ABSTRACT
CLARK, RACHEL KATHLEEN. Guide to the Vascular Flora of Kitty Hawk Woods (Dare County, North Carolina). (Under the direction of Alexander Krings).
One of ten reserves in the North Carolina Coastal Reserve system, Kitty Hawk Woods (Dare County, North Carolina) is a nationally significant natural area that is comprised of 760 ha of forested wetlands, sandy wooded ridges, and brackish marshes. It contains one of the last remaining examples of the rare Maritime Deciduous Forest (S1 G1) community type and the most extensive example of Maritime Swamp Forest (S1S2 G2) in North Carolina. Topographically, Kitty Hawk is unique along North Carolina‘s Outer Banks by being situated on a series of north to south running, parallel, relict beach ridges, which are evidence of the seaward development of the island. These relict dunes and intervening swales provide the framework on which the plant communities have developed. In order to effectively manage and preserve the biodiversity of KHW, comprehensive resource inventory and monitoring were identified as top priority management goals in the Kitty Hawk Woods
Guide to the Vascular Flora of Kitty Hawk Woods
(Dare County, NC)
by
Rachel K. Clark
A thesis submitted to the Graduate Faculty of
North Carolina State University
in partial fulfillment of the
requirements for the degree of
Master of Science
Plant Biology
Raleigh, North Carolina
2011
APPROVED BY:
_______________________________
______________________________
Dr. Jon M. Stucky Dr. H. Joseph Kleiss
________________________________
Dr. Alexander Krings
ii
BIOGRAPHY
Rachel was born in St. Louis, Missouri, where her interest in biology began at the early age of one when her parents adopted an Asian elephant in her name from the St. Louis Zoo. Frequent trips to the zoo to see her elephant helped develop her passion for conservation. Her parents were also members and frequent visitors to the Missouri Botanical Garden, but it was not until college that she really appreciated the time spent
accompanying her parents to the gardens.
She attended Beloit College in Beloit, Wisconsin. It wasn‘t until her junior year that she was able to enroll in an introductory botany course taught by her undergraduate advisor, Yaffa Grossman. It was during this course that Rachel‘s interest in botany began to grow and her interest in conservation expanded to include all the components of natural communities. As part of her coursework, she participated in a six month study abroad program in Tanzania. The program‘s emphasis on conservation biology and ecology meant spending
extended periods of time in Tanzania‘s national parks where she was lucky enough to see animals in the wild she had grown up watching at the zoo. Her independent project evaluating the effectiveness of a wildlife corridor in the Engare Nanyuki region of Tanzania served as the basis for her senior thesis and exposed her for the first time to GIS. She graduated in 2002 with a degree in Evolution, Ecology, and Behavioral Biology.
iii
ACKNOWLEDGEMENTS
I would like to thank everyone who contributed their time or expertise in helping me with this project, I could not have done it alone. I would like to thank my advisor, Dr. Alexander Krings for his patience,
guidance, help, and his enthusiasm for taxonomy and floristic studies. I would also like to thank my committee members, Dr. Joe Kleiss and Dr. Jon Stucky, for their thoughtful suggestions and assistance.
I thank the North Carolina Coastal Reserve system for access to Kitty Hawk Woods and providing collection permits. The reserve manager of Kitty Hawk Woods, Dr. Bo Dame, was an enormous source of encouragement and logistical support. I am indebted to Aaron McCall and The Nature Conservancy for the allowing me the use of the Nags Head Woods reserve house during the collection seasons. I thank Linda Craney, members of the Outer Banks NC Native Plant Society, and the OBX Paddler‘s Club for their gracious offers of lodging and meals. I would also like to thank Inger Seitz for the donation of kayaking equipment. For making my collection trips more enjoyable, I thank the following people who assisted me in the field: Ashley Craft (University of North Carolina-Wilmington), Kathy Mitchell (NC Aquarium-Roanoke) and Dr. Heather McGuire (Chowan University).
