens) constructed mainly cane bottomlands, swampy areas, hardwood and of branches, twigs, and leaves, although they can also mixed pine-hardwood forests, and dense thickets. incorporate bones, cattle dung, aluminum cans, and Diet includes vegetative matter, slugs, snails, fungi, shotgun shells. These middens may be 2 to 4 m (7 to 13 earthworms, insects, centipedes, spiders, and eggs of ft) wide and 1 m (3 ft) tall, built under rocky outcrops, turtles (Genoways and Choate 1998; McCay 2001). in brush piles, at the base of large trees, in trees, or in abandoned buildings and motor vehicles. Middens North American least shrews often inhabit dry grassy, may be maintained, modified, and occupied by gener- weedy, and brushy fields, but they may be present in ations of eastern woodrats, and they also are used as marshes, woodlands, and forests with maples, oaks, shelter by many kinds of animals (Best and Dusi 2014). and pines. Diet includes fungi, earthworms, centipedes, snails, slugs, spiders, insects, and some plant Hares, Pikas, and Rabbits material (Whitaker 1974). Three species of rabbits (swamp rabbit, eastern Southeastern shrews occur in moist to wet areas, cottontail, marsh rabbit) occur in southern Alabama. usually bordering marshes, swamps, or rivers. They The marsh rabbit has not been recorded from the also may be common in fallow or planted fields, dry greater Mobile-Tensaw River area, but is included upland hardwoods, planted pines, and dry sandy here because of reported occurrence in southern areas. Diet includes vegetation, slugs, snails, spiders, Baldwin County (Hall 1981). Marsh rabbits occupy centipedes, and insects (French 1980a; 1980b). damp or swampy lowlands, marshes, and brackish- water areas (Chapman and Willner 1981; Chapman Eastern moles inhabit moist, loamy, or sandy soils, and and Litvaitis 2003), all habitats present in the greater they are rare or absent in soils with dense clay, stones, Mobile-Tensaw River area. Swamp rabbits primarily or gravel. Woodlands and floodplains along rivers and occur in swamps, floodplains, marshes, and lowlands, streams are frequently occupied, but they may occur where they may construct shelters in vegetation, or in grassy habitats and cultivated fields. Diet includes they may use holes in the ground or in trees. Diet earthworms, insects, and some plant material (Arlton includes a variety of plants. Swamp rabbits are excel- 1936; Yates and Schmidly 1978). lent swimmers and will readily take to water to avoid predators (Chapman and Feldhamer 1981; Chapman Bats and Litvaitis 2003). In southern Alabama, some species of bats are active Eastern cottontails occupy diverse habitats, including throughout most of the year (e.g., eastern red bat, urban areas, agricultural fields, hedgerows, shrub- hoary bat, Seminole bat, Brazilian free tailed bat), lands, grasslands, swamps, and forests. Diet varies including warm nights in winter. Some migrate (e.g., seasonally and consists of a variety of plant material, eastern red bat, gray myotis), and some become including woody and herbaceous species. Eastern torpid or hibernate (e.g., big brown bat, Rafinesque’s cottontails are an important food for many raptors big-eared bat [Figure 16-2], tri-colored bat). Of the and mammalian carnivores. Shelters made by clear- 12 species of bats occurring in the greater Mobile- ing sites under vegetation, especially under clumps of Tensaw River area, 7 are common, 4 are rare (hoary grasses, are used for resting and hiding from predators bat, northern yellow bat, Rafinesque’s big eared bat, (Chapman et al. 1980; Chapman and Litvaitis 2003). little brown myotis), and 1 is endangered (gray myotis). There have been reports of parasites and rabies in Shrews, Moles, Desmans, and bats with geographic distributions that include south- Relatives ern Alabama (e.g., White 1959; LaVal 1967; Best et al. 1993; Hilton and Best 2000; Hester and Best 2007). In Three species of shrews (southern short-tailed shrew, the most thorough study of the region’s bats, Kilgore North American least shrew, southeastern shrew) and (2008) compiled data on relative abundance of 7 1 species of moles (eastern mole) occur in the greater species of bats occupying different-aged forest stands

124 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area in the Mobile-Tensaw Delta: Seminole bat (51 individuals), tri-colored bat (49), evening bat (32), eastern red bat (22), southeastern myotis (15), Rafin- esque’s big eared bat (4), and hoary bat (2). Two other species (Brazilian free-tailed bat, big brown bat) are common in the region, but they often roost in buildings and other human- made structures, which were uncommon in Kilgore’s (2008) study area. In addition, Brazilian free-tailed bats fly high and fast (Wilkins 1989; Harvey et al. 2011; Best and Dusi 2014), possibly at altitudes much higher than where Kilgore (2008) obtained bats for his study.

Carnivores Six families of carnivores represented Figure 16-2. The Rafinesque’s big eared bat Corynorhinus( rafinesquii) is a by 14 species occur in the greater rare species that occupies caves, abandoned buildings, and hollow trees in Mobile-Tensaw River area. Although the greater Mobile-Tensaw River area of Alabama. (Photo credit: J. Scott Altenbach.) carnivores are active during day and night, they generally are secretive and not easily observed (for example, the long-tailed weasel [Mustela frenata], Figure 16-3). Common species such as bobcats, coyotes, gray and red foxes, North American river otters, American minks, striped skunks, and raccoons are seen more often as roadkills on highways than alive. The cougar has been extirpated in Alabama and the native subspecies (Puma concolor cougar) is believed to be extinct throughout its range in eastern North America (USFWS 2015b). There are many reports of tracks and sightings of cougars in Alabama, including a report of a breeding pair in northern Baldwin County in 1984 (Mount 1984), but whether these individuals are from a natural population or released captives is unknown (Best and Dusi 2014). Large black Figure 16-3. The long tailed weasel (Mustela frenata), once a common spe- cats also have been reported through- cies throughout the southeastern United States, is currently uncommon in the greater Mobile-Tensaw River area and elsewhere in Alabama. (Photo out Alabama, but those large cats are credit: Troy L. Best.) not black cougars. During studies of geographic variation in cougars (Gay and Best 1995, 1996a, 1996b), no black

Biotic Landscape - Chapter 16 125 cougar was observed among hundreds of specimens In Alabama, humans are the only significant preda- examined from throughout North and South America. tors of adult feral swine (Sweeney et al. 2003; Best and Dusi 2014). The occurrence of the jaguarundi in Alabama is questionable because no specimen from the state has The white-tailed deer is a prized game species in the been examined. Sightings may have been of escaped greater Mobile-Tensaw River area and throughout or released individuals, or of some other species of most of its geographic range. This species inhabits mammal (e.g., large domestic cat). most terrestrial habitats, from fallow agricultural fields to dense forests. Diet includes agricultural crops (e.g., The last red wolf was killed in Alabama in 1917 corn, wheat, soybeans, alfalfa, fruit trees), grasses, (Nowak 1972), but the last record for Baldwin and forbs, fruits, mushrooms and other fungi, and succu- Mobile counties was in 1894 (Howell 1921). Red lent leafy vegetation; acorns are readily consumed wolves are critically endangered; the species now when available. White-tailed deer may cause signifi- exists in the wild only as a re-introduced population cant damage to agricultural crops, orchards, nurser- in eastern North Carolina (e.g., Phillips et al. 2003). ies, and ornamental plants in yards and gardens. In Prior to the mid-1900s, American black bears prob- Alabama, they are the most economically important ably occurred statewide in Alabama. Tracks, sightings, game species, with 300,000 to 500,000 white-tailed and other evidence indicate that a few individuals or deer harvested each year (Smith 1991; Best and Dusi small populations now exist throughout eastern and 2014). White-tailed deer cause considerable property southern parts of the state. Currently, the only breed- damage through collisions with vehicles on roads, ing populations appear to be in northeastern Alabama especially in rural areas. and in the greater Mobile-Tensaw River area (including parts of Baldwin, Clarke, Escambia, Mobile, and Ever-increasing Threats Washington counties). Diet is primarily vegetation, The greater Mobile-Tensaw River area is home to but insects, reptiles, birds, and mammals also are a biologically diverse fauna that is subject to ever- consumed opportunistically. In spring American increasing threats from encroachment by humans, black bears consume new growth of vegetation and loss of habitats, changes in global climate, invasions winter-killed carrion. In summer they eat herbaceous by exotic species, and other environmental pres- vegetation and fruits, and in autumn they feed primar- sures. With about 58 species of mammals, the greater ily on fruits and nuts. Adults have few predators, but Mobile-Tensaw River area contains one of the richest young may be preyed upon by bobcats, coyotes, and mammalian faunas in Alabama. Increases in popula- other American black bears. Most mortality is caused tions of humans and their associated domestic species by humans through shooting, trapping, and collisions (e.g., dogs, cats, cattle); construction of commercial with vehicles (Best and Dusi 2014). and residential structures and paving of highways and parking lots; extension of power grids, utilities, and Even-toed Ungulates pipelines; applications of pesticides, fertilizers, and In the 1500s, Spanish explorers introduced wild boars other chemicals; destructions of local flora and fauna into what is now the southeastern United States, from by exotic species (e.g., wild boars); and other pertur- their native range in northern Africa, Europe, and bations of this ecosystem, although of less conse- Asia. Feral swine now occur statewide in Alabama quence when assessed individually, add up to signifi- and they are the most destructive invasive species cant changes in our natural world. When coupled in the state. They are omnivorous and will consume with what may seem to be incremental increases in almost any type of plant or animal materials, includ- global temperatures and changes in weather patterns, ing agricultural crops, food discarded by humans, we should expect eventual reductions in some popu- carrion, small mammals, birds, turtles, snakes, and lations and loss of some species of mammals in the amphibians. By rooting large areas they have wide- greater Mobile-Tensaw River area. spread negative impacts, including destruction of Mammalian species especially vulnerable to human- crops, property, archeological deposits, native fauna, made environmental changes in the greater Mobile- and natural vegetation, as well as the disruption of Tensaw River area include the West Indian manatee, soils, seed banks, tree roots, and natural succession. southeastern pocket gopher, marsh rabbit, Brazilian

126 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area free tailed bat, Rafinesque’s big eared bat, gray myotis, Personnel of local, state, and national agencies and little brown myotis, American black bear, North organizations, landowners and land managers, and American river otter, long tailed weasel, American other informed citizens must work together if we are mink, eastern spotted skunk, and white tailed deer. to preserve our natural heritage for future generations.

Biotic Landscape - Chapter 16 127

CULTURAL LANDSCAPE

Alabama River, Dallas County, Alabama. Photograph courtesy of Hunter Nichols.

Cultural Landscape - Chapter 17 129

17. Archeology—Arrival to AD 1550 David W. Morgan, National Park Service

The other chapters in this volume illustrate many known as the Bottle Creek site (1BA2 [Smithson- ways in which the greater Mobile-Tensaw River area ian trinomials to identify archeological sites are is exceptional—ecologically, biologically, and geologi- provided]) (Bigelow 1853; Jones and DeJarnette 1933; cally. Superlatives also apply culturally, in the myriad Waselkov 1993; Brown 2003a). As 1 of only 2 prehis- ways humans have adapted to, made use of, and lived toric-era National Historic Landmark properties in within this unique environment. For a millennium or the state (Moundville is the other), Bottle Creek origi- more this area was a fulcrum of cultural interaction, a nated around AD 1250, and is characterized by a plaza major node on overland and canoe trails emanating and at least 20 mounds built atop a seasonally inun- from the continental interior and coastal societies to dated island (Figure 17-1). There is no other site like it the east and west. in scale on the central Gulf Coast, and no other coeval mound complexes (of any scale) within the greater Bottle Creek Mobile-Tensaw River area. The most significant site at this intersection was the Bottle Creek served as principal town for the Pensac- complex of monumental and residential earthworks ola culture, which spanned the greater Mobile-Tensaw

Figure 17-1. Topographic map of the Bottle Creek mounds (1BA2) (Waselkov 1993). Since creation of this map, several additional mounds have been discovered. (Image credit: Sarah Mattics, Center for Archaeological Studies, University of South Alabama.)

Cultural Landscape - Chapter 17 131 River area, westward along the Mississippi coast, and of monumental earthen construction (Dobyns 1983; eastward along the Florida panhandle (Stowe 1985; Ramenofsky 1987). Fuller and Brown 1995; Brown 2003a). The Pensacola culture, like other chiefly societies that flourishedcirca Short-term and long-term processes that led to the AD 1200 to 1550, was organized internally by social genesis of Bottle Creek remain murky. Indeed, the inequality that is partly reflected in the Bottle Creek archeology of the greater Mobile-Tensaw River area settlement pattern. Five large mounds—the biggest is very poorly understood. The few research projects one being a flat-topped pyramidal earthwork standing undertaken there offer us a patchwork of vignettes 16 m (52 ft) high—enclose an open plaza. The mounds scattered across a multi-millennial timeline (Figure themselves were platforms for residences occupied by 17-2). Consider, for example, what we do not know members of a hereditary elite who served as sociopo- about Bottle Creek. The place surely was occupied litical and religious community leaders. Surrounding seasonally because of the Delta’s regular, predict- the plaza and its ring of large mounds were numerous, able inundation every spring (Curren 1976; Knight smaller residential mounds that arose gradually from 1984; Brown 2003b). Where did the site’s occupants the accretion of household refuse. Inhabitants of reside during those inhospitable months? A partial these residential mounds are thought to have served answer may be found at the Crossroads Mounds site as retainers to the elite of the central mounds. (1BA356), which consists of 3 small earthen mounds arranged in an approximate equilateral triangle, No matter their social rank, all of the site’s occupants perched on the eastern bluffs overlooking the Tensaw consumed the same kinds of foods, with a heavy River on land owned by the US Army Corps of Engi- emphasis on corn (maize, Zea mays), brackish water neers (Figure 17-3). Two private archeological compa- clams (Rangia cuneata), and vast quantities of season- nies have conducted surveys in the vicinity of the site ally available small schooling fish of the Clupeidae in the last 2 decades (Mozingo and Whitley 2000), but family, like shad (Alosa sapadissima) and menhaden the age of this site remains uncertain. It may be affili- (Brevoortia patronus) (Quitmyer 2003; Scarry 2003a, ated with the Pensacola culture, and thus linked to 2003b). Corn was brought to the site off the cob, as Bottle Creek, or the site may predate Bottle Creek by shelled kernels, perhaps as tribute from the lowest as much as 1100 years. Similarly, Bottle Creek’s chiefly social class living elsewhere, presumably at the non- rulers are thought to have dominated non-mound mound sites of this era scattered throughout the and single-mound Pensacola centers in Mississippi greater Mobile-Tensaw River area (Brown 2003b). and northwest Florida (Knight 1984; Bense 1994, p Internal social differences are further suggested by 234-8; Blitz and Mann 2000), yet insufficient research differences in ceramic vessels resulting from food has occurred to test that hypothesis. Thus it remains preparation by nobility at the residential mounds, unclear if Bottle Creek represents the apex of a 3-tier slated for consumption by the paramount elites on or 2-tier settlement system, or if it sits in isolation as the larger mounds (Johnson 2003). The presence Moundville’s “gateway” community near the Gulf of yaupon holly (Ilex vomitoria) at the accretional Coast (Brown 2003b). mounds provides a clue to elite religious activities at the complex, as yaupon leaves were used to brew the A systematic and thorough exploration of such caffeinated Black Drink—principal ritual beverage of ancient social dynamics is one area of research that Pensacola and other similar Mississippian cultures. will prove fascinating in the years ahead. Future research should include a consideration of Bottle Use of the Bottle Creek site declined by AD 1550, Creek’s relationship with the next closest society of although the cause of this eclipse is unknown. Cycles comparable size and age—Moundville, located in the of social formation and dissolution were a regular part Black Warrior River valley—particularly with respect of the Southeastern cultural landscape throughout to architecture and other types of evidence unavail- the first half of the nd2 millennium AD, due to a variable in prior years. It also should consider how Bottle ety of interacting social and environmental factors. Creek articulated with the non-mound habitation Moreover, we now know that diseases introduced to sites found throughout the Mobile-Tensaw Delta the region by European explorers of the 16th century and surrounding uplands. Studies conducted already devastated indigenous populations and contributed to provide some tantalizing clues. Many researchers, for the collapse of chiefly power structures and cessation instance, suspect that the Bottle Creek complex was a

132 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figure 17-2. Timeline of cultures in the greater Mobile-Tensaw River area. (adapted from Fuller 1998, p 4, Figure 2).

Moundville development or intrusion (e.g., compare a study of animal remains at Bottle Creek identified DeJarnette 1952; Sears 1977; Brose et al. 1983; Fuller a single snail (Elimia hydei) endemic to the Black and Brown 1998, p 88; Brown and Fuller 1999; Brown Warrior River valley (Quitmyer 2003), which conjures 2003b; Fuller 2003). This argument is based mostly on an image of this tiny mollusk clinging to a southbound comparable early ceramic types and the lack of a signif- dugout canoe traveling down the Tombigbee River to icant older occupation at Bottle Creek. Intriguingly, the Mobile-Tensaw Delta.

Cultural Landscape - Chapter 17 133 Figure 17-3. Field Sketch of the Crossroads Mound (1BA356) when initially recorded in 1998 by Richard S. Fuller, David W. Morgan, and Jack W. Morgan (on file at the US Army Corps of Engineers, Mobile District Office). This is one of the only sites in the area besides Bottle Creek (1BA2) that exhibits multiple platform mounds, but its age and relationship to the larger mound complex remains unclear.

134 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Little Lizard Creek is probably no coincidence that these two subsistence To date, excavations at just a few archeological sites intensifications occurred at a time of population in the estuary proper—Little Lizard Creek (1BA195) growth, influx, and reorganization accompanying the and D’Olive Creek (1BA196 and 1BA251)—afford us genesis of Bottle Creek. a view of Pensacola culture life outside the mound center (Curren 1976; DeJarnette and Nielsen 1976; Earliest Human Use of the Greater Knight 1984; Morgan 2003a). Information from older Mobile-Tensaw River Area Late Woodland period (circa AD 400 to 1200) sites Some of the biggest gaps in our understanding of the (Shorter 1999; Detwiler 2000; Morgan 2001, 2003a; ways humans used the greater Mobile-Tensaw River Dumas 2007) suggests a great deal of continuity in the area relate to the periods preceding the Late Wood- ways the inhabitants of the greater Mobile-Tensaw land. Not surprisingly, we know the least about the River area exploited the Delta estuary and adjacent oldest inhabitants of southwest Alabama. We suspect uplands. During the Late Woodland period, people people entered the greater Mobile-Tensaw River area lived at small villages or hamlets in the estuary during approximately 12,000 years ago, as we know they did the summer and early fall, and in the uplands on a elsewhere in the Southeast (Anderson and Sassaman seasonal or permanent basis. While in the Delta, 1996). But, aside from isolated stone projectile points they collected clams (Rangia cuneata comprised or knives found on the hills overlooking tributary 99% of the mollusks eaten at Little Lizard Creek) creeks, no archeological sites from these early years and netted a specific suite of seasonally abundant, have yet been identified (Trickey and Holmes 1971). small freshwater and brackish water fishes (7 taxa at The most ancient archeological sites within the estu- Little Lizard Creek; 98% of the fish vertebrae recov- ary were destroyed as rivers meandered over thou- ered are smaller than 6 mm [0.2 inch] in diameter). sands of years, or now lay deeply buried under later Turtles were also favorite targets, and medium-sized flood deposits within the Delta or submerged off our mammals making use of the forested wetlands, such present coast by early Holocene climate change and as raccoon, were probably collected opportunistically. rising seas (Trickey and Holmes 1971; Curren 1976). Fruits, nuts, and wild and domesticated indigenous plants complemented the animal portion of the diet. The oldest human occupations documented in the Corn was grown in minor amounts for the first time Delta date to the Late Archaic and come primarily from in the greater Mobile-Tensaw River area after circa shell middens on the south side of Tensaw Lake, specif- AD 400, probably in small garden plots in and among ically sites named Bryant’s Landing 3 and 4 (1BA176 dwellings, and became a food staple after circa AD 900 and 1BA175, respectively; Figure 17-4) (Trickey and (Curren 1976; DeJarnette and Nielsen 1976; Knight Holmes 1971). The Late Archaic period (circa 4000 1984; Gremillion 1993; Detwiler 2000; Morgan 2001, to 1400 BC) in the Southeast in general represents a 2003a, 2003b; Scarry 2003a, 2003b). culmination of trends from the preceding 4 millennia: populations expanded, regional differentiation Notable against this backdrop of almost 1100 years continued, and social complexity became increas- of continuity are subtle variations in fish and clam ingly elaborated. Mound construction occurred at a exploitation that tell us a great deal about prehistoric scale never before seen in North America, and some life. For instance, around AD 1100, roughly coincident Late Archaic sites are considerably bigger than those with the initiation of mound construction at Bottle of prior eras. Inter-community exchange height- Creek, clam sizes in the midden at the Little Lizard ened noticeably as large quantities of non-utilitarian Creek site shrank significantly, suggesting people were goods were moved great distances. The increasingly taking smaller and smaller animals, intensifying pres- specialized Late Archaic tool kit reflects their ability sure on clam beds to a point where clam populations to exploit virtually all available major food resources. were unable to reach their prior size before being In the upland Southeast by 2000 BC, exploitation of harvested (Morgan 2003a). This hypothesis matches wild plant foods resulted in domestication of several data drawn from the oldest Pensacola culture clam weedy species, providing important additions to the refuse at Bottle Creek (Quitmyer 2003). Around this subsistence regime (Cowan 1985; Ford 1985a, 1985b; same time corn shifted from a minor dietary compo- Yarnell and Black 1985; Smith 1989, 1992a, 1992b, nent to a dominant cultigen at Pensacola culture sites 1995; Yarnell 1989, 1993; Watson 1991; Chapman and throughout the greater Mobile-Tensaw River area. It Watson 1993; Scarry 1993).

Cultural Landscape - Chapter 17 135 the massive Poverty Point site and the apparent role of Poverty Point culture as a broker of long-distance exchange have been debated for decades (e.g., Ford and Haag 1955; Ford and Webb 1956; Webb 1968, 1982; Gibson 1974, 1987, 1994a, 1994b; Jack- son 1989, 1991). It will be exciting in the years ahead to attempt to identify how the Bryant’s Landing sites, and the greater Mobile- Tensaw River area in general, articulate with this broader regional cultural pattern.

Figure 17-4. Archeologist Richard S. Fuller, then of the Alabama Museum of Natural His- In the greater Mobile- tory, standing next to the Bryant’s Landing 4 site (1BA175) site. Food debris in the form Tensaw River area, the of clam shells was deposited by the initial inhabitants, who resided there in the Late Archaic period around 2100 BC. Other people took up residence at the site in the Gulf Late Archaic is often inter- Formational period (the Bayou La Batre culture), the Middle Woodland period (Porter preted as ending around Hopewell culture), the Late Woodland period (Tate’s Hammock, Coden, and Tensaw 1400 BC with the intro- Lake cultures), and the Mississippi period (Pensacola culture). This is one of the most duction of fiber-tempered archeologically informative sites recorded in the Mobile-Tensaw Delta to date because ceramics. To highlight the of its long use and stratigraphic layering. (Photo credit: David W. Morgan.) transition to a ceramic technology, the time

between Late Archaic and Woodland periods in the The Late Archaic in the greater Mobile-Tensaw River South has been dubbed Gulf Formational (circa 1400 area presumably parallels these trends observed else- BC to AD 100). Sites of this time period can be diffi- where, although we have few data from which to draw cult to distinguish from Late Archaic sites without conclusions. People first began using the Bryant’s excavation. Gulf Formational sites tend to be smaller, Landing 4 site for clam exploitation around 2100 BC. were used less intensively, and lack evidence for long- Later, at the adjacent Bryant’s Landing 3 site, another distance inter-community material exchange. Settle- clam midden formed, this time associated with copi- ment patterns generally remained the same, and there ous amounts of broken fired clay balls. Trickey and were few elaborations to tool assemblages, treatment Holmes (1971) equated these clay balls to similar of the dead, or other aspects of material culture. “Poverty Point Objects” dating to circa 1700-1350 BC Pottery became increasingly diverse in terms of vessel in the lower Mississippi River valley (Gibson 1996a, forms (and presumably functions) and decorative 1996b). Baked clay objects are ubiquitous at the treatment. Poverty Point site, in Louisiana; thousands have been encountered lining hearths at contemporary sites, and Evidence from the Early Gulf Formational (circa so are conjectured to represent an innovative cooking 1400 to 700 BC) has been revealed by excavations method (Ford and Haag 1955, p 56; Ford and Webb at the Coon Neck site (1BA290), located on Doug- 1956, p 39; Saunders 2000, p 14). las Lake in the northeast portion of the delta (Stowe 1990), and at the View Point site (1BA281) on the In the lower Mississippi valley, major Poverty Point southeastern shore of Mobile Bay (Stowe 1990; Fuller centers are surrounded by smaller sites, comprising 1998, p 7). The former site, which rests atop an old hierarchical settlement clusters (Webb 1982; Bense terrace and dates roughly to the 14th century BC, has 1994, p 101; Gibson 1996b, p 4-6). Implications of fiber-tempered pottery, steatite vessels, stone points,

136 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area and other tools. View Point had a buried peat layer, The Woodland stage in the Mobile-Tensaw River Basin which has since eroded into the bay. Fuller (1998, p begins with what is considered the Middle Woodland 7) reported a calibrated radiocarbon date for the peat period (AD 100 to 400) elsewhere in the Southeast. It layer in the 1200s BC. Collections at low tide by avoca- is distinguished from the Late Gulf Formational by the tional archeologists produced several hundred sherds general trends noted above, and because of a notice- of grit-tempered pottery, a minor amount of fiber- able increase in interregional exchange and inter- tempered pottery, stemmed projectile points/knives, action related to the Hopewell Interaction Sphere. ground and polished stone objects, and steatite vessel Beginning in the 1st century AD, new sacred infor- fragments. mation, iconography, and artifacts diffused throughout eastern North America from several core areas in By the Late Gulf Formational period (circa 700 to the Ohio and Illinois river valleys. Hopewellian traits 100 BC) 4 major pottery traditions had spread across disseminated as far north as southern Ontario and the Southeast. The Gulf Tradition is represented in Maine, south down the Atlantic Coast to middle Flor- the greater Mobile-Tensaw River area by Bayou La ida, and west to Kansas City. Unlike Archaic cultural Batre culture, with sites in the Delta and as far north elaborations, which were gradual and encompassed as the lower Tombigbee drainage (Bense 1994, p 127; all aspects of life, the Middle Woodland Hopewell Brown 2004a). Most of what we know about it relates phenomenon appears to have been restricted mainly to its pottery, which consisted chiefly of grit tempered to religious, ceremonial, and mortuary activities. In vessels, often adorned with small feet (called “podal the core areas of the Midwest the Hopewell concept supports”), either left plain or decorated with stab- may well have been holistically transformative, but in and-drag incisions or dentate rocker stamping the greater Mobile-Tensaw River area, on the south- (Wimberly 1953, 1960; Bense 1994, p 124-30; Fuller ern Hopewell periphery, these religious paraphernalia 1998; Brown 2004a). Excavations are limited to units appear to have been incorporated into otherwise local dug in several stratified sites, including the Blakeley traditions. site shell middens (1BA229) at the head of Mobile Bay (Stowe 1977); Bryant’s Landing 3 and 4 (Trickey In the Hopewell core, elaborate mortuary ritual and Holmes 1971); the Griffin site (1CK45) in Clarke included crypt burials or pit tombs in earthen County (Chase 1972; Brose et al. 1983); the Bayou mounds, with one or multiple bodies placed with La Batre Shell Midden (1MB12-14) (Wimberly 1960, specialized grave goods obtained through long- Table 18); and on the eastern bluffs of the delta at distance exchange. Burials often were associated with the Hamilton Creek site (1BA354) (Morgan 2003a). charnel houses, designed to house the dead and ritual Research at these sites has focused on pottery and participants, built on earthen platforms or enclosed stone tool types, with little attention yet devoted to by geometrical earthworks. Based on burial analysis, questions of estuarine-adaptation, settlement pattern, Hopewell is thought to represent a “big person” soci- subsistence, mortuary ritual, or social organization. ety, in which charisma and ability, structured within a kin and lineage system, enabled certain individu- Life in the Greater Mobile-Tensaw als to acquire and conspicuously redistribute wealth, River Area during the Woodland Stage thereby enhancing their own and their lineage’s pres- Our portrait of prehistoric life in the greater Mobile- tige and social status. A kin-group’s achieved rank Tensaw River area in the subsequent Woodland stage is was ephemeral and could shift with the death of indi- somewhat more detailed. There are several hallmarks vidual leaders. Each Hopewell community appears to of life in the region during the Woodland period, have been autonomous, with no overarching politi- spanning circa 100 BC to circa AD 1000. One was the cal structure (Service 1962; Dragoo 1963; Sahlins use of pottery for cooking on a daily basis, as well as 1963, 1968, 1972; Caldwell and Hall 1964; Prufer accompanying innovations in pottery manufacturing 1964; Brown 1979; Goad 1979; Seeman 1979a, 1979b, and decoration. Another was the gradual introduction 1992; Braun 1986; Webb and Snow 1988; Clay 1992; of corn, and a third was a change in stone technology, Pacheco 1996). including introduction of the bow and arrow from the On the Gulf Coast, evidence of the Hopewell Interac- Plains. All of these changes influenced technological, tion Sphere takes the form of small conical mounds, economic, political, and social developments in the usually 1 to 2 m (3 to 7 ft) tall and about 15 m (49 ft) Southeast. in diameter, with northwest Florida and the greater

Cultural Landscape - Chapter 17 137 Mobile-Tensaw River area being the two areas most life continued much as they had in prior centuries. heavily influenced and perhaps interconnected (Brose With the loss of this shared ideology, it is perhaps no 1979; Walthall 1990, p 156; Brown 2004a). The local surprise that the Late Woodland period is character- manifestation in the greater Mobile-Tensaw River ized by greater regional variability. Gulf Coast soci- area, and extending into the lower Tombigbee valley, is eties continued to interact on a local level, with the the Porter culture. Walthall (1990, p 155-65) suggested Weeden Island culture adapting and evolving from that Porter people employed two settlement types and the prior Porter culture in the greater Mobile-Tensaw two distinct modes of burial. There were short-term River area, and the Santa Rosa-Swift Creek culture occupations within the Delta, bay, and coast focused developing in the Florida panhandle. Many aspects of on exploitation of the estuary and represented by the preceding Porter and Swift Creek cultures contin- shell middens, such as Bryant’s Landing 4 (Trickey ued, including modes of settlement, subsistence, and Holmes 1971). And there were upland communi- and mortuary ritual. Some well-known examples ties, such as Porter Village (1CK21) and McVay Village are the elaborate Kolomoki and McKeithen mound (1CK1), which focused on hunting and collecting complexes to the east in southwest Georgia and the plant resources from adjacent swamps (Wimberly Florida panhandle (Sears 1956; Milanich et al. 1984; 1960). The latter sites cover several hundred square Milanich et al. 1997; Williams and Elliot 1998). Within meters and, unlike the lowland sites, contain middens the greater Mobile-Tensaw River area, Late Wood- bearing sparse numbers of mollusks but relatively land mortuary ritual on the coast is represented by abundant quantities of stone artifacts. It is unclear small sand mounds, such as those at Seymour’s Bluff if these upland sites were seasonal or longer-term (Figure 17-5), many of which are now lost to develop- occupations. ment (Moore 1905; Fuller 1998; Morgan 2005).

Porter Hopewell burials were interred within habitation sites and within burial mounds away from residential areas (Walthall 1990, p 160). The best known of the mound sites is McQuo- rquodale (1CK25), north of the Delta, near modern day Jackson, Alabama (Wimberly and Tourtelot 1941; Walthall 1990, p 158). There Porter people constructed a conical mound, about 1 m (3 ft) high and 20 m (66 ft) in diameter, that covered a smaller earthen platform. Ten individuals were buried in the mound/platform with copper adornments, galena and mica, ground stone celts, and other exotic, non-local materials. Sites along the lower Tombigbee River contain fewer burials placed on the ground before being covered with dome-shaped mounds (Brown 2004a). In contrast, Figure 17-5. Example of a small, circa AD 400 to 700 Weeden Island pot exca- at places like Porter Village and the vated from the Late Woodland period Seymour (Parkway) Mound (1BA80) Blakeley site burials were found within complex in the 1930s by the Alabama Museum of Natural History. This is 1 the middens, seemingly with no or few of 3 whole vessels believed to have been unearthed by David L. DeJarnette and John L. Bruckner, along with a crew of 6 men, who dug a test trench exotic funerary goods (Moore 1905; into mound 1BA80 in 1937. Their photo catalog labels (1933.010.366, Stowe 1977). 1933.010.367, and 1933.010.368) cause some ambiguity as the 1933 prefix appears to reference the year the museum director, Walter B. Jones, visited The influence of Hopewell ideol- the site. The 1937 acquisition date is likely correct, as Jones made only sur- ogy waned across the Southeast face collections while DeJarnette and Bruckner’s methods would have been after AD 400, while other aspects of more likely to yield intact vessels. (Image credit: Eugene Futato, Office of Archaeological Research, Moundville, Alabama.)

138 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Judging from the variability in Late Woodland pottery speculation that intensification of corn horticulture styles, there appear to have been many cultures on prompted a population expansion and migration into the central Gulf Coast, and “influences seem to have the greater Mobile-Tensaw River area (Milanich et al. come from all directions simultaneously, as very 1997; Lolley 2003). Local traditions continued, repre- different groups chose to occupy the Mobile Basin” sented by relatively numerous Tates Hammock phase (Brown 2004a). The lower Tombigbee people who (AD 400 to 750) clam shell midden deposits in the worked the salt springs of Clarke County seem rela- Delta and oyster midden deposits on the Bay (Figure tively distinct in terms of pottery and, thus, may have 17-6). Similar sites are known from the subsequent been newcomers from the Alabama and Coosa river Coden (AD 750 to 1100) and possibly Tensaw Lake valleys (Brown 2004a; Dumas 2007). Small Weeden (AD 850 to 1100) phases (Fuller 1998; Brown 2004a). Island sites became common along tributaries in the Florida panhandle around AD 700, and this has led to Summary Archeological research in the greater Mobile-Tensaw River area has created a chronological and spatial framework that can now support more robust anthropological inquiries about early Native American residents. Where researchers have looked beyond basic culture history, at places like Bottle Creek, they are revealing the intricate and complex lives of the region’s original inhabitants.

But there is so much more to be learned. Exciting patterns in the archeological data are materializing that merit intensive scrutiny and the application of new analytical techniques in the years ahead. For instance, evident in the greater Mobile-Tensaw River area are vacillations between periods of great interregional interaction and status building with periods of regionalism and egalitarian social organization, a pattern seen elsewhere in the Southeast. It will be fascinating to search for the reasons for these long- term patterns as they were expressed in southwest Alabama. Viewed from a geographical perspective, the greater Mobile-Tensaw River area appears to have been a prehistoric cultural frontier, a fulcrum on which societies to the east and west met in a precari- ous balance. How did local peoples mediate that frontier? Notions of creolization may be worth applying to such pre-modern culture contact situations. Figure 17-6. The Briar Lake Slough Site (1BA270) prior to excavation in 1998. The tidally inundated shell and The greater Mobile-Tensaw River area’s past affords black-earth deposit dates to the circa AD 400 to 700 por- an ongoing opportunity to address the most contem- tion of the Late Woodland period, and is typical of such porary of issues, as the archeological record has the small habitation sites throughout the Delta. Less obvious are the overlying non-shell midden layers associated with unique ability to let us study human behavior across the Late Woodland Tensaw Lake Phase (circa AD 900 to thousands of years against the backdrop of this partic- 1100) and the much more recent circa AD 1550 to 1700 ular natural environment. Climate change, immigra- protohistoric period. The near invisibility of these non- tion, population growth, resource depletion, and shell layers during boat-based survey projects has caused social inequality are just some of the issues to which them to be under represented in archeological studies. these data can speak. (Photo credit: David W. Morgan.)

Cultural Landscape - Chapter 17 139

18. Archeology and History, AD 1550 to 1950 Gregory A. Waselkov, University of South Alabama

Hernando de Soto’s failed effort to conquer southeast- quickly established military and trade connections ern North America for Spain had broad and long-last- with the Mobilians and the Tomés, whose villages ing repercussions for the region’s indigenous peoples occupied high bluffs on the west banks of the upper and their environment. Even though the climactic Mobile and lower Tombigbee rivers, respectively. battle at Mabila, where Chief Tascalusa ambushed Mobilian and Tomé women cultivated extensive fields and nearly destroyed the conquistadors in 1540, took of maize, beans, squash, and other cultigens on the place probably some distance north of our study area east banks of those rivers, on natural levees border- (Knight 2009), native societies peripheral to the inva- ing the bottomland islands of the delta. However, sion route experienced population declines, collapse as native populations declined rapidly in following of chiefly power structures, an end to earthen platform years from exposure to newly introduced Old World mound construction, and abandonment of many agri- diseases, French colonists reclaimed abandoned cultural fields, towns, and hunting territories. Over fields and the rich soils of former Indian settlements the following century and a half, descendants of the for their plantations (Figure 18-1; Potter and Wasel- Mississippian peoples reorganized their societies and kov 1994; Waselkov 1997; Waselkov and Gums 2000, landscapes in novel ways that enabled them to with- p 7-21). stand later colonizing efforts by Europeans. After an initial tentative settlement on Biloxi Bay, in Evidence of the Earliest Mobilians 1702 the French established Mobile as their colonial capital, at Twenty-seven Mile Bluff on the Mobile Archeological surveys have identified clusters of small River, just downstream from the largest Mobilian sand burial mounds in the upper Delta and adjacent th th village. With its formal European-style town grid uplands that date to the late 16 and early 17 centu- covering about 40 ha, Old Mobile, as it came to be ries (Stowe 1978, 1981, 1982; Brose et al. 1983; Fuller known after abandonment in 1711, was the first colo- et al. 1984; Butler et al. 1997; Fuller and Brown 1998, nial town in the northern Gulf Coast (preceded only p 146-7). Neither these mounds nor their associated by failed settlement attempts by Luna in 1559 and village sites have yet been scientifically investigated. LaSalle in 1685, and by the successful Spanish military However, second-hand Spanish accounts between outpost at Pensacola in 1698). This important archeo- 1675 and 1693 reported many towns crowded into logical site has been intensively studied since 1989 the lower Alabama River area (Waselkov and Gums (Waselkov 1991, 2002, 2005; Moussette and Wasel- 2000, p 7). Among these relocated nations—refu- kov 2013, p 376-92), and has been determined eligible gees fleeing homelands further north to escape slav- by the Secretary of the Interior for designation as a ing raids by interior tribes (Ethridge 2010)—were the National Historic Landmark (NPS 2001). Mobilians, who moved south to the Mobile-Tensaw Delta sometime in the mid-17th century (Knight and The French selected this location for their capital Adams 1981; Lankford 1983; Goddard et al. 2004). for many of the same reasons that led the Mobilians, Mobilians discouraged a Spanish emissary from visit- and before them the Mississippians at Bottle Creek, ing their towns in 1686 due to a drought “so severe to settle near the heart of the Mobile-Tensaw Delta. that although they had cultivated the ground they had With the confluence of 2 major river systems not far secured no crop and all had dried and for many days to the north, this place controlled major canoe routes they had sustained themselves on shellfish” (Boyd into the heart of the continent. And Delta soils were 1937). the most productive, by far, for agriculture in the entire region, since silt naturally deposited by flooding Establishment of the Louisiane Colony annually renewed soil fertility. Less appreciated by the and Old Mobile French was the continuing sacred significance of the When a French expedition arrived on the Gulf Coast Bottle Creek mound site to the Mobilians. In a brazen in 1699 to establish the Louisiane colony, colonists act of desecration, early in 1702 Governor Iberville’s

Cultural Landscape - Chapter 18 141 Figure 18-1. Detail from the French map titled “Carte d’une partie du cours de la riviere de La Mobille et de celle des Chicachas,” attributed to Valentin Devin, circa 1725. This map segment shows a stretch of the Mobile River from Old Mobile (Vieux Fort) at Twenty-seven Mile Bluff to the Mobilian and Apalachee lands near modern day Mount Vernon, with old fields Ancien( desert), flooded fieldsdesert ( jnnondez), villages of the Mobilians and Apalachees, and French settlements at the village St. Hermine and the habitation of Monsieur Parent. (Image credit: Library of Congress, Geog- raphy and Map Collection, Washington DC)

brother, Jean Baptiste Le Moyne de Bienville, paid a religion and, with their church at Mobile and chapels Mobilian a musket . . . at several missions, introduced Roman Catholicism to the region. . . . to show him the place where their gods are. . . . It took a search to locate them on a little Salt and Trade hill among the canes, near an old village that is destroyed, on one of these islands. The gods Salt was another local resource exploited by humans were brought here. They are 5 images—a man, for at least a thousand years, processed from saline a woman, a child, a bear, and an owl—made of springs in the lower Tombigbee River valley. Archeo- plaster. . . . We have them at the settlement. The logical investigations in southern Clarke County at Indians who see them here are amazed at our the upper, middle, and lower salt works, centered boldness and amazed that we do not die as a on a large hill called Salt Mountain, reveal intensive result. exploitation by Late Woodland and Mississippian peoples (Spanos 2006; Dumas 2007; Brown 2009). Iberville determined to take these statues, stolen from Salt becomes a critical resource as subsistence shifts a temple atop one of the small mounds on the periph- to fulltime farming, which probably inspired the agri- ery of Bottle Creek, to France, although he consid- cultural Mississippians to colonize the northern Gulf ered them “not particularly interesting” (McWilliams Coast in the first place (Brown 2004b; Brown 2009, 1981, p 168-9). Despite this sacrilege, the Mobilians p 114-5). Tomés and Choctaws continued to exploit continued to venerate the ancient site, and excavations these salines in the colonial period, as did the region’s there revealed the residence of a native priest occu- 19th-century inhabitants (Figure 18-2). pied well into the 1730s (Silvia 1998; Brown 2003a). Louisiane But the French had symbolically challenged the old Until the French ceded this portion of to the British in 1763, they maintained substantial trade

142 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area societies led many petites nations to seek refuge in the Mobile area, including elements of the Apalachees, Chatos, Towasas, and Taensas. French households held enslaved members of the Chitima- chas, Alabamas, and other distant nations, and French traders ranged widely among the Creeks, Choctaws, Natchez, Tunicas and a multitude of other peoples. In this remarkably diverse linguistic environment, with over 20 native languages, a pidgin known as Mobilian (distinct from the language of the Mobilian people) became the lingua franca for intercultural relations (Crawford 1978; Drechsel 1997).

African Slaves Forced immigration of enslaved Africans, mainly from Senegam- bia, to Louisiane began on a large scale in 1719 (Hall 1992; Berlin 1998), coinciding with the establishment of large plantations by wealthy French colonists. West Africans were particularly sought for their knowledge of indigo and rice cultivation (Wood 1974; Carney 2001), which, along with maize and small acreages of cotton, sugar, and tobacco, formed the basis of crop th Figure 18-2. “Salines, or Salt Springs, along the lower Tombigbee River [in mod- husbandry throughout the 18 ern day Clarke County], as drawn by Thomas Freeman, Surveyor General for century. But plantation econo- Lands South of Tennessee, September 30, 1816” (Lowrie 1834). (Image credit: mies in the region were diverse Center for Archaeological Studies, University of South Alabama.) and relied equally on forest products and cattle raising (another

skill common among West Africans). Later British and with native societies of the region for deerskins, pelts Spanish plantations emulated the earlier French “long of furbearing mammals, and foodstuffs like maize, lot” plantation configuration, stretching inland from a dried fish, venison, and bear oil (Usner 1992; Wasel- waterway to provide each one with river access, flood- kov 2004; Miller Surrey 2006). In fact, Mobile (relo- plain arable soils, and either upland or bottomland cated by the French in 1711 to the area of the modern forest (Figure 18-3; Fabel 1988; Rea 1990). city’s downtown) became increasingly important as a market for the region’s Native Americans and The British Colonial Period remained so for at least 2 centuries (Waselkov and Gums 2000, p 35-44). In the early 18th century, French We are fortunate to have two exceptional travel- willingness to treat and trade with all indigenous ers’ accounts of the region’s natural and cultural landscape from the British colonial period. Bernard

Cultural Landscape - Chapter 18 143 Figure 18-3. Numbered long lot plantations in the Mobile-Tensaw Delta area, detail from “A Plan of part of the Rivers Tombecbe, Alabama, Tensa, Perdido & Scambia in the Province of West Florida,” by David Taitt, circa 1771. (Image credit: Library of Congress, Geography and Map Collection, Washington DC; G3971.P53, 1771.T3 Vault.)