I am grateful to the following people who provided assistance with some determinations: Paul Fantz (Liriope), Richard Leblond (Dichanthelium portoricense; Solidago villosicarpa), Dr. Jon Stucky, and Donna Wright. I thank the staff of DUKE and NCU for access to their collections. I am also grateful to Carol Ann McCormick (University of North Carolina-Chapel Hill), Tina Sekula (Louis Berger & Associates), and John Finnegan (NC Natural Heritage Program) for their time and advice.
For being generally outstanding, I would like to thank my fellow students of Drs. Stucky and Krings: Kelly Hines, Amanda Saville, Jenny Stanley, and Robert Thornhill. I would also like to recognize Kelly Hines for her dedication and companionship.
I would like to thank all my friends and parents for their support and encouragement throughout my entire graduate experience. I am also extremely grateful to Paul D‘Agostino for his unwavering emotional,
iv
TABLE OF CONTENTS
LIST OF TABLES ... v
LIST OF FIGURES ... vi
INTRODUCTION ... 1
METHODS ... 1
STUDY AREA ... 2
Setting ... 2
Geomorphology ... 3
Soils ... 4
Climate ... 5
Land Use History ... 5
PLANT COMMUNITIES ... 6
FLORISTIC SUMMARY AND DISCUSSION ... 9
TAXONOMIC TREATMENT ... 12
v
LIST OF TABLES
Table 1. List of North Carolina Natural Heritage Program‘s State Significantly Rare or Endangered Vascular
Plants vouchered or reported from Kitty Hawk Woods ... 177
Table 2. Scale used to estimate frequency of taxa at Kitty Hawk Woods ... 179
Table 3. Summary of the vascular plant groups at Kitty Hawk Woods ... 179
vi
LIST OF FIGURES
Figure 1. Map of northern Outer Banks and location of Kitty Hawk Woods. ... 180
Figure 2. Aerial photograph of Kitty Hawk Woods showing relict beach ridges. ... 181
Figure 3. Soil map units at Kitty Hawk Woods ... 182
Figure 4. Walter climate diagrams for Manteo and Cape Hatteras (Dare County, NC). ... 183
Figure 5. Distribution of species rich families by habit at Kitty Hawk Woods. ... 184
Figure 6. Log transformed species richness versus log transformed area for seven barrier island floras ... 184
Figure 7. Comparison of the distribution of species rich families between Nags Head Woods (NHW) and Kitty Hawk Woods (KHW). ... 185
Figure 8. Asplenium platyneuron ... 186
Figure 9. Azolla caroliniana ... 186
Figure 10. Woodwardia areolata ... 187
Figure 11. Woodwardia virginica ... 187
Figure 12. Pteridium aquilinum ... 188
Figure 13. Botrypus virginanus ... 188
Figure 14. Sceptridium dissectum ... 189
Figure 15. Osmunda regalis var. spectabilis ... 189
Figure 16. Osmundastrum cinnamomeum ... 190
Figure 17. Pleopeltis polypodioides ... 190
Figure 18. Thelypteris palustris var. pubescens ... 191
Figure 19. Juniperus virginiana ... 191
Figure 20. Taxodium distichum ... 192
Figure 21. Pinus echinata ... 192
Figure 22. Pinus taeda ... 193
Figure 23. Yucca filamentosa ... 193
Figure 24. Sagittaria lancifolia ... 194
Figure 25. Sagittaria latifolia ... 194
Figure 26. Narcissus pseudonarcissus ... 195
Figure 27. Nothoscordum bivalve ... 195
Figure 28. Arisaema triphyllum ... 196
Figure 29. Lemna obscura ... 196
Figure 30. Lemna valdiviana ... 197
Figure 31. Peltandra virginica ... 197
vii
Figure 33. Wolffia columbiana ... 198
Figure 34. Wolffiella gladiata ... 199
Figure 35. Sabal minor ... 199