144 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Romans described a 1772 hurricane that devastated by the sovereign Choctaw and Creek nations, within the LaPointe-Krebs plantation, and the great botanist boundaries confirmed by treaties in 1765. William Bartram spent the summer of 1775 exploring the Mobile-Tensaw Delta from his base at the Farmar The Americans established Fort Stoddert (in modern plantation (Harper 1958, p 250-66, 277-8; Wasel- day Mount Vernon) on the Mobile River as their mili- kov and Braund 1995, p 93-8; Romans 1999, p 90-1). tary and administrative seat once Spain transferred the Colonial plantations here were restricted to the west Tensaw and Tombigbee districts to US control. Not side of Mobile Bay, the Mobile-Tensaw Delta, and up until April 1813 did the US Army seize Mobile and the the Tombigbee River to McIntosh Bluff. Archeologi- rest of the modern day Alabama coast from the Span- cally investigated sites include the Rochon-Demouy- ish. Several archeological excavations in downtown Hollinger plantation (circa 1725 to 1848) at the mouth Mobile, particularly at the Gulf Coast Exploreum of Dog River; the Augustin Rochon plantation (circa Science Center, have revealed important aspects of 1750 to 1781) at the edge of the lower delta in modern the city’s colonial and early American occupations day Spanish Fort; the LaPointe-Krebs plantation (Harris and Nielsen 1972; Sheldon and Cottier 1983; (circa 1717 to 1940) in Pascagoula, Mississippi; and Gums and Shorter 1998). Mobile at that time had a the Badon plantation (1763 to 1780) in Blakeley State majority black population, including a substantial Park, on the site of an earlier Apalachee village and number of free blacks (Brown 2001). Completion in mission from 1733 to 1763 (Gums and Waselkov 1997, 1811 of the Federal Road, a military and postal road 2015; Gums 2000; Waselkov and Gums 2000). These from Milledgeville, Georgia, to Fort Stoddert, opened sites confirm the varied basis of colonial economy, the region to massive immigration across the Creek with an abundance of cattle remains, vats for tanning Nation from the southern states (Christopher and cowhides, and pitch and tar processed for naval stores Waselkov 2012). This immigration quickly shifted the from the region’s abundant pines. Live oaks were city’s demographics and strained relations between valued for ship’s knees, and pines for masts and spars, the Creek Indians and their American neighbors. while pine, cypress, and oak were exploited for clap- boards, shingles, and barrel staves. British and Span- The Redstick War and Alabama Fever ish colonists constructed water-powered sawmills In the summer of 1813, amid the War of 1812 (which could double as gristmills and cotton gins), between the US and Great Britain, a civil war in the such as Lizar’s mill near Twenty-one Mile Bluff. Early Creek Nation spilled over the border into Mississippi Americans built others, including Hinson and Kenne- Territory with the destruction of Fort Mims by the dy’s mill on Rain’s Creek near Stockton (Brooms and Redstick faction of the Creek Nation. This conflict, Smith 1996; Waselkov and Gums 2000, p 63-70; Smith known as the Redstick War of 1813-1814 (also known 2008, p 102-8). as the Creek War), transformed the southern social landscape. Within 7 months, US armies invaded the From British to Spanish to US Control Creek Nation from south, east, and north and crushed The greater Mobile-Tensaw River area changed hands the Redstick movement. General Andrew Jackson from British to Spanish control in 1780-1781 with the imposed a 21-million-acre land cession on the Creeks, conquest of British West Florida by Spanish General which opened large portions of Georgia and Alabama Bernardo de Gálvez (DuVal 2015). Each change in to American settlement. Jackson’s success against government increased the region’s ethnic diversity, the Creeks led to his command of US forces defend- as British, then Spanish colonial families joined and ing New Orleans and a victory that propelled him to intermarried with long-established French residents. the Presidency. In 1830, at Jackson’s urging, Congress In 1799 the United States acquired lands north of 31° passed the Indian Removal Act, which forcibly evicted longitude, a line established by surveyor Andrew Elli- most Native Americans from lands east of the Missis- cott that bisected the Mobile-Tensaw Delta. A stone sippi River. So the Redstick War had immense reper- demarcating the Ellicott Line between Spanish Flor- cussions for American history. The site of Fort Mims ida and Mississippi Territory still stands in modern is now a 5-acre state park and has been intensively day Axis. Although ostensibly everything between investigated archeologically (Waselkov 2006; Wasel- that international border and Tennessee belonged kov et al. 2006). to the US, American settlers were in fact hemmed in

Cultural Landscape - Chapter 18 145 Alabama Fever, a land rush complete with boom Native Americans in the Greater towns, followed in the wake of the Redstick War. Mobile-Tensaw River Area East of the lower Mobile-Tensaw Delta, the town of Southern Monroe County soils supported cotton Blakeley briefly competed with Mobile and for a time farming, but elsewhere in the greater Mobile-Tensaw boasted thousands of residents, but declined just as River area forest products and cattle raising (in woods quickly in the wake of a yellow fever outbreak (Stowe and canebrakes) continued to dominate the local 1977). On the lower Tombigbee River, the town of St. economy. Some Choctaws along the Mobile-Washing- Stephens experienced similarly rapid growth. From ton county border managed to avoid forcible removal 1817 to 1819 it served as capital of Alabama Territory, to the west in the 1830s, and their descendants today but declined precipitously with statehood and loss of are organized as the MOWA (Mobile-Washington) capital status to Cahawba (Lewis 2014). Both of these Band of Choctaws (Matte 2002). To the east, from remarkable archeological sites are now state parks, Tensaw to Atmore, a few Creeks received land grants although perpetually underfunded, underdeveloped, for assisting the US military during the Redstick War; and generally underappreciated. one of these tracts formed the nucleus of a reserva- In the 1820s, Mobile’s economy benefited enormously tion, today home to the Poarch Band of Creek Indi- from the rise of cotton plantations on the Black ans, a federally-recognized tribe (Paredes and Knight Belt soils of central Alabama, reflected in the prolif- 2012). eration of cotton warehouses near the city’s water- Throughout the 19th century, Native Americans in the front (Gums and Shorter 1998; Gums 1999; Shorter greater Mobile-Tensaw River area continued carrying and Mattics 2006). Steamboats on the Alabama and forest resources to the Mobile market, while main- Tombigbee rivers transported vast quantities of cotton taining many traditional subsistence practices. In downstream to Mobile (Figure 18-4), for export to 1875, Dr. Gideon Lincecum published an account of Europe, and were only challenged decades later by mass capture fishing by Chickasaw Indians in August railroads (Cline 1997, p 30-64; Fickle 2014, p 33). The 1828, during low water level above the Paineyigaby region’s maze of waterways and swamps occasionally shoal on the Tombigbee River. Although north of our hid runaway slave camps; militia destroyed one such study area, this sort of fishing was practiced through- maroon settlement on Hal’s Lake in 1827 (Anony- out the region for millennia. According to Lincecum, mous 1827; Riley 1906; Diouf 2014, p 153-5). the men went into the woods to dig buckeye (Aesculus rubra) roots, which they pounded and grated. An hour before sunset they carried baskets of root into the water, “plung- ing, or rather churning these violently as they moved along, [and] distributed the pulver- ized root everywhere the whole length of the deep hole.” Within an hour, fish “were visible everywhere on the surface of the water; not dead, but feebly moving about on their back.”

Over a 24-hour period, thousands of fish were Figure 18-4. “Scene on the Alabama River, Loading Cotton.” From Ballou’s Pictorial 13(22), taken from the river p 1, November 28, 1857. (Image credit: Center for Archaeological Studies, University of pool, split open, and South Alabama.)

146 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area dried over fires kindled beneath low wooden racks. (Bergeron 1991; Stowe 1977; Shorter 2001, 2008; Lincecum (1875) related: Sledge 2015, p 132-9).

It was a little curious to observe the different Following the Civil War degrees of power among the fish to resist the deleterious effects of the poison. The perch Post-war Mobile witnessed a redistribution of popu- and small scale fish came first, then the trout lation as freed urban and plantation slaves found new family, suckers, all the scale fish. Then the blue means of employment and modes of housing. Histori- cats, then the big, fat yellow cats, then the mud cal and archeological research has revealed little physical segregation of white and black residences in late- cats and eels, and, lastly, the soft shell turtle and th loggerheads [alligator snapping turtles]. . . . I did 19 century Mobile (Gums 1998; Wilkie and Shorter not know before that a deep hole in the river 2001; Wilkie 2003; Shorter et al. 2007). could be so densely populated. In 1873, the US Army converted Mount Vernon Arse- nal to Mount Vernon Barracks, where Geronimo and Impact of the Civil War 450 other Chiricahua Apaches were held as prisoners In 1830, the US Army established Mount Vernon of war from 1887 to 1894. The property was trans- Arsenal near the site of old Fort Stoddert. Just prior ferred to the State of Alabama, which repurposed to Alabama’s secession from the Union in 1861, state the buildings in 1905 as a mental hospital for Afri- militia seized the arsenal and Forts Morgan and can-American patients. In 1906, Dr. George Searcy Gaines at the mouth of Mobile Bay. The schooner reported an outbreak of pellagra among patients Clotilda, last slaving ship to enter the US, had slipped into the bay 6 months earlier, offloaded her human cargo, and been scuttled near Twelve-Mile Island— one of numerous shipwrecks awaiting discovery in the region’s waterways. Many of those Africans, Yorubas from Dahomey, lived out their lives in a post- war community called Africa Town, on land they purchased in north Mobile (Roche 1914; Diouf 2007; Robertson 2008; Sledge 2015, p 114-5, 261-5).

During the Civil War, as the federal blockade of southern ports became increasingly effective, thousands of workers, enslaved and free, took up residence at the saline springs in southern Clarke County, processing salt for the state and for personal use (Figure 18-5; Lonn 1933). Likewise, as imported coffee, chocolate, and Asian tea became nearly unobtainable, caffeine- bearing leaves of native yaupon holly (Ilex vomitoria) were harvested throughout the southern coastal plain and brewed into “Black Drink” (Hudson 1979, p 7).

War took a long time to impact Mobile directly. The submarine H.L. Hunley was built in the city, far from battlefields, and taken by rail to Charleston, where it sank a blockade ship. Eventually though, in August 1864, Confederate-held forts at the mouth of Mobile Bay fell to a federal invasion force. Still, the city’s 3 robust lines of fortification discouraged a direct Figure 18-5. Partially excavated Civil War-era furnace Union assault, so US forces marched north along the with steamship boiler converted for brine boiling, Fur- bay’s eastern shore to besiege entrenched Confeder- nace AA, Upper Salt Works, Clarke County, Alabama, ate troops at Spanish Fort and Blakeley, which fell in 2010. (Photo credit: Ashley Dumas, University of West Alabama.) April 1865 in two of the final battles of the Civil War

Cultural Landscape - Chapter 18 147 trucks are evident throughout the site.

Swamps called for different methods of tree harvest. Oxen proved useful there, too, until largely replaced by mechanized pullboats— barges with winches that hauled huge gum and cypress logs to waterways. Radial patterns visible in aerial and satellite views of the Delta are deep trenches left by pullboats, perma- nents scars that altered the topography and surface hydrology. Logs were also floated to waterways by Figure 18-6. Four-yoke team of oxen pulling a high-wheel cart, or log cart, hauling damming or channeling a large clamped log from a longleaf pine forest near Mobile, Alabama. Postcard printed in Germany for Raphael Tuck & Sons, Post Card Series No. 2248; published in small tributaries. Robert the United States by G. Van Antwerp & Son; postmarked May 8, 1907. (Image credit: Leslie Smith’s account of Center for Archaeological Studies, University of South Alabama.) high-water logging and raft- ing in the Mobile-Tensaw and correctly recognized their maize-based diet as a Delta is an invaluable firsthand description of a way contributing factor to this nutritional disease, consid- of life that ended just after World War II (Smith 2008, ered to that point a psychiatric disorder (Rajakumar p 42-53, 115-30). Tupelo gum and cypress logs, which 2000). Initially called Mount Vernon Hospital for the float, were bound together in rafts, while non-floating Colored Insane, and renamed Searcy Hospital in 1919, logs of oak and red gum were suspended beneath the facility was integrated in 1969, and closed by the pontoon “gunboats.” Watermen guided them, via river state in 2012. This virtually intact early-19th century current, to downstream mills in and around Mobile. US arsenal and nationally significant historic site remains closed, as buildings deteriorate and collapse. Turpentine distillation from pine resin was a major industry in the upland longleaf and slash pine forests The Logging Economy of the region from the 1850s to 1950s. At first, resin Forests of the uplands and bottomlands of the greater was collected in triangular troughs or “boxes” axed Mobile-Tensaw River area provided the basis of most into tree trunks and cleaned out monthly by dip iron. residents’ livelihoods until the mid-20th century, and to Trees exploited in this manner were vulnerable to a lesser extent they continue to do so today. Evidence disease and fire. Adoption after 1900 of the French of past logging is everywhere, in the region’s altered cup and gutter system, in which resin flowed from vegetation, land surfaces, and soils. In upland forests, grooved bark into ceramic or metal cups, revolution- teams of oxen hauled pine logs by high-wheel carts ized southern turpentine production (Figure 18-7) (Figure 18-6) to a proliferation of short-spur rail lines, (Sargent 1884, p 202, 516-8; Smith 2008, p 255-6; both narrow gauge and standard gauge. From there Fickle 2014, p 36-7, 81). the logs traveled by steam locomotive to sawmills from More recently, forestry has remained important, with 1875 to 1930, when trucks began replacing carts and local mills specializing in paper products, preserva- rails (Burnette 2006, p 269-75; Smith 2008, p 29-41, tive-treated wood, plywood, and laminates. In addi- 116, 123-4, 265-7; Fickle 2014, p 73, 87-8, 92-106). tion, the region’s forest-dwelling animals continue At the northeast end of the Old Mobile archeologi- to be intensively hunted, for subsistence and sport cal site, one can still discern the raised bed of a circa (Jordan 1999; Smith 2008, p 15-8, 96). Deer and 1900 “dummy line” that serviced a logging camp over- bears were extirpated early in the 20th century, but looking the Mobile River; soils compacted by logging

148 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area deer returned during World War II, and bears were On-going Connections reintroduced recently. Hunting clubs and wildlife A multitude of connections continue to tie the human management areas established in the early- and mid- population of the greater Mobile-Tensaw River area th 20 century are scattered throughout the middle and to the region’s natural resources. One need only recall upper portions of the study area, and recreational fish- the baptisms that once occurred in local springs and ing, canoeing, and kayaking draw substantial numbers streams (Jordan 1999, p 45, 99) to realize the depth of of people into the Delta. the bonds linking one to the other.

Figure 18-7. Turpentine gathering containers and dip iron, collected by Robert Leslie Smith from the Mobile-Tensaw Delta. The log is a longleaf pine stump with an axe-cut turpentine collecting box. (Photo credit: Center for Archaeologi- cal Studies, University of South Alabama.)

Cultural Landscape - Chapter 18 149

19. Modern Cultures of the Mobile-Tensaw Delta Ben Raines, Independent Environmental Reporter

Ride through the Mobile-Tensaw Delta any given day speckled trout fishermen number in the thousands, a and you will encounter pleasure seekers of all manner, virtual armada of center console fishing boats drift- from birdwatchers and duck hunters, to fishermen, ing across the big bays, choking the Raft and Tensaw kayakers and water skiers. But in the mix out on the rivers, or surrounding the Battery at the junction of water, you’ll also find a lot of people making money the Apalachee and the Blakeley. Come winter, the off the Delta. Commercial boats ply the lower Delta duck hunters appear in camouflage boats, hunting rivers for live shrimp for the bait shops that line the US from blinds built from the tall Roseau cane (Phrag- 98 Causeway. Restaurant owners run catfish box traps mites australis) that lines the inner creeks. Spring out of johnboats for the fresh Alabama River channel means professional bass tournaments and birdwatch- cats they will fry up and sell that night. Fishing guides ers numbering in the hundreds. And in the summer- number in the dozens. And most days, you are likely time pontoon boats and jet skis raft up around to see one of the big tour boats carrying 40 paying Gravine Island in the heart of the Delta. Hundreds of customers up river to see the sights. people crowd up on its sandy beach, boiling pots of crawfish, drinking beer, and getting sunburned while If your eyes are keen, you might also spy a few modern Sweet Home Alabama booms out of a ski boat stereo day bandits, perhaps poaching turtles from Big Briar, system. The beach makes a tall hill, about 15 ft (5 m) or scraping through Native American sites for artifacts high. Revelers up on top shout warnings to swimmers to sell on eBay. Game wardens routinely arrest scoff- if an alligator makes an appearance nearby. laws in the Delta, sometimes illegally netting mullet under cover of night in the Delta’s rivers, or even kill- When the US Army Corps of Engineers bought up ing anhingas for their elaborate tail feathers. much of the lower Delta in the 1980s, it changed the complexion of the place in some ways. Numerous The People of the Delta people who were living full time in the Delta were The Delta, in many ways, is the same as it has always forced out when it was discovered they did not have been: part playground, part natural bounty waiting legal title to the marshy land beneath the ramshackle to be reaped and brought to market. The players may homes they had built. The federal purchase, coupled have changed over the years—for instance, the logging with the arrival of modern game and fish regulations, crews are gone and no one grazes cattle on the islands hastened the end of the Delta’s resident trapper/fisher anymore—but the landscape that has supported vari- population, people who had eked out a living entirely ous human endeavors for millennia remains. from what they could pull from the great swamp. Even so, there are likely more people out in the Delta on a Most of the people you see in the Delta are locals, and day-to-day basis today than at any point in its modern most of them have grown up visiting the big swamp history. with their families. For those unfamiliar with navi- gating the Delta’s twisting and shoal-ridden waters, Stories from Delta Locals making that first trip into the giant swamp can be Lucy “Pie” Hollings has been watching the world forbidding. Even more than the alligators and snakes, drift by her “little piece of paradise” on the Tensaw many fear getting hopelessly lost, or breaking down River on the Delta’s eastern edge her entire life. She deep in the interior. There are countless boat owners inherited Cloverleaf Landing, where she charges $4 to in the area surrounding the Delta who say they prefer launch a boat, from her father. Most days, she can be the open waters of Mobile Bay, where navigation is found sitting on her houseboat next to the ramp. The easy, and help is never far. Delta has defined her life. As she says,

The type of folks you’re likely to meet in the Delta I don’t want for anything. Everything I’ve ever depends largely on what time of year it is. In the fall, needed was right here in the Delta. I have fish

Cultural Landscape - Chapter 19 151 to eat, crabs. People bring me things when they Sitting in the bait shop where his son now works, come to the boat ramp. It’s just a wonderful life. Larry went on: My foreparents was Indians. My grandmama was Cherokee. So I enjoy living off the land. It’s about all we’ve ever known and it has been Very seldom go to the store and buy food. Don’t good to us. A lot of people would die to have have to. our sunsets every afternoon, overlooking Mobile, same thing with sunrises when we’re Her brother, Sylvester Crooks, hunts wild hogs in out shrimping. It doesn’t get much better. the Delta. The hogs, domesticated pigs turned feral, number in the hundreds of thousands in the Delta, Jimbo Meador likes to say he grew up in the Delta by most estimates. With teeth to rival a German shep- and it taught him everything he ever needed to herd, and the ability to reproduce when they are know. Meador earned pocket money as a teen trap- just 6 months old, the invasive animals cast a long ping around the edges of the Delta and its adjoining shadow over Delta life, consuming everything from swamps. In 1957, at age 15, he entered and won one baby deer and ducks to the tender roots and tubers of the first nutria rodeos in the nation, killing the larg- of the wetland plants that hold the soft Delta mud in est nutria in the tournament. Even today, at 73, he still place. Hog hunting has emerged as a major sport in spends long nights gigging frogs in Little Bateau, his the Delta, with several butcher shops in the area now favorite of the Delta’s inner lakes. Meador now runs specializing in dressing the wild pigs. 17 Turtles Outfitters, which specializes in providing guided trips in the Delta, and renting kayaks and Crooks doesn’t use a gun when he hunts. Just his dogs stand-up paddleboards to people who want to explore and a knife. The dogs run the pig down and corner it the area. For him, the Delta provides a place of soli- somewhere, and then Crooks jumps on its back and tude amid the technology that surrounds modern life. slits its throat. He won’t let anyone accompanying him In Jimbo’s words: bring a gun, he says, for fear they will shoot one of his prized dogs. I spent most of my life over there. Grew up in the Delta. I’ve seen what’s transpired. One of Larry Scott and his twin brother Barry own Scott’s the reasons I’m doing what I’m doing [the kayak Landing, a boat launch and bait shop on the edge of business] is because I want to make more people the Causeway. The shop was owned by their father aware of the treasure we have. People pass it on when the Scott boys were young, but sold in their the Interstate, but nobody really knows what’s teens, when the name was changed to Mizell’s Land- out there. I’ve seen it change. ing. The Scott boys bought it back decades later. Fish- ing the Delta, particularly the Blakeley River, has been Over the years, I was obsessed with duck hunt- a way of life for their family for 4 generations. Larry, ing over there. I ran a trap line and sold furs. I who lives on a houseboat behind the bait shop with shrimped. I did crabbing. I’ve done a whole lot various pets, including a raccoon, relates: of fishing there. Frog gigging. I don’t want to say it, but I used to hunt alligators, and I used We were fishing from the time we were real to eat them. The snipe hunting. Oh gosh, I love small. Our grandfather had a seafood market the Delta for snipe hunting. I had trot lines up in Mobile, and my father shrimped and fished there, caught catfish, bream fishing, bass fishing, for him. By the time we were 4 or 5 years old, speckled trout, redfish, sheepshead. The beauty Daddy had us in a boat. of it is you got freshwater fishing and saltwater fishing. I have caught bass, speckled trout, In the 1970s, the Scotts’ father nearly froze to death flounder and redfish all in the same spot, on the while trying to run a commercial gill net on one of same bait, on the same morning. the coldest days on record in south Alabama. After his boat ran aground, the elder Scott tried to wade “Delta Rats” to shore, but collapsed due to hypothermia. When he was finally rescued, lying in the shallows, his body There’s a sort of fraternity of Delta lovers, people who cocooned in a layer of ice, his eyeballs were frozen in call themselves “Delta rats.” They might be commer- place, staring straight ahead. Miraculously, he made a cial shrimpers, or water skiers, or recreational trout full recovery and went back to fishing. fishermen. It doesn’t matter. They are people who love

152 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area to be out in the Delta, and their diverse backgrounds two leading candidates for US Representative for offer proof that the Delta is big enough, and change- District 1 spent more time talking about the future able enough, that just about anyone can find whatever of the Mobile-Tensaw Delta than any other topic. At they may be looking for. issue was the question of whether the Delta might one day become a national park, a proposition backed by I, personally, have fallen under the Delta’s spell. For former Representative Jo Bonner before his resigna- the last 15 years I’ve worked my way deeper and tion. Both men competing for Bonner’s seat were deeper into its interior, and gotten to know more adamantly opposed, arguing that the Delta belongs to and more of these self-described Delta rats. I learned the people of south Alabama, and it would amount to my way around by running aground and getting lost, a horrible crime to allow federal officials any domin- the best way according to the old timers, who warn ion over the precious wetland. that the Delta is not easy. GPS [global positioning system] is unreliable because the shorelines change There remains a local group campaigning for some sort and reshape with every spring flood. Getting lost and of federal status for the Delta, perhaps as a national finding your way out teaches you a certain respect for preserve, which would still allow unrestricted hunting the place, they say. And it teaches you to pay attention. and fishing, among other activities. In the eyes of those pushing for federal recognition, such protection could Now I guide people into its inner reaches, to places prove vital to the long-term viability of the Delta’s rich seldom seen. I look forward to July, when the giant habitats. They cite two remarkable but opposing facts yellow flowers of the American lotus bloom, tens of about Alabama and its wild environs. thousands of golden heads nodding in the summer breeze. And to October, when the trout and redfish First, the state leads the nation in aquatic diversity, run up the rivers in great numbers, and it is possible with more species of fish, mussel, snail, and crayfish to catch 40 or 50 before going to work in the morn- than any other state. Most of those creatures can be ing. I love February, when the entire Delta disappears found in the rivers of the Mobile Basin, which drain beneath 10 ft (3 m) of water in the spring flood and into the Delta. The second often cited truth about you can drift through the forest amongst the trees, Alabama is that the state ranks as the king of extinc- floating over places where you’ll be walking on dry tions for aquatic creatures, responsible for roughly ground in a few months. I’ve learned where I can go in half of all extinctions that have occurred in the conti- May and play a recording of a Prothonotary Warbler nental US since the 1800s. and call 2 or 3 of the tiny, canary yellow birds right to my boat. Obviously, a state cannot forever wear both the crown for most aquatic diversity and the crown for I see the same faces at the boat ramp again and again. most extinctions. Unfortunately, the current political We trade notes on what’s happening where—birds climate in the state suggests the scale may be tipping diving over schools of menhaden at the mouth of toward more extinctions. The Alabama Legislature Crab Creek; a new log down on the way into Jessa- has provided zero funding to the state’s environmen- mine Bayou; a colony of Yellow-crowned Night tal agency, the Alabama Department of Environmental Herons nesting in One-Mile Creek—the way people Management, for the last several years. And Alabama talk about old friends. The boat ramp crowd includes ranks dead last among the 50 states in terms of money doctors, lawyers, shrimpers, mechanics, teachers, spent on environmental protection. In many ways, the artists, and commercial fishermen, because the Delta future of the Delta rests squarely on the backs of the as the common thread makes for easy friendships. The people who use it most: the kayakers, the fishermen, big swamp has defined the communities that surround the birders, the water skiers, the ecotour guides, and it, in both Mobile and Baldwin counties. And it is the the charter captains. central focus for recreation for thousands of people living in those counties, year round. For all the majesty of the Mobile-Tensaw Delta, you often hear the same lament from old timers: “You Looking to the Future should have seen it then.” Often the losses are plain to see, from bays in the lower Delta taken over with inva- Perhaps the greatest testament to the importance of sive milfoil or filled with dredge spoil, to duck hunters the Delta as a cultural touchstone for south Alabama returning to the dock with no ducks on cold winter was revealed in the last congressional election. The

Cultural Landscape - Chapter 19 153 mornings. But to focus too much on those voices Finch B, Young BM, Johnson R, Hall JC. 2012. from the past ignores the fecundity of the present. It Longleaf, far as the eye can see: a new vision of is possible to venture out most fall mornings in the North America’s richest forest, with a foreword Delta and catch 50 speckled trout before 9 AM, some- by EO Wilson. Chapel Hill NC: University of thing possible in only a few places on Earth. Likewise, North Carolina Press. birders heading out during the spring migrations often add dozens of species to their life lists in a single Gosse PH. 2013. Letters from Alabama, chiefly morning. The Delta is like that, so rich you can’t help relating to natural history, edited by GR Mullen but be impressed. and TD Littleton. Tuscaloosa AL: University of Alabama Press. I often think of the Delta the way a man at a boat ramp described the fishing one morning. “Just put some- Key W. 2015. Among the swamp people: life in thing wiggly on a hook and you’ll catch something,” he Alabama’s Mobile-Tensaw River delta, with said. “The Delta’s easy, no matter what you’re after.” illustrations by K Mercer. Tuscaloosa AL: University of Alabama Press.

Suggested Readings Nation F. 2002. Where the wild Illicium grows: Bartram W. 1998. The travels of William Bartram, historic plants of south Alabama and the central naturalist’s edition, edited by F Harper. Athens Gulf coast. Bay Minette AL: Baldwin County GA: University of Georgia Press. Historical Society.

Duncan RS. 2013. Southern wonder, Alabama’s Smith RL. 2008. Gone to the swamp: raw materials surprising biodiversity. Tuscaloosa AL: for the good life in the Mobile-Tensaw delta. University of Alabama Press. Tuscaloosa AL: University of Alabama Press.

154 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 20. National Historic Landmarks David B. Schneider, Schneider Historic Preservation; Bonnie L. Gums, University of South Alabama; Gregory A. Waselkov, University of South Alabama; George W. Shorter, Jr., retired archeologist, Mobile, Alabama; Richard S. Fuller, retired archeologist, Mobile, Alabama

National Historic Landmarks (NHLs) are places objection, 1 of 5 such properties in the country. The deemed nationally significant by the US Secretary of National Park Service is tasked with reporting on the Interior for their exceptional value in interpret- the conditions of landmarks through periodic status ing the historical heritage of our country. Slightly updates provided by owners of National Historic more than 2500 sites are so recognized nationwide, Landmarks. This chapter summarizes status updates compared to about 85,000 historic properties listed on the 6 National Historic Landmarks in the greater on the National Register of Historic Places, a sister Mobile-Tensaw River area, as compiled by the authors program also administered by the National Park in a lengthier report (Schneider et al. 2015). Service. One might legitimately characterize National Historic Landmarks as the best of the best. National Bottle Creek Site (Archeological Site Historic Landmarks include both privately and publi- 1BA2; 1994 NHL Designation) cally owned properties, and participation in the Bottle Creek, located on Mound Island in the Mobile- program is voluntary. Like National Natural Land- Tensaw Delta, is the largest Mississippian archeo- marks, another Department of the Interior program logical site on the north-central Gulf Coast, from the that recognizes our country’s outstanding biological Florida Panhandle to the Mississippi River (Figures and geological places, National Historic Landmarks 20-1 and 20-2). One of Alabama’s most significant are so designated to promote the preservation and protection of these gems, and more generally to raise public awareness of our nation’s historical heritage.

There are currently 6 National Historic Land- marks in the greater Mobile- Tensaw River area: Bottle Creek Site (Archeological Site 1BA2), Government Street Presbyterian Church, Fort Morgan, Mobile City Hall (Southern Market), the USS Alabama, and the USS Drum. A 7th place, the Old Mobile Site (Archeological Site 1MB94), has been determined eligible for National Historic Landmark status, but has not been so desig- Figure 20-1. Location of Bottle Creek Site National Historic Landmark, Baldwin County, nated because of landowner Alabama (7.5-minute Stiggins Lake, Alabama, 1983 USGS quadrangle).

Cultural Landscape - Chapter 20 155 Figure 20-2. Topographic map of Bottle Creek (Waselkov 1993). (Image credit: Sarah Mattics, Center for Archaeological Studies, University of South Alabama.)

prehistoric sites, second in size only to Moundville, 2012). Those explorations uncovered prehistoric and Bottle Creek served as a social, political, religious, and historic Indian house floors, as well as evidence of trade center for a society named by archeologists as mound construction methods. Excavation trenches the Pensacola culture, part of the Mississippian tradi- into Mounds A and B documented the sophisticated tion, from around AD 1200 to 1550. The site is owned engineering involved in their construction (Brown by the State of Alabama and administered by the 2003a). On November 30, 2014 Richard Fuller discov- Alabama Historical Commission. ered a previously undocumented mound, inspired by a search of LiDAR imagery by archeologist Jason There are 19 documented earthen mounds at Bottle Gardner (Fuller 2014). Based on these various inves- Creek. Most are platform mounds that would have tigations, Bottle Creek is considered one of the most held wooden temples and dwellings. Five of the significant archeological sites in the north-central mounds, including the largest and tallest (at over 45 Gulf Coast region. ft [14 m]), Mound A, encircle an open plaza or public space. Apart from a few outlying mounds, the others The site is monitored by officers from the Alabama line the wetlands on the north and west edges of Department of Conservation and Natural Resources, the site. The village site around the mound complex who check for evidence of looting or other distur- covers at least 46 acres (0.2 km2). Shell middens repre- bances. There has been little recent evidence of loot- senting food remains of the residents occur through- ing for artifacts, but rooting by wild hogs is damaging out the site, and there are at least 3 borrow pits dug for the site surface, a fairly recent phenomenon. Damage soil to build the mounds. to trees has occurred from hurricanes, most recently with Hurricane Ivan (2004) and Hurricane Katrina The mounds at Bottle Creek were first mapped in (2005). Heavy rains from hurricanes and tropical 1853 (Bigelow 1853), followed by limited excavations storms can erode the earthen mounds, as happened in in 1905 and in 1931 to 1932 (Sheldon 2001). In 1990, the mid-20th century when Mound A suffered a major staff from the University of South Alabama’s Center slump on its western face. for Archaeological Studies and volunteers from the University of Alabama, University of Southern Missis- Historic Blakeley State Park, Five Rivers, and private sippi, and Auburn University-Montgomery created ecotourism companies offer boat trips to Bottle Creek a modern topographic map of the mounds at Bottle that bring several hundred people to the site annually. Creek (Waselkov 1993). Between 1991 and 1994, the Maps of the local waterways distributed at fish camps, University of Alabama’s Gulf Coast Survey conducted bait stores, and other outlets mark the site, which is the first intensive excavations at Bottle Creek, directed also a highlight on the Delta’s Bartram Canoe Trail, by Ian Brown (Fuller and Brown 1998; Brown 2003a, so many people find their own way to the mounds.

156 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area As the site has become better known in recent years, distyle-in-antis façade, the building is noted for its evidence of illicit digging on the site has declined, intact Greek Revival-style interior. The sanctuary has which suggests that higher public visibility and more seen only minor modifications since original construc- visitors may deter vandalism and looting. tion in 1838. This privately owned church is an active religious facility and is open to the public. Bottle Creek is one of the few major Mississippian sites in the southeastern United States that has not Government Street Presbyterian Church is in excel- been developed for tourism, due to its isolated loca- lent condition, little changed in appearance since tion. This heavily forested site in the Mobile-Tensaw construction. The building was last rehabilitated in Delta gives visitors a sense of the rich natural environ- 2002 with new paint, carpeting, rewiring, and instal- ment that supported Native American societies for lation of a new organ, in conformation with the Secre- thousands of years. tary of the Interior’s standards (Morton et al. 1997). Government Street Presbyterian Church is very Government Street Presbyterian well maintained, with an active program of cyclical Church (1992 NHL Designation) maintenance and restoration overseen by a property Located at 300 Government Street in Mobile, manager and building committee. The city’s historic Government Street Presbyterian Church is significant preservation ordinance requires prior approval of any as one of the oldest and least-altered Greek Revival- proposed exterior work. style houses of worship remaining in the United As Mobile is prone to hurricane and tornado activ- States today (Figures 20-3, 20-4, 20-5) (Hamlin 1944; ity, weather poses the primary threat to Govern- Kennedy 2010). Along with a distinctive monumental ment Street Presbyterian Church. Hurricanes have

Cultural Landscape - Chapter 20 157 Figure 20-4. South façade detail, Government Street Presbyterian Church, Mobile, Mobile County, Ala- bama. (Photo credit: David B. Schneider, March 2015.)

Figure 20-5. Interior sanctuary, Government Street Presbyterian Church, Mobile, Mobile County, Alabama. (Photo credit: David B. Schneider, March 2015.)

158 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area damaged the church in the past and are likely to be a 20-9) (Bergeron 1991; Hearn 1993). Fort Morgan was threat in the future. The surrounding urban area has constructed between 1819 and 1834 as a pentagonal, changed considerably since the building’s construc- masonry, coastal defense fortification. Extensively tion, with most of the adjacent historic streetscapes damaged during the Civil War, when a central citadel already lost prior to landmark designation. Govern- was destroyed, the fort was remodeled in 1895 to 1898 ment Street Presbyterian Church now exists within an with reinforced concrete gun emplacements. Frame essentially modern urban setting. At the present time, barracks, officers’ quarters, hospital, and other struc- Government Street Presbyterian Church is in excel- tures were added when the fort was reactivated as a lent condition, is well maintained by ownership that Spanish-American War training base. All but 4 frame recognizes and appreciates it historical and architec- structures have been lost to hurricanes after landmark tural significance, and the overall threat level is mini- designation in 1960. mal, except as already noted for the unpredictable nature of the region’s extreme weather events. Fort Morgan was recorded in 1937 as archeological site 1BA186. After World War II the state-owned Fort Morgan (1960 NHL Designation) military reservation became a park overseen by the Alabama Department of Conservation, then by the Located at the entrance to Mobile Bay, in Baldwin Alabama Historical Commission since 1977. Prior County, Fort Morgan played a significant role in the to state ownership, many individuals removed arti- 1864 Civil War battle of Mobile Bay, when Admiral facts from Fort Morgan by careless digging and metal Farragut’s Union Navy fleet ran past the fort to seize detecting. Most of the fort’s cannons were sold for control of the bay, losing the ironclad USS Tecum- scrap in the early-20th century (England et al. 2000, p seh to a mine in the process (Figures 20-6 through 48).

Figure 20-6. Location of Fort Morgan National Historic Landmark, Baldwin County, Alabama (7.5-minute Fort Morgan, Alabama, 1958 USGS quadrangle).

Cultural Landscape - Chapter 20 159 Limited amateur and professional archeological investigations have occurred at Fort Morgan over the last 50 years. In 1970, 3 of the fort’s 4 cisterns were excavated (Keebler 1970). A 1979 survey of Mobile Point, where Fort Morgan sits, recorded fort-related surface features (Stowe et al. 1979), and a Troy State University team investigated the drainage system between casemates in 1991. In 1993 archeologists surveyed an area north of the park’s museum prior to relocation of the historic Figure 20-7. Entrance at north rampart, Fort Morgan, Baldwin County, Alabama. 1872 metal lighthouse. (Photo credit: David B. Schneider, March 2015.) Expeditions 21 and 22, educational programs organized by the Alabama Museum of Natural History, were held at Fort Morgan in 1999-2000, directed by University of South Alabama archeologist George Shorter (Shorter 2001). These extensive excavations focused on Battery Dupor- tail and the Citadel that once stood in the center of the fort. Massive brick remains of the Citadel and a large privy pit yielded Civil War munitions, such as a wooden sabot and a Federal 3-inch (8-cm) Schenkl shell, as well as medicine bottles, uniform buttons, and smoking pipes. Archeological testing by Panameri- can Consultants in 2008 of the fort’s terreplein and parapet revealed a clay surface of the original War of 1812 fort (Carruth 2008). These investigations amply demonstrate the existence of significant archeological deposits throughout Fort Morgan.

Fort Morgan is generally in good condition and is subject to a continual program of maintenance and restoration consistent with the Secretary of the Inte- rior’s standards (Morton et al. 1997). In recent years, restoration has concentrated on reconstruction of the east rampart and addressing the fort’s continuing drainage problems. The surviving frame buildings at Fort Morgan and concrete artillery batteries are stable. An iron lighthouse installed in 1872 was sold to a scrap dealer in 1966, only to be returned to the fort in 1991, then again dismantled in 2003 for restora- Figure 20-8. Interior vaulted casemates, Fort Morgan, tion. Funding has not been available to complete the Baldwin County, Alabama. (Photo credit: David B. project. Schneider, March 2015.)

160 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Fort Morgan retains most of its essential historic design elements. Historic fabric and materials are respected during ongoing maintenance and restoration work and original workmanship of Fort Morgan remains visible. The setting has been compromised to some degree by gas wells in Mobile Bay and the Gulf of Mexico that are readily visible from the fort. Fort Morgan is well maintained by the Alabama Historical Commis- sion, which recognizes and appreciates its historical and architectural significance. The overall threat level is minimal, Figure 20-9. Officer’s quarters, Fort Morgan, Baldwin County, Alabama. (Photo credit: except for the unpredictable David B. Schneider, March 2015.) nature of the region’s extreme weather events. Ongoing restoration efforts are attempting to address chronic Montuse’s tavern (circa 1815), and 1830s cotton ware- water penetration that adversely affects the masonry houses and commercial buildings (Gums and Wasel- structure, which remains an ongoing conservation kov 1997; Gums and Shorter 1998, 2000). concern. The old Mobile City Hall buildings were rehabilitated Mobile City Hall (1973 NHL in 2000 to 2001 to serve as the city’s history museum. Designation) This project resulted in some alterations, but was generally consistent with the Secretary of the Interi- Long known as the Southern Market and Municipal or’s standards for rehabilitation (Morton et al. 1997). Building, Mobile City Hall, constructed in 1855 to The essential character-defining elements of the th 1857, exemplifies a mid-19 -century trend to combine buildings’ design, materials, and workmanship remain multiple civic functions in public structures (Figures intact and were enhanced through appropriate repair 20-3, 20-10, 20-11) (Holmes 1987; Ewert 1993). This and restoration work. complex of 4 Italianate, stuccoed brick buildings links smaller flanking structures to the main gabled build- At the present time, the buildings are in excellent ings with crenellated, arched wing walls. The lower condition, are well maintained by ownership that floors of the buildings on South Royal Street housed recognizes and appreciates their historical and archi- a market, with upper floors for municipal offices. By tectural significance, and the overall threat level is the mid-20th century the entire complex was occupied minimal except for the unpredictable nature of the by City Hall and continued to serve as such until the region’s extreme weather events. The structures buildings were severely damaged by Hurricane Fred- suffered substantial damage from flooding during eric in 1979. Hurricane Katrina in 2005, which necessitated extensive repairs. The surrounding urban area has Mobile City Hall was built on a site that straddles the changed considerably since construction and most colonial era riverfront. Between 1997 and 2000, two of the surrounding historic elements of the adjacent major archeological investigations of the site, desig- streetscapes had been lost prior to landmark designa- nated 1MB189, uncovered deposits up to 5 m (16 ft) tion. Mobile City Hall is now, and was at time of desig- deep that date to the 18th- and early-19th centuries. nation, a historic building complex that exists within These deposits relate to the northeast bastion of Fort an essentially modern urban setting. Condé, a Spanish colonial residence (circa 1800),

Cultural Landscape - Chapter 20 161 Figure 20-10. Mobile City Hall, Mobile County, Alabama. (Photo credit: David B. Schnei- der, March 2015.)

Figure 20-11. Interior first-floor lobby of the History Museum of Mobile, in Mobile City Hall, with 1936 Works Progress Administration murals by John Augustus Walker. (Photo credit: David B. Schneider, March 2015.)

162 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area USS Alabama (1986 NHL Designation) USS Alabama is a South Dakota-class battleship (designated BB-60 by the US Navy), commissioned in 1942, that spent 40 months in active service during World War II, participating in 26 engagements (Figures 20-3, 20-12, 20-13, 20-14) (Sumrall 2001; Arnhart 2007). The ship was decommissioned in 1947 and transferred to the State of Alabama in 1964 for use as a museum (although the federal government retains ownership). The battleship is administered by the Figure 20-12. The USS Alabama. (Photo credit: David B. Schneider, March 2015.) USS Alabama Battleship Commission, which operates the USS Alabama Battle- ship Memorial Park. The curatorial staff maintains an active program of restoration and maintenance, and has developed interpretive exhibits throughout the vessel, with considerable sensitivity to the ship’s historic fabric. The ship is nearly unique among World War II vessels in never having undergone updating for use in subsequent wars.

USS Alabama is in good condition and is subject to a continual program of maintenance and restoration work that is consistent with the Secretary of the Interior’s standards (Morton et al. 1997). Other than enhancement through restoration of damage caused by Hurricane Katrina, as well as ongoing maintenance and repair work, the exterior has remained unaltered since decommissioning. The battleship is located in Mobile Bay at a waterside park in which it and the submarine USS Drum are interpreted. USS Alabama was damaged by Hurricane Katrina, which caused it to list 12° in its cofferdam. It has since been righted and restored and a new protective cofferdam has been created. At the present time, USS Alabama is in good condition, is well maintained by ownership that recognizes and appreciates it historical and architectural significance, and the overall threat level is minimal, except for the unpredictable nature of the region’s extreme weather events.

Figure 20-13. The USS Alabama. (Photo credit: David B. Schneider, March 2015.)

Cultural Landscape - Chapter 20 163 Figure 20-14. Galley of the USS Alabama. (Photo credit: David B. Schneider, March 2015.)

Figure 20-15. USS Drum at Battleship Alabama Memorial Park, Mobile, Alabama. (Photo credit: David B. Schneider, March 2015.)