Figure 36. Tillandsia usneoides ... 200
Figure 37. Commelina erecta ... 200
Figure 38. Bolboschoenus robustus ... 201
Figure 39. Carex alata ... 201
Figure 40. Carex comosa ... 202
Figure 41. Carex complanata ... 202
Figure 42. Carex glaucescens ... 203
Figure 43. Carex hyalinolepis ... 203
Figure 44. Carex lupulina ... 204
Figure 45. Carex muehlenbergii ... 204
Figure 46. Carex seorsa ... 205
Figure 47. Carex stipata ... 205
Figure 48. Cladium jamaicense ... 206
Figure 49. Cyperus croceus ... 206
Figure 50. Cyperus echinatus ... 207
Figure 51. Cyperus esculentus ... 207
Figure 52. Cyperus filicinus ... 208
Figure 53. Cyperus grayi ... 208
Figure 54. Cyperus haspan ... 209
Figure 55. Cyperus odoratus ... 209
Figure 56. Cyperus polystachyos ... 210
Figure 57. Cyperus retrofractus ... 210
Figure 58. Cyperus retrorsus ... 211
Figure 59. Cyperus rotundus ... 211
Figure 60. Cyperus strigosus ... 212
Figure 61. Dulichium arundinaceum ... 212
Figure 62. Eleocharis albida ... 213
Figure 63. Eleocharis flavescens ... 213
Figure 64. Eleocharis obtusa ... 214
Figure 65. Eleocharis parvula ... 214
Figure 66. Eleocharis quadrangulata ... 215
viii
Figure 68. Fimbristylis autumnalis ... 216
Figure 69. Fimbristylis castanea ... 216
Figure 70. Rhynchospora colorata ... 217
Figure 71. Rhynchospora corniculata ... 217
Figure 72. Rhynchospora inexpansa ... 218
Figure 73. Rhynchospora scirpoides ... 218
Figure 74. Schoenoplectus americanus ... 219
Figure 75. Schoenoplectus pungens ... 219
Figure 76. Schoenoplectus tabernaemontani ... 220
Figure 77. Scirpus cyperinus ... 220
Figure 78. Eriocaulon decangulare ... 221
Figure 79. Limnobium spongia ... 221
Figure 80. Iris virginica ... 222
Figure 81. Sisyrinchium atlanticum ... 222
Figure 82. Juncus acuminatus ... 223
Figure 83. Juncus coriaceus ... 223
Figure 84. Juncus dichotomus ... 224
Figure 85. Juncus effusus ... 224
Figure 86. Juncus marginatus ... 225
Figure 87. Juncus megacephalus ... 225
Figure 88. Juncus roemerianus ... 226
Figure 89. Juncus scirpoides ... 226
Figure 90. Juncus tenuis ... 227
Figure 91. Triglochin striata ... 227
Figure 92. Listera australis ... 228
Figure 93. Spiranthes praecox ... 228
Figure 94. Spiranthes vernalis ... 229
Figure 95. Tipularia discolor ... 229
Figure 96. Andropogon ternarius ... 230
Figure 97. Andropogon virginicus ... 230
Figure 98. Anthoxanthum odoratum ... 231
Figure 99. Aristida lanosa ... 231
Figure 100. Aristida purpurascens ... 232
Figure 101. Arthraxon hispidus ... 232
ix
Figure 103. Arundo donax ... 233
Figure 104. Axonopus fissifolius ... 234
Figure 105. Briza minor ... 234
Figure 106. Bromus commutatus ... 235
Figure 107. Cenchrus spinifex ... 235
Figure 108. Cenchrus tribuloides ... 236
Figure 109. Chasmanthium laxum ... 236
Figure 110. Chasmanthium sessiliflorum ... 237
Figure 111. Dactylis glomerata ... 237
Figure 112. Danthonia sericea ... 238
Figure 113. Dichanthelium commutatum var. commutatum ... 238
Figure 114. Dichanthelium dichotomum ... 239
Figure 115. Dichanthelium laxiflorum ... 239
Figure 116. Dichanthelium oligosanthes ... 240
Figure 117. Dichanthelium scoparium ... 240
Figure 118. Digitaria ciliaris ... 241
Figure 119. Digitaria ischaemum ... 241
Figure 120. Digitaria sanguinalis ... 242