164 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area USS Drum (1986 NHL Designation) USS Drum is a Gato-class submarine (designated SS-228 by the US Navy), commissioned in 1941, that represents the standard design for American fleet submarines early in World War II (Figures 20-3, 20-15, 20-16, 20-17). This vessel served throughout the war, sinking 15 enemy ships and earning 12 battle stars. USS Drum was decommissioned in 1946, served as a training vessel for the Navy Reserve, and was donated to the USS Alabama Battleship Commission in 1969 for display at USS Alabama Battleship Memorial Park in Mobile, Alabama. Initially moored in Mobile Bay, USS Drum was damaged by storm surge during Hurri- cane Georges in 1998 and subsequently moved on shore. The ship has suffered considerable rust damage to its below decks prior to being relocated to a fixed aboveground platform. Ongoing restoration is underway to repair the damage.

USS Drum is in good condition and is subject to a continual program of maintenance and restoration that is consistent with the Secretary of the Interior’s standards (Morton et al. 1997). USS Drum is well maintained, but still requires restoration work. All of the essential character-defining elements of the USS Drum remain intact, and historic fabric and materials are carefully respected during ongoing repair and restoration. Integrity of setting has been compromised to a degree by its removal from the water and Figure 20-16. View across the foredeck of USS Drum. placement on land. However, this action was neces- (Photo credit: David B. Schneider, March 2015.) sary for the long-term preservation and interpretation of the submarine. While the ownership’s maintenance program and emergency management plan minimize threats to the vessel, a major storm could pose a threat to the submarine given the unpredictable nature of the region’s extreme weather events.

Figure 20-17. After torpedo room of USS Drum. (Photo credit: David B. Schneider, March 2015.)

Cultural Landscape - Chapter 20 165 Summary tornadoes and each has previously suffered significant The 6 National Historic Landmark resources assessed damage from storms. While steps have been taken in this study retain integrity, are well maintained, and to minimize this risk at some of the individual land- are continuing established maintenance and resto- marks, there is nothing that can prevent damage in the ration programs. None of the landmarks faces any event of a major storm. Most of the owners of these immediate threat to their current status. The common landmarks have developed an emergency response threat to each of the landmarks is weather related, plan to respond to natural and other disasters. as all are located in a region prone to hurricanes and

166 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 21. A Socioeconomic Overview of the Greater Mobile-Tensaw River Area Ermanno Affuso, University of South Alabama

For the purposes of this socioeconomic overview, the greater Mobile-Tensaw River area is considered as the Alabama portion of the Mobile-Tensaw watershed that spans an area of approximately 7600 km2 (2900 mi2; Figure 21-1A). This region includes stakeholders from the counties of Baldwin, Clarke, Escambia, Mobile, Monroe, and Washington. The total population of the area is 704,106, with 37.6% identifying as African American, 60.1% Caucasian, and the remaining 2.3% Hispanic and other races (US Department of Commerce 2013). A graphical analysis of the age distribution of the population of this area is illustrated in Figure 21-1B.

The population pyramid reveals a near-stationary pattern where the age groups are similar in percent- age terms, except for the upper tail of the age distribution. This result contrasts with the constrictive pattern of the US national population pyramid that is similar to Western European countries (Cohen 2003; Pollack 2005). Median age of the population is 38 years, in line with the national trend and that of other developed countries (United Nations 2002). The dependency ratio for this region (calculated as the ratio of people Figure 21-1A. Geography of the study area. who are not potentially in the labor force to those typically in the labor force) is approximately 71%, which is slightly above the national dependency ratio of 67% estimated by Vincent and Velkoff (2010). (To be consistent with the national estimates of Vincent and Velkoff [2010], the dependency ratio in the current study is calculated as (# of people 0 to 19 + # of people 65 and above)/(# of people 20 to 64). The assumption that the segments of the population age 15 to 19 and 65 and above are not in the labor force is quite strong. However, the dependency ratio in this case should be considered as a combined ratio of youth and old-age group to the population aged 20 to 64 years.) Figure 21-1B. Demography of the study area.

Cultural Landscape - Chapter 21 167 Socioeconomic Analysis that residents who belong to the first quartile of the According to the US Census Bureau, the population income distribution ($3,062 to $17,002) are located increased by 8.9% between 2000 and 2013. In 2013, mainly in the northern part of the Mobile-Tensaw 1.7% of the population was composed of new resi- watershed and in the Mobile metropolitan area (Figure th dents who relocated to the Mobile-Tensaw region 21-2A). Residents belonging to the 4 quartile of the from other counties in Alabama, 2.2% of the popu- income distribution ($26,507 to $39,073) are located lation were new residents who relocated from other in the southern parts of the watershed in Mobile and states in the US, and 0.3% of the population consisted Baldwin counties. The portions of the region with of new foreign residents. In the same year, income the largest unemployment rate are those areas in the per capita was $21,503, 22.6% of the population lived northern part of the watershed (Figure 21-2B). These below the poverty line, and the unemployment rate spatial trends may predict the geographic economy was approximately 7.95%. The proportion of the of the greater Mobile-Tensaw River area with major population with less than a high school education economies located in the southern part of the water- was 17.3%, while 29.3% of residents were reported to shed and minor economies that occupy the northern hold an associate’s degree. part of the watershed. A disaggregated geospatial analysis of per capita Overview of Economic Activities income and unemployment rate of this region is The County Business Pattern dataset of the US reported in Figure 21-2. The analysis, based on data Census Bureau was used to identify major and minor at the census tract level available from the US Census economies of the study area, as well as the main Bureau’s American Fact Finder (US Cenus Bureau economic activities sorted by number of industrial [date unknown]a), shows a spatial dependence of establishments and annual employee payroll. Note these two socioeconomic factors. In fact, it appears that the County Business Pattern was built by the US

Figure 21-2. A disaggregated geospatial analysis of A) per capita income and B) unemployment rate for the region.

168 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Census Bureau upon the Business Register, which is each sector. Table 21-2 reports the annual payroll a comprehensive database of business establishments of each industry in the 6 counties of the study area. at county level. Business Register information is confi- Compared to the previous considerations based dential, according to Title 13 and 26 of the US Code, on number of establishments, the payroll analysis so data that would disclose the operation of an indi- provides more accurate information on the state of the vidual employer are not reported in the public data- economy in an attempt to identify the major industrial set. Further, the US Census Bureau adopts the noise areas that characterize the region. infusion method, which adds a bias (2 to 5% noise) to protect sensitive data prior to publication (US Cenus Descriptive statistics show that manufacturing is the Bureau [date unknown]b). In the case of the current leading industrial sector in the area with a payroll study, information on annual payroll of some of the volume of approximately $2.6 billion, represent- industrial sectors may be subject to noise infusion. ing 26.0% of the annual volume of employee payroll However, even if there is a small bias in part of the across the entire industry. Among the top 5 indus- information on the County Business Pattern reported trial sectors, healthcare and social assistance is the by the public agency, these data still provide a valid second largest economic sector that paid its employ- snapshot of the current state of economic activity in ees approximately $1.22 billion (12.2%), followed the greater Mobile-Tensaw River area. by the construction sector with $913 million (9.1%), retail and trade with $882 million (8.8%), and whole- Data on number of establishments and annual sale trade with $824 million (8.2%). When retail employee payroll of the industrial sectors in the coun- and wholesale are combined, the entire trade sector ties of Baldwin, Clarke, Escambia, Mobile, Monroe, amounts to approximately 17% of the value of total and Washington were extracted from the national wages received by workers in this region. The agri- dataset and reported in Tables 21-1 and 21-2. Accord- cultural, forestry, fishing, and hunting sector paid its ing to the most recent available data, in 2013 there were employees approximately $59 million, only 0.6% of 15,357 industrial establishments in the area of study. total salaries across the reported industries. As expected, retail trade is the sector with the largest number of enterprises (2840) representing 18.5% of The geographic distribution of annual payroll clearly total establishments. Interestingly, there were 1379 shows two outliers: Mobile and Baldwin counties establishments in the healthcare and social assistance with approximately $7.37 billion and $1.65 billion sector (9.0% share of total establishments) mainly total aggregate value of wages paid by the entire indus- located in the counties of Baldwin and Mobile (88% try of the area. These two counties represent 73.7% of total establishments within the sector). These data and 16.5% of the total annual payroll volume of the may be consistent with the demographic structure industry, respectively. Consequently, these two coun- of the population, with a 22.6% old age dependency ties can certainly be considered the major economies ratio. Furthermore, Serow (2001) finds scientific of the greater Mobile-Tensaw River area. Figure 21-3 evidence of a retirement migration towards coastal illustrates a graphic summary of the geographic distri- locations of the southeastern United States that may bution of annual payroll across the various segments potentially shift the demand for healthcare services. of the industry of this region. The number of establishments in the accommodation Major Economies of the Greater and rood services sector was 1296 (8.4%), while in the Mobile-Tensaw River Area agriculture, forestry, fishing, and hunting sector there were 179 enterprises representing less than 1% of the 1) Mobile Metropolitan Area total number of firms in this area. By looking at the Metropolitan Mobile is located on the western shore spatial distribution of the industrial sectors, it appears of Mobile Bay over an area of 562 km2 (217 mi2) with a that 55.8% of the enterprises are located in Mobile population of 413,188 inhabitants (US Census Bureau County, followed by Baldwin County hosting 31.7% [date unknown]a). As illustrated in Figure 21-4, this of total establishments, while Washington County has metropolitan area is the crossroads of two main high- only 217 firms representing 1.4% of the total share. ways: Interstate 10 (a transcontinental highway that connects Santa Monica CA to Jacksonville FL) and Annual salaries received by employees can be consid- Interstate 65 (connecting Mobile AL to Gary IN). In ered a proxy of the value of the industrial output per addition, the presence of an international port and a

Cultural Landscape - Chapter 21 169 NA 1.52% 3.35% 3.70% 4.70% 4.76% 5.67% 7.53% 8.09% 8.44% 0.08% 8.98% 0.24% 9.08% 0.51% 0.72% 0.97% 1.00% 1.25% 10.93% 18.49% 100.00% Sector Share 12 37 78 233 515 568 722 731 871 110 149 154 192 Total 1,156 1,242 1,296 1,379 1,394 1,678 2,840 15,357 100.00% 2 1 0 2 8 1 1 9 1 0 13 14 12 19 10 19 14 35 31 25 217 1.41% Washington 2 9 6 0 2 7 3 3 18 19 12 22 34 12 30 32 16 47 93 26 393 2.56% Monroe 9 95 18 36 79 41 130 313 323 398 452 580 646 638 681 758 843 975 107 8,567 1,445 Mobile 55.79% 7 0 6 8 5 1 19 23 35 18 18 23 65 62 52 61 37 94 19 723 170 4.71% Escambia 6 0 2 7 1 4 11 30 26 12 14 14 64 24 48 53 30 76 27 586 137 3.82% Clarke 2 8 80 63 11 21 11 39 118 151 282 239 224 335 487 475 456 454 451 964 4,871 31.72% Baldwin Arts, entertainment, and recreation Information Transportation and warehousing Transportation Manufacturing Real estate and rental and leasing Real estate and rental Administrative and support and waste Administrative and support services management and remediation Wholesale trade Finance and insurance County Share County Share Construction Total Accommodation and food services Accommodation and food Industries not classified Health care and social assistance Health care Mining, quarrying, and oil and gas extraction Professional, scientific, and technical services scientific, and technical Professional, Utilities Other services (except public administration) Other services Management of companies and enterprises Retail trade Agriculture, forestry, fishing and hunting forestry, Agriculture, Industrial Sector Educational services Business pattern in the greater Mobile-Tensaw River area: number of establishments in 2013 (US Cenus Bureau [date unknown]a). in 2013 (US Cenus of establishments River area: number Mobile-Tensaw in the greater 21-1. Business pattern Table

170 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area NA 1.89% 1.97% 2.66% 3.83% 3.95% 5.01% 5.13% 0.00% 6.09% 0.36% 8.24% 0.41% 8.81% 0.54% 9.13% 0.59% 1.38% 1.83% 12.22% 25.97% 100.00% Sector Share 93 Total 35,656 41,523 54,374 58,617 188,845 196,750 266,298 383,112 395,274 501,637 513,265 609,101 824,300 882,244 913,514 138,525 183,504 100.00% 1,223,008 2,599,800 10,009,440 0 0 0 0 0 0 0 0 0 609 1,467 5,561 2,088 2,442 1,853 5,896 1,998 5,772 8,410 3.50% 349,860 313,764 Washington 0 0 0 0 0 0 984 2,969 3,857 4,774 4,315 7,363 1,644 1,170 4,801 1.19% 26,312 10,233 17,098 10,401 23,008 118,929 Monroe 0 93 24,800 45,324 23,660 78,882 Mobile 73.66% 147,492 179,626 177,751 181,835 175,429 306,549 373,971 431,993 509,302 691,666 482,402 774,138 846,330 7,373,141 1,921,898 0 0 0 6,946 5,362 2,240 7,303 3,743 6,304 9,050 8,065 1,380 2.96% 10,800 17,765 16,245 13,614 34,543 12,547 40,787 99,878 296,572 Escambia 0 0 0 0 0 163 1,923 5,926 6,942 6,386 4,286 2,101 6,483 3,196 1,107 2.20% Clarke 15,170 32,189 10,063 29,085 95,018 220,038 0 0 0 656 24,174 16,759 64,993 32,701 86,800 71,599 87,782 35,656 16,560 56,172 16.49% 185,017 100,451 310,116 116,834 275,388 169,242 Baldwin 1,650,900 Information Utilities Management of companies and enterprises Other services (except public administration) Accommodation and food services Transportation and warehousing Transportation County Share Finance and insurance Total Administrative and support and waste Administrative and support services management and remediation Industries not classified Professional, scientific, and technical services scientific, and technical Professional, Educational services Wholesale trade Arts, entertainment, and recreation Retail trade Mining, quarrying, and oil and gas extraction Construction Agriculture, forestry, fishing and hunting forestry, Agriculture, Health care and social assistance Health care Real estate and rental and leasing Real estate and rental Manufacturing Industrial Sector 2013 Annual payroll (expressed in thousands of dollars) of industrial sectors in the greater Mobile-Tensaw River area, shown by county (US Cenus Bureau by county (US River area, shown Mobile-Tensaw sectors in the greater dollars) of industrial in thousands of payroll (expressed 21-2. 2013 Annual Table [date unknown]a).

Cultural Landscape - Chapter 21 171 Figure 21-3. Geographic distribution of industrial sectors.

regional airport makes Mobile a hub for distribution and transportation logistics. In fact, the Alabama State Port Authority reported an overall annual tonnage of 29 million tons (26 billion kg) for the fiscal year 2014. Primary cargos (outbound/inbound) include coal, metals, forest products, containers, frozen poultry, cement, chemicals, and project cargo. With over $500 million of direct and indirect tax impact, the total economic value of the port of Mobile is approximately $18.7 billion (Alabama State Port Authority 2015).

Data on Gross Domestic Product (GDP) for the Mobile metropolitan area from 2001 to 2013, available from the US Bureau of Economic Analysis, were used to calculate the GDP deflator for the study area as 100% times the ratio of the nominal GDP to the real GDP (2009 = 100). The GDP deflator was then used to calculate the inflation rate in the Mobile metropolitan area. Figure 21-5A shows nominal and real GDP expressed in 2009 constant dollars. A qualitative assessment, based on the diagram of the real GDP, shows a positive trend of economic growth from 2001 to 2013, most readily evident Figure 21-5B. Note that the economic growth rate peaked in 2006, the year Figure 21-4. Metropolitan area in Mobile-Tensaw watershed. after Hurricane Katrina, with approximately a 5.9% increase in real growth compared to the previous year.

172 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Figure 21-5. Macroeconomic indicators of Mobile metropolitan area: A) nominal and real GDP expressed in 2009 constant dollars; B) inflation rate and economic growth rate from 2002 to 2013.

These data are consistent with Mobile having been been in a range between 1.3 and 3.9 %, with a defla- moderately damaged by Hurricane Katrina in 2005. tionary 1.1% in 2009. The area may have attracted residents from neigh- boring counties who suffered major injuries from the 2) Comparison of the Mobile Metropolitan storm. Garber et al. (2006) found that employment in Area with Daphne-Fairhope-Foley the Mobile area had grown by 3.1% in August 2005, The micropolitan area (defined as an urban area with when the hurricane struck, but Katrina did not stop a population greater than 10,000 but less than 50,000) th the employment growth. In fact, in the 4 quarter of of Daphne-Fairhope-Foley is the second largest econ- 2005, Mobile experienced storm-related job growth. omy of the greater Mobile-Tensaw River area. This Economic growth slowed down in 2007 and increased area occupies 125 km2 (48 mi2) of the Eastern Shore again in 2013, when the economic growth rate of the of Mobile Bay, with a total population of 187,114 resi- metropolitan area was 2.9%. The inflation rate has dents in 2013 (US Census Bureau [date unknown]a).

Cultural Landscape - Chapter 21 173 A comparison of the industrial sectors of the Daphne- than their corresponding population ratio (which is Fairhope-Foley micropolitan area and the Mobile 413,188/187,114 ≈ 221%). Pie charts illustrate a disag- metropolitan area is based on business pattern survey gregated comparison by the top 8 industrial sectors data of the US Census Bureau for 2013. As in the previ- (Figure 21-6). ous analysis, total annual payroll per industrial sector is used as a proxy for the value of industrial produc- Industrial sectors such as wholesale trade, adminis- tion. Total value of wages received by the workers trative and support, waste management, transporta- across all industries in the Mobile metropolitan area tion and warehousing, management of companies was approximately $5.53 billion versus $1.50 billion and enterprises, real estate, and others have been received by the employees in the Daphne-Fairhope- aggregated in one class called “other services” that Foley area. Volume of the payroll in the Mobile metro- represents 30% of the total annual industry payroll politan area is 368% of Daphne-Fairhope-Foley, more in the Mobile metropolitan area versus 26% of the

Figure 21-6. Comparison of the industry of A) Mobile metropolitan area, and B) Daphne-Fairhope- Foley micropolitan area.

174 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Daphne-Fairhope-Foley area. Healthcare and social comparison of the accommodation and food services assistance appears to be the largest industrial sector in sector in these two areas is noteworthy. In the Mobile the Mobile metropolitan area and the second largest metropolitan area this sector accounts only for 3% sector in Daphne-Fairhope-Foley, representing 17% of total annual payroll across all industries ($174 of total annual payroll in both areas. Manufacturing is million), while in the Daphne-Fairhope-Foley area the second largest sector in Mobile with $923 million this is the third largest sector with 11% share among in total payroll (17% sector share) versus $145 million all industries and $165 million in annual payroll. This in the Daphne-Fairhope-Foley (10%, third largest data may reflect the geographic proximity of this area sector). The manufacturing sector of Mobile includes to the coast, which potentially attracts more tourists aviation and aerospace, chemical, maritime, steel, and compared to the metropolitan area of Mobile. others (Mobile Area Chamber of Commerce 2015). Minor Economies of the Greater The largest industrial sector, ranked by total annual Mobile-Tensaw River Area payroll, in the Daphne-Fairhope-Foley area is retail and trade, with $290 million annual payroll (19% Marginal economies are spread over the rural terri- sector share). The same sector is ranked 4th in the tory of the greater Mobile-Tensaw River area, outside Mobile metropolitan area, with a total annual payroll the main urban areas already analyzed. According of $474 million (8% sector share), after the finan- to the USDA, in 2014 Baldwin, Escambia, Monroe, cial sector. The two areas have different industrial and Mobile counties topped the state production patterns. While the economy of Mobile is primar- of peanuts with 95.3, 72.6, 33.1, and 26.7 million ily dominated by manufacturing industry, Daphne- lb (43, 33, 15, and 12 million kg), respectively. The Fairhope-Foley is mostly a residential area, which may same year, Escambia and Monroe counties were the increase demand for retail and trade services. In addi- largest producers of cotton in Alabama with 40,300 tion, given the proximity of the two areas connected and 31,200 bales, respectively (USDA-NASS [date by several routes, including Interstate 10, a consistent unknown]). number of workers reside in Daphne-Fairhope-Foley Total value of agricultural products of the greater and commute daily to Mobile for work. Mobile-Tensaw River area is available from the The geographic spillover of demand for housing in Census of Agriculture, a survey conducted every 5 these two areas, which may include potential differ- years by the National Agricultural Statistical Service ences in neighborhood quality, could be a topic of (NASS) of US Department of Agriculture. These academic research by urban economists. However, data have been deflated using the Consumer Price

Figure 21-7. Total value of agricultural production in constant 2012 dollars.

Cultural Landscape - Chapter 21 175 Index (2012 = 100), South Region, released by the US Summary Census Bureau (Figure 21-7). According to the survey, The major conclusion derived from this short total value of agricultural output of the area increased descriptive analysis of socioeconomic indicators by 9.7% from 2002 to 2007 and by 27.9% from 2007 of the greater Mobile-Tensaw River area is that a to 2012. The commercial value of agricultural output spatial trend exists in the economic development of was approximately $267 million in 2012. At a qualita- the area. In this region it is possible to distinguish a tive level, the chart shows that agricultural production predominantly rural economy in the northern part in Mobile County declined slightly in the last decade of the Mobile-Tensaw watershed and a major emerg- of the survey, while agricultural activities of Bald- ing economy in the southern part, towards the coast. win, Escambia, Monroe, and Washington counties This spatial trend is not new. It has persisted since the increased. For example, Monroe County experienced boom of the cotton trade in the 19th century, when the a sharp decline in the value of its agricultural output economies of riverside commercial cities such as New of approximately 80% from 1997 to 2002. However, Orleans, Natchez, and Mobile flourished downstream after 2002 the total value of the agricultural product of agricultural areas (O’Sullivan 2007, p 22). in the county increased from $5.2 million in 2002 to $10.4 million in 2007 (99.6% real growth) to reach $32 million of 2012 (209% real growth). Monroe County had the best relative performance in terms of increased value of agricultural product in the entire area.

176 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area 22. Conservation and Tourism Development in the Greater Mobile-Tensaw River Area Michele L. Archie, The Harbinger Consulting Group

In 2013 and 2014, my firm consulted with an infor- (Archie and Mullins 2013) with more than 3 dozen mal coalition of organizations on potential economic business leaders, local government officials, land impacts of conservation designations in the greater managers, scientists, conservation leaders, and tour- Mobile-Tensaw River area, and on compatible tourism promoters from 7 southwest Alabama counties ism development. A public dialogue was taking (Baldwin, Clarke, Escambia, Mobile, Monroe, Wash- shape—perhaps re-emerging—in the greater Mobile- ington, and Wilcox). These interviews, and obser- Tensaw River area about the essential character and vation of emerging forums for regional collabora- significance of the place and the feasibility of coor- tion such as the Delta Roundtable, suggest that area dinated, landscape-level conservation to protect residents and communities may be on a path toward it. This emerging public dialogue is reminiscent of developing—or at least considering—a regional iden- conversations that have occurred in other places tity that also makes ecological sense. coming to grips with efforts to align natural resource management and socioeconomic development with This research confirms that a tremendous personal maintaining ecological integrity and health. Paral- affection for the Delta is shared by many across the leling Rick Reese’s (1985) popular encapsulation of region. The interviews also revealed a common view the policy debate around the Greater Yellowstone that the Delta and surrounding uplands and commu- Ecosystem in the mid-1980s (Clark and Zaunbrecher nities are a largely untapped resource for economic 1987), several key themes are emerging in the greater growth that complements the existing economy and is Mobile-Tensaw River area: compatible with maintaining both the area’s ecological health and its rural character. 1. The Delta is a very special, and in some ways unique, place. The evolution of a regional identity that corresponds with a functional natural system may be motivated 2. The Delta is not an isolated feature; rather, it in part by these common perceptions, as well as by exists in a larger ecological context that includes shared desires, such as increasing public recreational the surrounding uplands and an extensive river access to the scenic environments of the Mobile- drainage. Tensaw Delta and surrounding uplands, providing 3. This ecosystem is an extraordinary national trea- better protection for culturally significant features and sure, encompassing the nation’s richest aquatic diverse habitats, and ensuring the ecological health of system and some of the world’s richest temperate already conserved areas. forests. Local organizations are exploring a variety of ways 4. Most resource management decisions in the to expand conservation and recreational access in greater Mobile-Tensaw River ecosystem are the greater Mobile-Tensaw River area. Communi- made in a fragmented manner that does not rec- ties across the region are developing and promoting ognize the area as an ecological unit. nature, heritage, and cultural tourism in and near the 5. The environmental integrity of the greater Delta. The natural areas at the core of this ecosystem Mobile-Tensaw River ecosystem is imperiled are seen by many as assets that could be leveraged to by activities and developments, and the task attract more tourists, greater investment, and new of conserving and maintaining its integrity is residents and businesses. incomplete. In this chapter I draw on my research in the greater Mobile-Tensaw River area, particularly on interviews

Cultural Landscape - Chapter 22 177 Tourism is Underdeveloped in the Internet search covering the entire 7-county region Greater Mobile-Tensaw River Area yielded just 6 additional service providers. According to census data, between 2001 and 2010, tourism made a steady contribution to the region’s Factors Contributing to economy. In each of the 4 counties for which figures Underdevelopment of Tourism in the were reported, two sectors closely related to tourism Greater Mobile-Tensaw River Area maintained a relatively consistent share of the overall My research suggests 4 important factors, described economy, as measured through the proxy of private below, that contribute to underdevelopment of tour- employment (US Department of Commerce 2001). In ism in the greater Mobile-Tensaw River area. These 2013, as in years past, Baldwin County had the stron- factors focus on nature, heritage, and cultural tourism, gest concentration of tourism jobs, with a total of 30% consistent with sustaining the character and ecologi- of private employment in accommodation and food cal integrity of the area’s landscapes and waterways. services and in arts, entertainment, and recreation (Table 22-1; US Department of Commerce 2013). 1) Key Elements of Visitor and Transportation Infrastructure Are Missing or Not Well These data suggest that the north-south spatial trend Developed in the area’s economic development identified by High-quality, year-round lodging is in short supply, Affuso (this volume) is also reflected in economic especially in the upper reaches of the Delta and adja- activity related to tourism. The concentration of cent uplands. Facilities are not large enough to accom- tourism-related jobs in Mobile and Baldwin counties modate group tours, and there are few lodging options reflects a bias in visitation toward the more developed that appeal to nature and heritage travelers looking for coastal and urban areas to the south. The northern, authentic, local experiences. more rural portions of this region that encompass the Delta and bluffs see significantly less economic activ- Public and shared private transportation networks are ity from tourism. lacking. Without a private vehicle, it is difficult to reach destinations in and around the Delta from Mobile and The relative underdevelopment of tourism in these the coastal areas that attract the bulk of the region’s more rural and inland parts of the greater Mobile- visitation. A recent tourism strategic plan for Mobile Tensaw River area is further illustrated by business (Dow 2012) identified high-speed ferry connections activity related to wildlife tourism. A recent study as a key to tying together attractions, small towns, of wildlife tourism in coastal areas along the Gulf of beaches, ecotours, and the downtown Mobile Mari- Mexico (Stokes and Lowe 2013) identified 89 compa- time Center. The plan also suggests developing other nies that provide wildlife watching, fishing, and land- and water-based transportation services, includ- hunting guide services in Baldwin and Mobile coun- ing water taxis, expanded bus tours, and easy-access ties. Only 4 of these offered services in the Delta. An rental cars to connect tourists to places they want to visit or to Table 22-1. Concentration of jobs in tourism-related industries, 2013 (US Department key hubs for visitor activity. of Commerce 2013). Expanding regional transporta- Accommodation and Arts, entertainment, Total private-sector tion links beyond Mobile and food services share of and recreation, share of jobs in two tourism- Baldwin counties—or even to County private employment (%) private employment (%) related industries northern Baldwin County, a Baldwin 20.8 2.0 12,426 crucial point for accessing the Clarke 9.1 0.1 615 Delta—may not be viable with- Escambia 9.9 7.4 1,730 out significant added attractions Mobile 8.9 1.3 15,076 to draw more local and out-of- Monroe 7.4 0.4 391 area visitors into the northern Washington 1.1 0.1 67 part of this region. The north- Wilcox 8.7 0.2 154 ernmost terminal considered in a decade-old feasibility study Total reported jobs in two industries, 2013 30,459 for a Mobile Bay ferry was in

178 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Daphne, in Baldwin County south of the Delta (Day 4) Regional Collaboration on Tourism is Just Wilburn Associates 2004). To connect visitors with Beginning and is Not Well Developed Outside destinations further north, ferry service would need Baldwin and Mobile Counties to connect public and private transportation provid- Most efforts to promote nature, heritage, and cultural ers serving these areas. Such services currently do not tourism have focused on specific communities or exist, except for small-scale operations focused on counties, and most attempts to collaborate across commuters and persons with special needs (Archie community or county lines have been limited in 2014a). scope, aimed at establishing small-scale partnerships and specific efforts. A more ambitious collaborative 2) Public Recreational Access is Limited effort began in 2014, when representatives of several Despite a roster of more than 40 public and private tourism promotion organizations in Mobile and Bald- fishing access sites and launches, the Delta is accessi- win counties formed the Regional Tourism Council to ble mostly by private boat. Few companies rent canoes advise the Coastal Alabama Partnership on policies and kayaks or offer guided tours (Archie 2014b). Fuel needed to develop and promote tourism regionally. In availability limits the range of powerboats traveling May 2015, the council launched a collaborative “Visit through or deep into the Delta. The number, location, Coastal Alabama” website. and quality of public land and water access are seen as impediments to expanding guided and outfitted expe- Mobile and Baldwin counties anchor the region’s riences for visitors (Archie 2014c). population and developed visitor services and attractions. But focusing on those two counties alone leaves Interviews identified other limitations on public recre- out a) a significant portion of the geography of the ational access: few developed camping opportunities area, and b) a good share of the opportunity to include in the upper reaches of the Delta and surrounding the upstream portions of the Delta and surrounding bluffs; a lack of long-distance or connected networks uplands and communities as part of a common visi- of hiking, biking, and horseback trails; and underuti- tor destination. The ground seems fertile for devel- lized or closed recreational sites managed by agencies oping active partnerships to develop and promote without a primary focus on natural area conserva- nature and cultural tourism in the Delta region. Many tion and recreation, such as the US Army Corps of key players are acknowledging the potential benefits, Engineers. and even the necessity, of regional collaboration to leverage and protect the area’s natural and historical 3) The Area and Its Existing Nature, Heritage, resources, and to attract more visitors to rural areas and Cultural Tourism Opportunities Are (Archie 2013). Fragmented, Not Well Promoted, and the Area is Not Easily Identifiable as a Distinct Opportunities to Align Conservation Destination and Tourism/Economic Development The Delta itself is not well known outside the region, Local organizations—including a nascent regional nor is there a flagship attraction within the Delta and leadership forum known as the Delta Roundtable— surrounding area to draw significant numbers of have been exploring a variety of ways to expand and nature, heritage, and cultural visitors. coordinate conservation and recreational access in the greater Mobile-Tensaw River area, including large- Tourism promoters in the area have recognized this scale conservation agreements with private landown- kind of tourism as an important avenue for increasing ers and enhancing and expanding state conserva- economic activity in the area. This focus is seen as a tion lands. National support, including broadening way to complement an already strong, beach-focused the role of the National Park Service in the region, is tourist trade, and to help draw visitors to less known also being explored to complement and extend these and less traveled parts of the region. Lack of fund- efforts. ing for marketing and promotion of facilities, sites, and services offered in the Delta is a major hurdle in As these explorations mature, connections between increasing visibility and visitation, especially for more conservation and leveraging the region’s rich natural rural and remote communities and sites. and heritage resources for shared economic benefit are emerging.

Cultural Landscape - Chapter 22 179 The majority of conserved land in the Delta and surrounding uplands is in the floodplain. Among the highest conservation priorities in the region are natural areas adjacent to the floodplain, contiguous blocks of longleaf pine and related habitats, and unique ecotypes away from the floodplain. Depending upon the management regime adopted, adding these priority landscapes to the region’s conserved-lands portfolio could help boost the region’s appeal to visitors by increasing land-based recreation opportunities, which are popular among visitors, even in areas Figure 22-1. Most popular recreational activities based on visitor studies in 4 water- that are seen as largely water- and wetlands-focused National Park Service areas: New River Gorge National River oriented (Figure 22-1). in 2004; Congaree National Park in 2005; Big Cypress National Preserve in 2007; and Everglades National Park in 2008 (Papadogiannaki et al. 2008; NPS [date unknown]). Another way in which conservation and tourism development could reinforce each other in the greater Mobile- Table 22-2. Everglades National Park, visitor group participation in selected guided and Tensaw River area is through self-guided activities for winter and spring 2008 (Papadogiannaki et al. 2008; NPS [date unknown]). expanding guide and outfitting services. Visitor research Guided # of visitor groups Self-guided # of visitor at other destinations may participating on day of groups participating on day provide a window into types Activity survey of survey of experiences for which Boat tour 428 47 travelers prefer to use guides Tram tour 256 or outfitters. For example, Walking/hiking 65 683 visitor surveys in the Florida Photography/painting 30 531 Everglades—a landscape that, Boating 30 57 like the Delta, is dominated by Canoeing/kayaking 30 85 water and wetlands—suggest Birdwatching 12 114 that some types of visitor Bicycling 12 114 activities lend themselves to Camping 6 85 guided experiences better than others (Table 22-2). their clients minimize the impact and risks associated A study of Montana’s outfitting and guiding industry with recreation. In the Montana study, visitors rated (Nickerson et al. 2007) identified several advantages their connection to nature as the most valued part of of promoting guided recreation. Outfitters both help their outfitted trip. attract visitors to Montana and benefit from existing visitation, and visitors who participate in an outfit- Further, guides and outfitters have a business inter- ted trip tend to spend more money than those who est in—and most often a personal commitment to— do not. Visitors learn about the area’s natural and conserving the areas and resources on which their cultural history from their guides. Guides can help

180 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area profits depend (Nickerson et al. 2007). Mark Grisham points, 3 land-based camping areas, and 4 floating (personal communication, unreferenced, 2013), a campsites for paddlers only. A southward expansion longtime member of the Grand Canyon River Outfit- is underway to extend the trail to the 5 Rivers Delta ters Association, points out that guides and outfitters Resource Center. can play an important leadership role with other local businesses, influencing the path of tourism develop- The Bartram Canoe Trail was started in the mid-2000s ment and conservation. Grisham suggests that devel- by the Alabama State Lands Division and the Alabama oping a business alliance can help strengthen guides, Department of Conservation and Natural Resources, outfitters, and other businesses, enable collabora- mostly on land secured by Alabama Forever Wild. tion, and expand the influence of member businesses Between 2004 and 2012, ADCNR recorded 762 reser- in working with government partners, whether land vations for the 4 floating campsites designated for use management agencies, permitting agencies, or tour- by paddlers only. Beyond that, trail use is not tracked. ism promotion boards. Because it is a young and relatively unknown water trail, Bartram may have a long way to go before reach- Interviews in the greater Mobile-Tensaw River area ing the “full use” levels of more mature trails. confirm that this kind of leadership and collaboration is seen as necessary for sustaining public support for Comparing the Bartram Canoe Trail with other trails conservation. Alabama’s Forever Wild conservation provides a sense of its potential as a draw for paddlers. land trust program was created in 1992 by a constitu- Averages derived from studies of 6 paddling destina- tional amendment that passed with 83% of the popu- tions that include 75 or more waterway miles (120 lar vote (League of Women Voters 2012). The program km) provide data—including average expenditure was reauthorized in 2012 with 75% voter support per trip—needed to estimate both likely potential use (Ballotpedia.org [date unknown]). Recently, Forever and economic impact for the Bartram Canoe Trail in Wild has withheld support for acquisition of high its current configuration. At 200 miles (322 km) of quality lands surrounding the delta that could have waterway and an average of 199 paddlers/mi/yr (124 addressed issues of large landscape scale connectivity paddler/km/yr), the Bartram trail would host 39,800 and ecological processes; and it is expected that such annual visitors. At an average visitor expenditure of acquisitions would likely require state-federal part- $67 per trip, the total visitor spending would be $2.67 nerships, e.g. Land and Water Conservation Fund. million. Using a multiplier to estimate the secondary and indirect impacts of this spending, the annual Raising local awareness of, and appreciation and impact of these paddlers would be $4.4 million in support for, protecting the area’s natural, historical, sales at local businesses, and 58 jobs (a rough calcu- and cultural resources is also important because word lation that does not attempt to differentiate between of mouth is one of the primary drivers of visitation to local and non-local visitation). regional tourism trails, byways, and other natural and historical attractions. A study of visitors to Minne- Managing for Shared Benefits sota’s Paul Bunyan Scenic Byway, for example, found As the local exploration of ways to expand and coordi- that over 25% of byway users discovered it through nate conservation, recreational access, and economic word of mouth, about 20% through signage, and 15% development in the greater Mobile-Tensaw River area on maps (Liechty et al. 2010). Along the Crooked continues, it will be important to continue to identify Road, a music heritage trail spanning 10 counties in and refine the region’s shared goals for these activi- southern Virginia, almost half of study participants ties. These goals can help guide strategic decisions discovered the Crooked Road venue they were visit- and collaborative action, for example deciding which ing on the recommendation of friends or coworkers public and private conservation and land manage- (Jones 2008). ment agencies and approaches are the best fit. Potential Economic Benefits of the Conservation acquisitions and designations are not Bartram Canoe Trail guaranteed, by themselves, to improve recreational The state-run Bartram Canoe Trail in the Mobile- access, to boost visitation, nor even to maintain or Tensaw River Delta includes roughly 200 mi (320 km) improve ecological integrity. How these areas are of day-use and 1-to-2-night routes, several access managed matters, as does what happens outside their boundaries. A review of World Heritage Sites

Cultural Landscape - Chapter 22 181 (Rebanks Consulting and Trends Business Research as being able to work in complementary ways with 2009) recently concluded that even with this interstate and local agencies that focus more on enforce- nationally prestigious status, increased visitation and ment. That juxtaposition raises an important point economic gain are not givens. Without additional for local consideration. Different public agencies and efforts, sites and their host communities may see a 0 to private organizations involved in conservation and 3% increase in visitation. But for a small and growing land management have different priorities, mandates, number of World Heritage Sites, most designated rela- expertise, assets, and rules for budgeting and allocat- tively recently, the status has been a powerful catalyst ing their resources. These differing priorities can result for socioeconomic change. The study attributes these in marked differences in natural area management differences to specific interventions based on shared and visitor services that significantly affect conserva- intentions to use the designation to catalyze change. tion and economic activity from tourism. Examples of Among the study’s conclusions: “World Heritage Site some of these points of comparison and their likely status is what you make of it.” impacts on conservation and tourism goals can be found in 2 studies comparing the US Forest Service A preliminary evaluation by Harbinger Consulting and the National Park Service as potential managing Group (2015a, 2015b) examined how a National Park agencies for specific public lands (Kathy and Steve Service unit, such as a National Park and Preserve, Berman Environmental Law Clinic 2008; Harbinger might help develop the economic potential of the Consulting Group 2011). patchwork of natural, cultural, and historical sites in the Mobile-Tensaw Delta and surrounding region. Inhabiting the Ecosystem This preliminary study found that economic impacts could vary widely depending upon many factors, In the 1990s, land managers in Alberta, British Colum- such as how the park and preserve is configured; the bia, and Montana drew the first maps of what would types of visitor facilities and services developed; the become known as the Crown of the Continent, an extent to which surrounding communities support, ecological region 3 times the size of Connecticut that promote, and enhance visitors’ park experience; and encompasses Waterton-Glacier International Peace how well discreet sites and events are coordinated Park, forested mountain ranges, river valleys, and and promoted to create more cohesive offerings for windswept prairies. At a public meeting in those early visitors. days, a man stood up and expressed a sentiment not uncommon in rural western communities beset by The study found that by its 10th year a National Park dramatic changes in natural resource management and Preserve, complemented by additional natural and the global economy: “We don’t want to be part and cultural tourism development in the Delta region, of your stinkin’ ecosystem” (personal communication could support up to 1074 new jobs in the 7-county with S. Thompson, unreferenced, 2014). region. Those jobs would be primarily in areas away For a time, land managers carefully drew “their” from the beaches, where most current tourism activity ecosystem boundaries to exclude human commu- is focused. Further benefits are anticipated. For exam- nities. But in areas like the Crown of the Continent, ple, a park-centered alternative transportation system Greater Yellowstone, and the greater Mobile-Tensaw could help connect the Delta region with established River area, it is difficult to avoid the conclusion that population and tourism centers around Mobile Bay human communities do not really have the choice to and the Gulf Coast. Park visitors could also represent opt in or opt out of the ecological systems they inhabit. significant business for a proposed Mobile Bay ferry Yet interestingly, the re-emerging conversation about system and other regional transportation services. protecting the essential character and ecological integ- In an interview one local business owner stated that rity of the greater Mobile-Tensaw River area suggests he believes an expanded National Park Service role that evolving a regional identity that makes ecological in the area could provide recreational access and sense might also make economic sense. visitor interaction that would help more people see, enjoy, and come to know the Delta. He sees NPS

182 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area SYNTHESIS

Sea encroachment, Mobile Bay, Alabama. Photograph courtesy of Hunter Nichols.

Synthesis - Chapter 23 183

23. A Greater Mobile-Tensaw River Area— Connections and Consequences Glenn E. Plumb, National Park Service; Gregory A. Waselkov, University of South Alabama; C. Fred Andrus, University of Alabama; Bill Finch, Earthword Services

Alabama citizens have consistently recognized and to those viewing the region from the air. As described praised the qualities of their own Mobile-Tensaw Delta by Ward et al. (this volume), the Delta’s water level as an area of local, regional, and national significance. can change from minute to minute, driven variously Ben Raines has characterized the Delta as it has been by tides, precipitation, wind, and drought. Bars and and is still now for many, part playground and part channels reposition over days or weeks in response natural bounty, a landscape of wondrous complex- to shifting currents. Changes occurring at the scale of ity and diversity that has been home to humans and decades or centuries can be seen in floodplain oxbow supported human endeavors for millennia (Raines, lakes, created from cut-off sections of river, and in this volume)—a landscape of such dynamic complex- winding swaths of deep green vegetation growing in ity that, if one does not deliberately seek to under- rich soils of infilled former channels. stand, can be quite forbidding, if not downright dangerous. The people who live in and near the Delta Finch and Plumb (this volume) describe how the know and respect this place. Mobile-Tensaw Delta and particularly the lower Alabama River and its tributaries slice through unusu- Yet there is so much more to learn about the Delta. ally complex and highly exposed Cenozoic and Plio- How do the physical properties of the area’s geology cene-Pleistocene strata, resulting in steep riparian and hydrology underpin diverse and dynamic biotic bluffs, coves, and the Red Hills erosional cuestas that systems, and how do they together shape the ecologi- in places drop several hundred feet to the river bed. cal setting for human relations with the Delta’s lands, Such features, highly unusual in the Southern Coastal waters, flora, and fauna? Plain, are believed to have been shaped by the interaction of uplift of ancient seabeds and large- and small- A combination of personal experience, natural and scale fluvial processes. The cause of the uplift remains cultural history, and science helps us understand that a matter of speculation, but active tectonics appears dynamic ecological and cultural connections bind responsible for the highest rate of elevational change together the Mobile-Tensaw Delta. This report’s (approximately 4 mm/yr [0.2 inches/yr]) of any place contributors have explored how this landscape’s in the Southeastern coastal plain. This vertical relief connectivity is organized and pointed out some conse- and the complex mineralogy associated with it have quences if our collective stewardship fails to preserve had profound effects on the ecosystems surrounding the Delta we have inherited. the Delta and also on the development of the rivers that wind through these steep headlands. As with Lay of the Land other rare areas of high relief or unusual soil charac- This landscape has been described as resembling a teristics within the coastal plain, such as the Apala- giant dowsing rod pointing south to the ocean, where chicola bluffs region, this combination of factors has the Tombigbee and Alabama river valleys merge into resulted in high total species diversity and unusual the extensive Mobile River valley, which forms a floral and faunal endemism (Duncan, this volume). “bayhead” delta at the head of Mobile Bay (Rodriguez, this volume). Deltas form at places where rivers meet Looking beneath the modern land surface reveals oceans. The dynamic waters of ocean coasts and rivers a deeper history to those who understand the never cease to sculpt the land they contact. The way language of sedimentology and geology. Rodriquez river currents shape the land is evident to any visitor to (this volume) helps us appreciate how the location the Mobile-Tensaw Delta today, and is especially clear and morphology of the valley is controlled, in part,

Synthesis - Chapter 23 185 by ancient tectonic events related to the creation the waters of the Mobile-Tensaw River system move of the Gulf of Mexico. More recently, over the past through 7 major channels. South of the confluence of few hundred thousand years, the valley experienced the Alabama and Tombigbee rivers, the Mobile River multiple episodes of erosion and deposition related to splits to form the Tensaw River, which runs along the climate cycles and associated fluctuations in sea level eastern side of the Delta, and the Mobile River, which that repeatedly exposed and inundated the continen- runs along the western side. The Mobile River Basin, tal shelf of the northern Gulf of Mexico. the 6th largest in the US and the largest (44,000 mi2 [114,000 km2]) east of the Mississippi River that drains Rodriguez (this volume) interprets this record of deep into the Gulf of Mexico, covers most of Alabama, time to show how changes in sea level over the past parts of Mississippi and Georgia, and a small portion million years or so shaped the Delta in the past, and of Tennessee (Ward et al. 2005). It has, by far, the larg- may influence its future. The Delta has occupied its est discharge (79,300 ft3/s [2246 m3/s]) of any Gulf current location for only the past few thousand years. of Mexico river system east of the Mississippi River Prior to that, it repeatedly migrated seaward, then embayment, representing 35 times the discharge of all landward, with falling and rising sea levels responding the other eastern Gulf Coast river systems combined. to the advance and retreat of continental ice sheets. Therefore, the Delta receives water and materials that Glacial ice never reached the Delta, buts its effects did. emerge from an expansive river system and serves as The last time the Delta sat at its current location a critical buffer for coastal areas to the south. Abun- was about 120,000 years ago. Evidence from the last dant rainfall and runoff in the basin supply the Delta 9000 years shows that the Mobile Bayhead Delta with ample water each year. Indeed, it is this abun- has responded to small increases in the rate of sea dance within the basin that ultimately creates the vast level rise. This knowledge raises questions about the wetland landscape we now observe as the Delta. sustainability of the bayhead delta’s present extent Ward et al. (this volume) helps us understand the in the face of projected sea level rise from global Delta’s complex geomorphology of landforms and warming. In this broader context, taking a long view, aquatic habitats, created by thousands of years of the Delta seems like a living organism, adapting and geological, fluvial, and geomorphic interactions. In changing to internal and external conditions. In some addition to active distributary channels, other major sense the Delta is simultaneously ephemeral and eter- landscape components include islands, flood basins, nal. As long as river water reaches the ocean, it will levees, bays, abandoned channels, lakes, sloughs, exist in some manner. and bayous. The levees observed frequently along As climate changed the Delta over the ages, so did distributary channels develop from deposition of it also influence the area’s ecosystems. Stults and sand and silt suspended in flood waters that overtop Axsmith (this volume) describe how even during river banks, and they build in height over time from times of extensive glaciers and sea level change, some repeated flooding. Flood basins are major landscape of the trees that define today’s Delta forests were pres- features that receive flood water when levees are over- ent, though pines were far less abundant. The Delta topped, or receive inflow through breaches (crevasse forests were not so profoundly impacted by distant channels) when water levels are below levee height. ice sheets as environments just to the north. Deeper in The dynamic ebb and flow of the Delta is seen as time, before the era of ice sheets, plants familiar to us inflowing floodwaters later drain back through these today were preceded by evolutionary ancestor species breaches into distributary channels, or over low and increasingly exotic taxa occupying very different points in levees. environments. Nevertheless, the comparative stabil- The greater Delta area is one of the most important and ity of plant species is remarkable and may reflect the valuable hydrologic resources in the US. It has existed stabilizing climatic influence of nearby Gulf waters. for tens of thousands of years, and currently receives water, sediments, and solutes from a substantial area As Goes the Water of Alabama and adjoining states via the river system The Delta is considered a drowned alluvial plain and of the Mobile River Basin. The Delta, which receives valley, where rivers flow through a relatively flat land- nutrients from this vast upgradient river system via the scape in broadly meandering channels and often over- Mobile River, also likely functions as a nutrient reten- top their banks. Ward et al. (this volume) describe how tion and transformation zone between the Mobile

186 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area River and Mobile Bay, thus lessening nitrate-nitrogen inflow, water quality on most of the Delta’s blackwater export to Mobile Bay and fragile coastal areas. Sea streams is determined by local conditions and hydro- level changes and water flow from the basin have over logical connectivity to the surrounding uplands. Rela- time carved an extensive and complex riverscape of tively small changes to freshwater inflow as a result of channels, lakes, bays, bayous, and bottomland forests, house construction, road building, or groundwater which both harbor extensive wildlife resources and pumping can result in rapid loss of stream head and act to sequester sediments, nutrients, metals, and rapid replacement by siltier waters from the Delta’s pesticides draining from the basin upstream. It is brownwater systems, or by more saline tidal waters. important to recognize that human activities in the Such changes can dramatically alter floral and faunal Mobile River Basin have profound impacts on the communities. In the longleaf forests above these Delta downstream. We do not currently have a thor- blackwater streams, mesic flatwoods and pitcher plant ough understanding of the fates of water and mate- bogs exhibit extraordinarily high diversity (up to 60 rial flow in the Delta, nor of the myriad roles of biota species/m2 [6 species/ft2]) in part because the plants in mitigating nutrient and other chemical inputs that have carefully partitioned habitat as a result of minute otherwise would be transported into Mobile Bay, and (1 cm [0.4 inch]) changes in depth to the water table. thence to the Gulf. Without this knowledge, effective Road building and house construction can severely management of the Delta will remain elusive. impair the sheet-flow hydrology that feeds these highly diverse wetland systems, resulting in the loss of Connectivity, the Whys and hundreds of species of plants and insects from rela- Wherefores tively small areas. An area’s ecosystems are defined by plant and animal The greater Mobile-Tensaw River area hosts a diverse communities that represent a dynamic temporal assemblage of freshwater crustaceans notable for and spatial mosaic of interconnected patterns and their ecological and economic importance. Stoeckel processes. Taking into account evidence of interacting and Helms (this volume) describe the importance geomorphologic, hydrologic, and ecological consid- of the salinity gradient where, in freshwater non- erations, a greater Mobile-Tensaw River area can tidal portions of the system, the crustacean fauna is be seen to include the Mobile-Tensaw Bottomlands dominated by crayfish, but, as tidal influences and and Delta, and adjacent contributing watersheds, salinity increase in the Mobile-Tensaw Delta, fiddler bluff lands, and pinelands. All told, the area extends crabs gradually replace crayfish in semi-terrestrial north from the head of Mobile Bay across the deltaic environments and Blue Crabs replace crayfish as the floodplain, including locally rising watersheds and dominant aquatic crustacean. Finally, as the Mobile- streams and the forests that shelter them, through the Tensaw Delta empties into Mobile Bay, a wide range Red Hills region that overlooks the northern extent of crabs, prawns, and shrimps depend on estuaries as of the Delta. We can appreciate a greater Mobile- critical habitat for various life-history stages. Stoeckel Tensaw River area as a large landscape of intimately and Helms then demonstrate how protection and connected ecosystems without any formal jurisdiction maintenance of dynamic natural flows, seasonally across northeastern Mobile County, southeastern flooded areas, groundwater fluctuations, and general Washington County, southern Clarke County, south- hydrologic connectivity are of considerable impor- western Monroe County, western Escambia County, tance to the conservation of crustaceans in the greater and northern Baldwin County (Figure 23-1). Mobile-Tensaw River area. This approach includes Within this area, there is ample evidence of tightly careful consideration of future construction of dams coupled ecological drivers and stressors that interact and impoundments, road crossings, and bridges across temporal and spatial gradients, resulting in a where maintenance of salinity gradients is paramount. high degree of ecological connectivity that underpins Alterations in freshwater inflow/saltwater intrusion and essentially defines the area. Finch and Plumb (this patterns, as a result of water withdrawal for agricul- volume) describe how hydrological alteration affects tural and industrial uses, and drought, can poten- both blackwater streams and upland pitcher plant tially have devastating effects. Large groundwater bogs surrounding the Delta. Unlike water conditions withdrawals have been reported in the coastal zones of the interior Delta, which are primarily controlled of Baldwin and Mobile counties. Such changes in by inflow from hundreds of miles upstream or by tidal supply, coupled with changes in rainfall patterns (e.g.,

Synthesis - Chapter 23 187 Figure 23-1. The greater Mobile-Tensaw River area. (Image credit: Mark Bailey, Glenn Plumb, Bill Finch, Greg Waselkov, and Fred Andrus.)

188 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area drought), are viewed as a major threat to the sustain- uplands and wetlands (e.g., indigo snakes, black bear, ability of the Blue Crab fishery of Alabama. freshwater turtles, numerous bird species). Williams et al. (this volume) describe how the three coun- Heterogeneity within and between upland and ties surrounding the Delta support what is likely the wetland landscape communities is promoted and greatest turtle diversity in the world. Protection of the mediated by fire, which makes continuity between Delta alone, without giving attention to the surround- vegetation communities and across broad landscapes ing upland habitat, is not likely to benefit turtles, since particularly important. Finch and Plumb (this volume) some of these turtle species are primarily upland describe how human-induced habitat fragmentation species and use the Delta rarely or not at all. Most of has changed the fire ecology of the greater Mobile- those that do are threatened primarily by loss of suit- Tensaw River area so that sources of ignition must be able upland (non-flooded) habitat for egg laying and artificially introduced into ecosystems and the fires hatchlings. A great number are lost while trying to lay must be managed on a regular basis. Loss of connec- eggs on the grassy highway medians that run through tivity between longleaf uplands and surrounding the lower Delta. broadleaf and wetland communities has resulted in greatly reduced fire frequencies in those areas. These Historical Connections “ecotonal” marginal areas are integral to the existence of many if not most of the listed engendered, According to the Alabamas, descendants of the origi- threatened, and special concern species (e.g., Red nal inhabitants of the place that would become the Hills Salamander, Reticulated Flatwoods Salamander, state of Alabama, their ancestors “sprang out of the American chaffseed, pot of gold lily, Wherry’s pitcher ground, between the Cahawba and Alabama Rivers” plant). In the greater Mobile-Tensaw River area, along (Lankford 2011, p 111). Prior to that distant time, the lower Alabama River at the top of the Delta, tree there had been only the Above World of flying crea- species diversity ranks among the highest of any place tures and the Under World of swimming creatures, in North America, with some 25 species of oaks, 7 until an earth-diver created the Middle World where species of magnolias, and many other species occu- humans reside. While visiting some of his Alabama- pying a very small area. There is indication that the Coushatta Indian friends in 1930s Texas, Martin Red Hills of southwest Alabama support the highest (1977, p 2-3) transcribed a traditional Alabama legend oak diversity (28+ oak species) for North America. about an earth-diver, a crayfish who accomplished Occasional fire trickling in from the adjacent longleaf this feat. forests is necessary to maintain diversity, and loss of “Once, long ago, before the time of the oldest people, connectivity between these areas and the surround- water covered everything.” A log raft drifted on the ing longleaf ecosystem has resulted in decline in oak great ocean, occupied only by some small animals. regeneration and the decline or loss of many other “Land is somewhere beneath the water,” said their species. leader, Horned Owl. “Who will make it appear?”

Borden and Major (this volume) describe how habitat Beaver, a strong swimmer, tried first, but he tired diversity, environmental patchiness, and natural and and returned to the raft. Frog tried next, but Garfish human-induced disturbances affect the area’s ability chased him, and he too returned to the raft. Horned to serve as a breeding ground for seasonally resident Owl, in some desperation, asked Crawfish to look for birds and as a stopover refuge for refueling migra- land, and Crawfish eagerly answered, “I am ready.” tory birds. The greater Mobile-Tensaw River area Garfish did not think him good to eat, and he did not supports high bird diversity (over 200 species) and tire like Beaver. Finally he reached the floor of the provides critical habitat for avian populations across great ocean. the Eastern Gulf Coastal Plain and much of the East- ern Deciduous Forest ecoregions. Best (this volume) Crawfish used his wide tail “to scoop mud into a great and Plumb and Finch (this volume) also describe chimney. He worked rapidly, building it higher and how movement of wildlife between floodplain and higher, where it began to spread and form a mass of uplands has already been compromised by growth soft earth. The birds and animals on the raft looked of residential and commercial areas, roads, and other at the new earth and agreed that Crawfish had done developments. Many endangered or rare species are a good job.” highly dependent on continued connectivity between

Synthesis - Chapter 23 189 This origin myth of the Alabamas, and its parallels by other southeastern Indians, accounts for creation of the land, the “earth island,” which Mississippian societies replicated in microcosm with their earthen platform mounds (Knight 2006). Mound A at the Bottle Creek archeological site, in the heart of the Mobile-Tensaw Delta, stands today as an impressive example of this Mississippian cosmo- logical symbol.

Likewise, the discovery in 1922 of a massive stone pipe carved in the shape of a crayfish, near the large Mississip- pian shell mound on Dauphin Island, a site subsidiary to the Bottle Creek chiefdom, is most enlightening (Figure 23-2). Sculpted in the 11th century at the Figure 23-2. A cast of a flint clay stone pipe, created in the form of a crayfish greatest of the Mississippian centers, effigy at Cahokia in the 11th century AD and discovered in 1922 by Theodore Cahokia, in the central Mississippi R. Vaughan, Sr., on Dauphin Island. The earth-diver, Crawfish, created the River valley, this remarkable portrait earth island according to Mississippian mythology. (Courtesy of Theodore R. of the land’s creator was buried in the Vaughn, Jr., and Gregory Vaughn.) 13th century at the southern edge of the North American “earth island” (Emerson et al. 2003). A more recent archeological discovery in Orange Beach, a 10th-century crayfish effigy modeled in clay as if crawling over the lip of a pottery vessel, perhaps represents the legendary earth-diver’s “great chimney,” and testifies to the time depth of this ancient belief in this part of the world (Price 2009, Figures 6-10b) (Figure 23-3).

Morgan (this volume) describes how this region’s time depth of human history can be read literally in archeological layers buried by the near bottomless alluvium of the Mobile- Figure 23-3. A stylized crayfish effigy on the rim of a Weeden Island Punc- Tensaw Delta. In fact, only the most tated ceramic vessel, Bayou St. John site (1BA21), Late Woodland period, recent 3000 years of the delta’s arche- Orange Beach, Alabama. (Courtesy Center for Archaeological Studies, Uni- ology sits close enough to the present- versity of South Alabama.) day land surface to be accessible for study. One must look to the adjacent bluffs and hilly uplands on either side of the delta to find archeological exposures documenting an additional 8000 or more years of human occupation

190 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area in the greater Mobile-Tensaw River area. In “The American occupants proved adaptive and the transi- Forks” between the Tombigbee and Alabama rivers, in tion was long and contested (Waselkov, this volume). Clarke County, stone tools dating from 11,000 to 3400 Although the long-sought battle site of Mabila, where years ago (the Paleoindian and Archaic periods) are Hernando de Soto’s Spanish army narrowly escaped frequent finds, especially around outcrops of Talla- destruction at the hands of the Chief Tascalusa’s hatta Formation sandstone and agate, both quarried by warriors, remains undiscovered, many other sites of ancient flintknappers. But equally impressive are the national historical significance are located in the area. Delta middens at Douglas Lake and Bryant’s Landing, The first location of colonial Mobile, earliest French where fired clay balls from Late Archaic earth ovens town successfully established on the Gulf Coast, has and Gulf Formational pottery, the region’s oldest, are been extensively investigated by archeologists and visible in eroding banks at low water. determined eligible for designation as a National Historic Landmark, an honor already bestowed on 6 A magnificent Middle Woodland shell midden faces treasured sites and structures in the greater Mobile- the lower Delta at Blakeley State Park, a prime candi- Tensaw River area (Schneider et al., this volume). date for National Historic Landmark status as well Other likely candidates for NHL status include Fort as a refuge for rare, endemic, shell-midden adapted Stoddert, Old St. Stephens Park, Fort Mims Park, plants. David Morgan’s analysis of a midden at Little Historic Blakeley State Park, Africatown (Figure 23-4), Lizard Creek is, so far, the only modern study of a and the site of Mount Vernon Arsenal/Barracks and Late Woodland site in the Delta proper, although a Searcy Hospital. Most of these important historic sites flurry of Woodland research on the Alabama coast are owned by the State of Alabama and administered suggests what potential these data-rich sites offer for by a variety of agencies that struggle to maintain them understanding long-term human adaptation to the for public interpretation. Searcy Hospital complex, region’s fisheries (Morgan 2003a; Price 2008, 2009; where several 1830s arsenal buildings have collapsed Reitz et al. 2013; Hadden 2015). Biologists in search in recent years, remains in the greatest jeopardy. of historic baseline data on fish and shellfish species ranges and population structures, as well as paleocli- In recent years the region’s historical connections to matic data, should consider collaborating with arche- forest products industries, especially tree harvest and ologists, who have been retrieving faunal samples from sites threatened by unceasing waterfront development, a catastrophic oil spill, and shoreline and riverbank erosion. Vastly improved methods of stable isotope and elemental analysis of animal growth structures, such as teeth and mollusk shells, are beginning to reveal the selective and seasonal nature of hunting and fishing in the greater Mobile-Tensaw River area with a level of precision previously thought unachievable. There is some irony in the timing of these analytical break- throughs, which are occurring as sites are disappearing at an alarming rate. Preservation of some of these increasingly scarce middens has become a matter of urgency.

The arrival of Europeans and Afri- Figure 23-4. Photograph of Cudjo Lewis, enslaved in Africa and transported cans—the latter most often in bond- to Mobile in 1861 aboard the Clotilda, at home in Africatown, circa late age—eventually transformed the 1920s. (Overbey Collection, Doy Leale McCall Rare Book and Manuscript region’s cultural landscape, but Native Library, University of South Alabama.)

Synthesis - Chapter 23 191 turpentine production, are celebrated at Stockton Despite their importance to humans as disease Sawmill Days. Sponsored by the Stockton Heritage vectors, we have barely begun to count the unfathom- Association, the annual event hosts a variety of logging ably diverse insect taxa of the Delta (Burkett-Cadena, and folk craft demonstrations, including participation this volume; McCreadie and Adler, this volume). We by the Gees Bend quilters, local beekeepers, and a have a better handle on fishes, with the knowledge working oxen team. These days logging occurs most that Alabama supports more fish species than any intensely in the uplands of the greater Mobile-Tensaw other state. The Mobile Basin alone accounts for most River area, since the largest paper and pulp compa- of that. Its endemic fauna once included 40 fish, 33 nies sold most of their holdings in the Delta to the mussels, and 110 aquatic snails. Though extinctions US Army Corps of Engineers or to Forever Wild, or, in this basin exceed any other river system in North as International Paper Company did with the Bottle America, it still contains 60% of North America’s Creek archeological site, donated Delta lands to the mussel species, 52% of its turtle species, 43% of its State of Alabama. Those sales by wood products snail species, and 38% of its fish species (DeVries and companies led to the closure of many generations-old Wright, this volume; Gangloff, this volume; Williams fishing and hunting camps in the Delta, with lingering et al., this volume). The Cahaba, a single small river hard feelings on the part of the dispossessed to this tributary to the lower Alabama, supports more than day. Still, vast areas are devoted to game management twice as many fish species as the entire state of Cali- and hunting remains a major way the public interacts fornia. The turtle diversity concentrated in the hydro- with that landscape. logical unit surrounding the lower Alabama and lower Tombigbee rivers and Mobile-Tensaw Delta is rivaled A Greater Mobile-Tensaw River Area globally only by a much larger area in northwest India. In 1973, Professor Mount of Auburn University Terrestrially, the area is equally significant. Duncan described the Delta of the Mobile Bay drainage as one (this volume) describes some of the reasons for the of the most important wetland areas in the country. high biodiversity and endemism of the greater Mobile- Despite then prevalent land and water uses, Professor Tensaw River area. The Red Hills region at the head of Mount felt the area retained many of its natural values the Delta is now recognized as the national center of and was of considerable national importance. In a oak diversity. Surveys in pinelands on the bluffs near time before email and social media, he sought out and the Delta have revealed some of the highest counts of talked with many people who shared his assessment vascular plant species anywhere in North America, up and also felt strongly that the area’s resources and to 60 species/m2 [6 species/ft2]. Terrestrial endemism values deserved protection (Mount 1973; Plumb, this in the region ranges from the tree-like Red Hills azalea volume). In 1979 the National Park Service, through to one of the world’s most distinctive salamanders. its responsibilities to identify and recognize National But, importantly, we are acquiring a better under- Natural Landmarks, updated Professor Mount’s standing of the natural extent of the greater Delta, report and indicated that (NPS 1979): what it encompasses, and its critical context. While the delta still retains its natural values We have begun to better appreciate that the Delta to a significant degree, threats to the integrity region is much broader than the deltaic portion of of the natural environment do exist. If these the floodplain. And the floodplain’s significance— threats are left unchecked, it is very likely that ecologically, biologically, historically, culturally—is the significant natural values would be lost. almost entirely dependent on the lands surrounding As this present report well documents, much has been it. Locals have long recognized this fact. When they learned about the significance of the greater Mobile- refer to the Delta region, they draw from their expe- Tensaw River area in the intervening years. Yet we also rience. The floodplain is simply the cornerstone of a concede that we have only begun to appreciate and much larger area of contiguous wildness, history, and protect one of North America’s and the world’s most unusual biodiversity. Recent scientific evidence and significant natural, historical, and cultural regions. popular accounts of the Delta region, including the The Mobile River Basin, whose waters feed the Delta, PBS movie America’s Amazon, extend the concept of is now recognized as one of the temperate world’s the Delta up the lower Alabama River deep into the most biologically diverse freshwater environments. Red Hills, and west and east into the longleaf pine region that surrounds it.

192 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area There is merit in denoting a Greater Mobile-Tensaw River Area. The deltaic floodplain is one of the best preserved in North America and is extraordinarily rich in comparison to any other delta. But the Delta itself is not the richest or even the most biologically and historically significant portion of the Delta region. So many of the globally significant attributes we associate with the Delta region—much of its turtle diversity, its fish diversity, its flowering plant diversity, even the Delta black bears, the last breeding population of bears in Alabama—are rarely or never found in the floodplain itself, but rather spend most of their lives Figure 23-5. “Mobile, taken from the marsh opposite the city near Pinto’s residence,” in the bluffs, pinelands, broad- watercolor by William Todd, 1841. (Courtesy of the Miriam and Ira D. Wallach Division of Art, Prints, and Photographs: Print Collection, New York Public Library leaf forests, and feeder streams http://digitalcollections.nypl.org/items/510d47d9-7c72a3d9-e40-e00a18064a99.) that surround the Delta.

Similar reassessments have given us a much better It’s no accident that the city of Montgomery, near understanding of the Delta’s human history. The the head of the Alabama River, was the Cradle of the rivers of the Mobile Basin were great corridors of Confederacy, and that Mobile and its surrounding North America’s pre-Columbian and modern history. Delta and Bay was a strategic location from the start of Shellmounds, complex architectural features of the Civil War until its very last battle. And it’s no acci- some of the region’s earliest human communities, dent that the Alabama River and its surrounding land- abound. The north-central Gulf Coast’s largest and scape became the cradle of the Civil Rights movement most significant Mississippian-era city, a sprawling where the march from Selma to Montgomery became complex of mounds, plazas, and canals, sits in the a turning point in the nation’s history. midst of the Delta. The Redstick War, instigated in the Mobile River Basin, ushered in the Jacksonian popu- The evidence is clear that for millennia, the Greater list movement that forcibly removed most American Mobile-Tensaw River Area has been a place where Indians to lands in the west, and thereby changed the people have gathered to share in nature’s beauty and social fabric of the eastern United States. bounty to improve their quality of life. More people are exploring and using the Greater Mobile-Tensaw The Alabama and Tombigbee rivers and their River Area at the present than perhaps at any time in surrounding landscape was also a central corridor its long history. Thus, there are now more opportuni- for 19th-century American politics. These rivers were ties than ever for continuing our education and study major highways for the antebellum South’s cotton and of the Greater Mobile-Tensaw River Area, and shar- slaves, wealth and oppression (Figure 23-5). Slave- ing in stewardship that will ensure its continued well- holding counties along these rivers influenced national being as central to the American experience. politics in the mid-19th century, and the community of Africatown, overlooking the Delta, traces its origins to the Clotilda, last slave ship to dock in North America.

Synthesis - Chapter 23 193

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232 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area List of Contributors

Diamond-backed Watersnake (Nerodia rhombifer). Photograph courtesy of Hunter Nichols.

Peter H. Adler is an entomologist with emphasis Energy Economics and Urban Studies, and has given a in medical entomology. Professor Adler received his number of invited lectures in the US and abroad. His PhD from Pennsylvania State University and has been primary research interest is environmental and natu- at Clemson University since 1984. He has conducted ral resource economics. In addition, he is interested research from the tropical rainforests to the high in computational economics and spatial econometrics arctic, authored several hundred publications, includ- applied to regional science. ing the award-winning book The Black Flies (Simuli- idae) of North America (co-authored with Douglas C. C. Fred Andrus is Associate Professor and Chair of Currie and D. Monty Wood, Royal Ontario Museum, the Department of Geological Sciences at the Univer- 2004), and served as mentor for dozens of graduate sity of Alabama. His research interests center at the students. His current interests are the systematics and intersection of geology, paleontology, and archeol- ecology of black flies and the structure and function ogy. He often analyzes the geochemistry of skeletal of the butterfly proboscis. remains (e.g., sea shells, coral, vertebrate remains) from fossil deposits and archeological sites to infer Ermanno Affuso, Assistant Professor of Econom- past climate and environmental conditions. Most of ics at the University of South Alabama, earned his his research in the greater Mobile-Tensaw River area PhD in Applied Economics from Auburn University focuses on the growth and chemistry of marsh clam and an MS degree in Civil Environmental Engineer- shells (Rangia cuneata), a common shell in archeolog- ing from the Bari Polytechnical University, Italy. He ical sites and fossil deposits, as a way to measure past is a member of the American Economic Associa- human-environment interaction. tion and the Urban Economics Association and has published in several scholarly journals, including

List of Contributors 233 J. J. Apodaca is a professor of conservation biol- and Southwestern Association of Naturalists. He ogy at Warren Wilson College and Senior Co-Chair has served as Editor of the Journal of Mammalogy, for Partners in Amphibian and Reptile Conservation Mammalian Species, and The Southwestern Natural- (PARC). His research interests include combining ist, and he recently co-authored Mammals of Alabama genetics techniques with spatial ecology to inform with Julian Dusi (University of Alabama Press, 2014). the management of imperiled species and using geographic information systems to identify priority Joel A. Borden is an Instructor in Biology at the areas for the conservation of amphibians and reptiles. University of South Alabama where he teaches General He has been conducting herpetological conserva- Biology, Ornithology, and Terrestrial Vertebrate Biol- tion research for nearly 15 years and has led efforts to ogy. In addition, he has over 10 years of research identify Priority Amphibian and Reptile Conservation experience studying vertebrate communities along Areas in the southeastern United States. the central Gulf Coastal Plain. Joel has a BS in Biology from Jacksonville State University and an MS in Biol- Michele Archie is a principal at The Harbinger ogy from the University of South Alabama, where he Consulting Group, specializing in economic analy- studied the effects of silviculture practices on herpe- sis and community engagement around nature, heri- tofauna in southern Alabama pine ecosystems for his tage, cultural tourism, and protected areas. This work, thesis. He is currently examining how disturbance spanning more than a dozen years, includes analysis affects ecosystem functions in the Mobile-Tensaw and projections of economic benefits, evaluation of River Delta and Mobile Bay Estuary. management alternatives, and strategy development for leveraging natural and historical resources and Nathan D. Burkett-Cadena is Assistant Profes- special designations for community and economic sor at the Florida Medical Entomology Laboratory, a benefit. Her contribution to this report draws on Research and Extension Center of the University of research conducted in 2013-14 on potential economic Florida. His current research focuses on the ecology impacts of conservation designations in the Mobile- of viruses transmitted by mosquitoes and other blood- Tensaw Delta and bluffs region, and on compatible feeding arthropods. Burkett-Cadena has co-authored tourism development. more than 30 scientific articles on topics related to mosquito ecology, virus transmission, and novel meth- Brian Axsmith is a Professor in the Biology Depart- ods for collecting insects of medical importance. He is ment at the University of South Alabama, where he author of the book The Mosquitoes of the Southeastern teaches Ecology and Evolution, Evolutionary Biology, United States, released by University of Alabama Press and the Evolution of Vascular Plants. He is also Grad- in 2013. uate Coordinator for the department’s MS program. His main research interests are the Mesozoic paleo- Larry Davenport holds a PhD in biology from botany of eastern North America and China, and the University of Alabama and is currently Professor the Neogene of the Gulf Coastal Plain. Much of his of Biological and Environmental Sciences at Samford work has dealt with the evolution of conifers, but he University, Birmingham. In 2007, Dr. Davenport was has worked on nearly every major group of vascular named Alabama Professor of the Year by the Carn- plants. When not working on fossil plants, he plays egie Foundation for the Advancement of Teaching. bass in several local groups. He is an expert on Alabama’s plant life, aquatic plants, wetlands, and rare species, including the Cahaba Lily Troy L. Best completed BS degrees in Biology, and Confederate Daisy. His most recent research Anthropology, and Secondary Education at Eastern focuses on the potential effects of climate change on New Mexico University and MS and PhD degrees Alabama’s plant life and the history of botany. He in Zoology at the University of Oklahoma. He is also writes the Nature Journal column for Alabama Professor of Biological Sciences at Auburn Univer- Heritage magazine, and a collection of those essays sity and Curator of the Auburn University Collection appeared in Nature Journal (University of Alabama of Mammals. In addition to Alabama, he has stud- Press, 2010). ied mammals in Argentina, Canada, Kenya, Mexico, South Africa, Swaziland, and much of Europe. He has received honors from the American Society of Mammalogists, Southeastern Bat Diversity Network,

234 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area Dennis DeVries is a Professor and an Assistant Michael Gangloff is an Assistant Professor of Director for Research Programs in Auburn Univer- Biology at Appalachian State University in Boone, sity’s School of Fisheries, Aquaculture and Aquatic NC. He received his PhD from Auburn University in Sciences, where he is involved in research, teaching, 2003 and has been working with freshwater mollusks and part-time administration. He teaches a course in in Alabama and the southeastern US for 17 years. Fish Ecology and 2 courses in the Auburn University Gangloff has authored more than 30 peer-reviewed Sustainability minor. His research program addresses publications, many of which focus on the ecology questions related to aquatic food web interactions, and systematics of the Southeast’s unique freshwater fish early life history, and conservation and ecology mollusk assemblages. of threatened or endangered species. DeVries has published more than 80 peer-reviewed papers and Bonnie Gums is a Research Associate at the Center book chapters, and has edited 1 book. for Archaeological Studies at the University of South Alabama in Mobile. She received her BA and MA from R. Scot Duncan is a Professor of Biology at Southern Illinois University focusing on French colo- Birmingham-Southern College. His upbringing in a nial archeology in southwestern Illinois. She joined family of ardent birdwatchers shaped his conserva- the Center for Archaeological Studies in 1995 and tionist outlook and career as a scientist. Scot received directs cultural resource studies focusing on historic a BS in Biology from Eckerd College (1993), and an archeological sites along the northern Gulf Coast. MS (1997) and PhD in Zoology (2001) from Univer- Her most recent publication is entitled Archaeology sity of Florida. He and his students study the ecology at LaPointe-Krebs Plantation on the Mississippi Gulf of endangered species and threatened ecosystems in Coast (co-edited with Gregory A. Waselkov, Missis- Alabama. Scot wrote the award-winning Southern sippi Department of Archives and History, 2015). Wonder: Alabama’s Surprising Biodiversity (Univer- sity of Alabama Press, 2013, foreword by Dr. Edward Craig Guyer is a Professor in the Department O. Wilson). The book interweaves several scientific of Biological Sciences and Curator of Herpetology disciplines to explain why Alabama hosts more species at Auburn University. He received a BS in Zoology than any other state in the eastern US. at Humboldt State University, an MS in Biology at Idaho State University, and a PhD in Biology from the William Finch, senior fellow with the Ocean Foun- University of Miami. His work centers on population dation, is a well-known writer and natural historian. and community ecology of southeastern amphibians He has served as Director of Conservation with the and reptiles, especially of the Lower Coastal Plain. He Alabama chapter of The Nature Conservancy, as has produced 27 graduate students, authored more executive director of Mobile Botanical Gardens, and than 100 scientific articles, and authored or edited 4 as assistant managing editor at the Mobile Press Regis- books, including Turtles of Alabama (with Mark A. ter, where his environmental writing received numer- Bailey and Robert H. Mount, University of Alabama ous regional and national awards. Finch is the author Press, 2015). of Longleaf: Far as the Eye Can See, published in 2012 by the University of North Carolina Press. Brian Helms is Assistant Research Professor in the Department of Biological Sciences at Auburn Univer- Richard S. Fuller, now retired, has practiced sity and manages the Invertebrate Collection at the archeology on the US Gulf Coast for 40 years, at vari- Auburn University Museum of Natural History in ous times affiliated with Harvard University’s Lower Auburn, Alabama. His research primarily focuses on Mississippi Survey, the University of South Alabama’s freshwater environments spanning organismal biol- Archaeology Laboratory, the University of Alabama’s ogy, system ecology, and resource management. This Gulf Coast Survey, and with Coastal Environments, includes investigating the natural history, population Inc. His specialty is Native American ceramics, with a connectivity, ecology, and conservation of crayfishes; particular interest in potteries created by the Missis- quantifying physical and biological effects of human sippian, protohistoric, and early historic period Indi- disturbance on stream systems; and development of ans of the north-central Gulf Coast. tools for stream assessment and restoration. He is also enjoys student mentoring, teaching, and biodiversity conservation outreach, but really he just digs crayfish.

List of Contributors 235 Alex Maestre is a civil engineer from the Univer- in federal, university, and private sector archeological sidad de Los Andes (Colombia), where he devel- enterprises. oped environmental programs and models for large marsh ecosystems. He developed strategies for the Hunter Nichols is a photographer and filmmaker implementation of surface water management plans based in Birmingham, Alabama. He has been produc- in compliance with federal and local regulations. At ing conservation-related productions since 2006 the University of Alabama, his dissertation involved and graduated from Auburn University with a BA in the development of the National Stormwater Qual- Radio, Television, and Film. His most recent feature River Dreams ity Database. As a post-doctoral fellow, he studied the film, , chronicles a 44-day solo canoe estimation of nutrient fluxes for the Mobile Alabama trip from Birmingham, Alabama to Horn Island off River System and use of cyberinfrastructure for study- Mississippi’s Coast. You can find his photography and ing coastal hazards and ecological protection. Maestre other film projects at www.hnproductions.com. is currently working at the NOAA National Water Glenn Plumb is the National Park Service Chief Center in Tuscaloosa. Wildlife Biologist, having worked as a biologist at C. Smoot Major is a Senior Instructor/Research Grand Teton National Park 1990-1993, Badlands Faculty member of the Department of Biology at the National Park 1993-1998, and Yellowstone National University of South Alabama. Previously he served as Park 1998-2010. He served in Mobile, Alabama on the State Ecologist and Natural Heritage Coordinator for the Department of Interior Strategic Science Group the State of Tennessee, as well as coordinator for the and Incident Command Team for Deepwater Horizon central basin’s Partners in Flight program, a coopera- and developed a lasting interest and fondness for the tive partnership of agencies and organizations dedi- Mobile-Tensaw River area. cated to conserving land birds, both resident and Ben Raines has explored and reported on Alabama’s migratory. His research now focuses primarily on the wild places for 15 years. His print work has garnered Mobile-Tensaw river basin. He has mentored 11 grad- numerous national environmental journalism awards, uate students and 2 honors students in research rang- and America’s Amazon, a film he wrote and produced, ing from ecotoxicology and plant community ecology is airing across the country on public television. His to avifauna and herpetofauna. underwater film work has appeared in documentaries John W. McCreadie is an insect ecologist, on the Discovery Channel and National Geographic with a special interest in aquatic insects. Professor Television. He reports on nature for the Alabama McCreadie received his PhD from Memorial Univer- Media Group and guides sightseeing and fishing trips sity, St. John’s, Newfoundland in 1991 on the ecology in the Delta and Mobile Bay, and scuba diving trips in and cytogenetics of larval black flies (Diptera: Simuli- the Gulf of Mexico. idae). He has been at the University of South Alabama Antonio B. Rodriguez is a Professor at the since 1999 and has undertaken research in Canada, University of North Carolina at Chapel Hill, Institute the US, the Galapagos Islands, and mainland South of Marine Sciences and has been working there since America. Currently, his primary interests are in the 2005. He graduated from Hamilton College in 1994 structuring processes responsible for aquatic commu- with a BA in Geology and Rice University in May 1999 nities and insect-fungal symbiosis. with a PhD in Geology and Geophysics. After gradu- David W. Morgan, PhD, works for the National ation, he stayed at Rice as a Postdoctoral Research Park Service, where he serves as the Regional Arche- Associate (May 1999 to January 2000) and Lecturer ologist for the Southeast and Director of the South- (January to July 2000). In August 2000, he moved to east Archeological Center (SEAC). His research the University of Alabama, Department of Geologi- focuses on the prehistoric and historic southeastern cal Sciences as an Assistant Professor and left in 2005 United States, particularly on underwater archeologi- as a College of Arts and Sciences Leadership Board cal methods and resource management; the archeol- Faculty Fellow and the George Lindahl Fellow. His ogy of the African Diaspora and creolization; settle- research is on estuarine evolution and he examines ment and subsistence patterns of the indigenous Gulf anthropogenic and climate impacts on bayhead-delta, Coast; and changing concepts about how heritage is bay, marsh, oyster-reef, and beach environments perceived and valued. Prior to SEAC, Morgan worked along the northern Gulf of Mexico and southern Atlantic Coast.

236 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area David B. Schneider’s firm, Schneider Historic teaches biology students and continues her paleobo- Preservation, LLC, established in 1999, provides a tanical research at the University of South Alabama. full range of historic preservation consulting services. Mr. Schneider’s historic preservation career spans Amelia K. Ward is Professor Emeritus in the 34 years, during which time he has successfully Department of Biological Sciences at the University of completed a diverse range of projects for both private Alabama, where she served as Director of the Center and public sector clients in 15 states and one other for Freshwater Studies for 19 years. She received her country. He specializes in community preservation PhD from Michigan State University and currently planning, design review in historic districts, historic serves as treasurer and on the Board of Directors resource documentation, and certification of projects of the Association of Ecosystem Research Centers for historic rehabilitation tax credits. (AERC) and on the Board of Directors of the NSF- funded National Ecological Observatory Network George W. Shorter, Jr., now retired, had a success- (NEON). Her research interests are in ecology of ful career as a landscape architect before discovering microbes and aquatic plants in lakes, streams, rivers, the joys of archeology and obtaining an MA degree in and wetlands, with emphasis on productivity, nitro- that field from Louisiana State University. During his gen cycle processes, interactions with higher trophic 20-year career at the University of South Alabama’s levels, and roles in nutrient/organic matter retention Center for Archaeological Studies, Shorter investi- and transformation. gated Late Woodland sites in the Forks region of the Mobile-Tensaw Delta, a French colonial settlement on G. Milton Ward is Professor Emeritus in the Dauphin Island, and Fort Morgan, scene of the battle Department of Biological Sciences at the University of Mobile Bay in 1864. More recently he has devoted of Alabama, Tuscaloosa. An Alabama native and 1969 years to the study of Old St. Stephens, Alabama’s terri- graduate of the University of Alabama, he received his torial capital from 1817 to 1819. PhD from Michigan State University in 1977. Ward’s research specialty is aquatic ecology, particularly Jim Stoeckel is an Associate Professor in the School streams, rivers, and wetlands, and he has published of Fisheries, Aquaculture, and Aquatic Sciences at over 30 refereed articles and book chapters focused Auburn University where he teaches courses on aqua- on aquatic ecology and hydrologic/ecological inter- culture and conservation. He studies aquatic inverte- actions in lotic and wetland ecosystems. He is a past brates with an emphasis on freshwater mussels and president of the Alabama Section of the American crayfish in the southeastern United States. His research Water Resources Association and is currently Curator examines effects of stressors, such as suspended of Insects at the Alabama Museum of Natural History. solids, agrochemicals, and altered flow regimes, on mollusks and crustaceans, and how these stressors Gregory A. Waselkov, Professor of Anthropology may affect population dynamics and viability. Stoeckel at the University of South Alabama in Mobile, directs has authored over 30 peer-reviewed publications. He the Center for Archaeological Studies and writes on is a member of the Freshwater Mollusk Conservation Southeastern Indians, French colonists, and shell William Bartram on the Society, and serves as Secretariat for the International middens. His books include Southeastern Indians Association of Astacology. (University of Nebraska Press, with Kathryn Braund, 1995), A Conquering Spirit: Fort Debra Z. Stults earned an MS in Biology and PhD Mims and the Redstick War of 1813-1814 (University of in Marine Sciences, with studies focusing on paleo- Alabama Press, 2006), and Archéologie de l’Amérique botany, palynology, paleoclimatology, and paleo-CO2 coloniale française (Lévesque éditeur, with Marcel atmospheric concentrations during the Late Neogene. Moussette, 2013). He is a past editor of the journal She is currently involved in research on Upper Creta- Southeastern Archaeology and current president of the ceous through Pleistocene floras from various sites Southeastern Archaeological Conference. along the Mid-Atlantic to Gulf of Mexico Coastal Plains. The route to paleobotany was not a direct one, as a BS degree in Medical Technology attests. In addition to her clinical duties at the University of South Alabama Children’s and Women’s Hospital, she

List of Contributors 237 Jennifer M. (Jen) Williams serves as the Federal of Science, the Crafoord Prize (equivalent of the Coordinator for Partners in Amphibian and Reptile Nobel, for ecology) of the Royal Swedish Academy Conservation (PARC), a position hosted by the Wild- of Sciences, and the International Prize of Biology of life Conservation Branch in National Park Service. Japan; and in letters, two Pulitzer Prizes in non-fiction, Jen is the only full-time, paid national employee for the Nonino and Serono Prizes of Italy and COSMOS PARC and is afforded the opportunity to work with 11 Prize of Japan. For his work in conservation he has agencies that serve on PARC’s Federal Agency Steer- received the Gold Medal of the Worldwide Fund for ing Committee, as well as many other partners across Nature and the Audubon Medal of the Audubon Soci- the US and internationally. She obtained her PhD in ety. He is currently Honorary Curator in Entomology Forest Resource Science from West Virginia Univer- and University Research Professor Emeritus, Harvard sity, her MS in Wildlife and Fisheries Science from University. Pennsylvania State University, and her BS in Biology from Gannon University. Russell Wright is an Associate Professor and Extension Specialist in the School of Fisheries, Aqua- Edward O. Wilson is generally recognized as one culture, and Aquatic Sciences at Auburn University. of the leading scientists in the world. He is also recog- Dr. Wright teaches courses in fisheries management nized as one of the foremost naturalists in both science and biology. His extension program broadly covers and literature, as well as synthesizer in works stretch- aquatic natural resource education and manage- ing from pure biology across to the social sciences and ment, with specific focus on recreational fishing, pond humanities. Wilson is acknowledged as the creator of management, and conservation issues. Dr. Wright’s two scientific disciplines (island biogeography and research addresses both basic and applied questions sociobiology), three unifying concepts for science in systems from coastal waters, rivers and streams, to and the humanities jointly (biophilia, biodiversity private farm ponds. His efforts have produced over studies, and consilience), and one major technologi- 50 peer-reviewed journal articles and book chapters, cal advance in the study of global biodiversity (the as well as many extension publications and digital Encyclopedia of Life). Among more than 100 awards resources. he has received worldwide are the US National Medal

Photograph of American Coots (Fulica americana), Mobile-Tensaw Delta, Alabama. Photograph courtesy of Hunter Nichols (http://www.hnproductions.com).

238 A State of Knowledge of the Natural, Cultural, and Economic Resources of the Greater Mobile-Tensaw River Area ON THIS PAGE Photograph of Mobile-Tensaw Delta, Alabama. Photograph courtesy of Hunter Nichols (http://www.hnproductions.com).

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