Figure 121. Digitaria villosa ... 242
Figure 122. Distichlis spicata ... 243
Figure 123. Echinochloa crus-galli ... 243
Figure 124. Echinochloa walteri ... 244
Figure 125. Eleusine indica ... 244
Figure 126. Elymus glabriflorus ... 245
Figure 127. Eragrostis capillaris ... 245
Figure 128. Eragrostis curvula ... 246
Figure 129. Eragrostis spectabilis ... 246
Figure 130. Festuca subverticillatus ... 247
Figure 131. Glyceria septentrionalis ... 247
Figure 132. Leersia oryzoides ... 248
Figure 133. Lolium perenne ... 248
Figure 134. Microstegium vimineum ... 249
Figure 135. Panicum amarum ... 249
Figure 136. Paspalum setaceum ... 250
x
Figure 138. Paspalum vaginatum ... 251
Figure 139. Phragmites australis ... 251
Figure 140. Piptochaetium avenaceum ... 252
Figure 141. Poa annua ... 252
Figure 142. Polypogon monspeliensis ... 253
Figure 143. Schedonorus arundinaceus ... 253
Figure 144. Setaria magna ... 254
Figure 145. Setaria parviflora ... 254
Figure 146. Setaria pumila ... 255
Figure 147. Sorghastrum elliottii ... 255
Figure 148. Spartina cynosuroides ... 256
Figure 149. Spartina patens ... 256
Figure 150. Sphenopholis nitida ... 257
Figure 151. Sphenopholis pensylvanica ... 257
Figure 152. Sporobolus clandestinus ... 258
Figure 153. Tridens flavus ... 258
Figure 154. Triplasis purpurea ... 259
Figure 155. Vulpia octoflora ... 259
Figure 156. Zizania aquatic ... 260
Figure 157. Pontederia cordata ... 260
Figure 158. Stuckenia pectinata ... 261
Figure 159. Ruppia maritima ... 261
Figure 160. Polygonatum biflorum ... 262
Figure 161. Smilax bona-nox ... 262
Figure 162. Smilax glauca ... 263
Figure 163. Smilax laurifolia ... 263
Figure 164. Smilax rotundifolia ... 264
Figure 165. Smilax smallii ... 264
Figure 166. Sparganium americanum ... 265
Figure 167. Typha angustifolia ... 265
Figure 168. Typha latifolia ... 266
Figure 169. Liquidambar styraciflua ... 266
Figure 170. Alternanthera philoxeroides ... 267
Figure 171. Amaranthus cannabinus ... 267
xi
Figure 173. Toxicodendron radicans ... 268
Figure 174. Asimina parviflora ... 269
Figure 175. Centella erecta ... 269
Figure 176. Cicuta maculata ... 270
Figure 177. Lilaeopsis chinensis ... 270
Figure 178. Ptilimnium capillaceum ... 271
Figure 179. Sanicula canadensis ... 271
Figure 180. Sium suave ... 272
Figure 181. Spermolepis divaricata ... 272
Figure 182. Asclepias incarnata ... 273
Figure 183. Asclepias lanceolata ... 273
Figure 184. Asclepias tuberosa ... 274
Figure 185. Asclepias variegata ... 274
Figure 186. Ilex opaca ... 275
Figure 187. Ilex vomitoria ... 275
Figure 188. Aralia spinosa ... 276
Figure 189. Hydrocotyle ranunculoides ... 276
Figure 190. Hydrocotyle umbellata ... 277
Figure 191. Hydrocotyle verticillata ... 277
Figure 192. Acanthospermum australe ... 278
Figure 193. Achillea millefolium ... 278
Figure 194. Ambrosia artemisiifolia ... 279
Figure 195. Baccharis halimifolia ... 279
Figure 196. Bidens bipinnata ... 280
Figure 197. Bidens laevis ... 280
Figure 198. Chrysopsis gossypina ... 281
Figure 199. Chrysopsis mariana ... 281
Figure 200. Cirsium horridulum ... 282
Figure 201. Conyza canadensis ... 282
Figure 202. Elephantopus nudatus ... 283
Figure 203. Elephantopus tomentosus ... 283
Figure 204. Erigeron annuus ... 284
Figure 205. Eupatorium capillifolium ... 284
Figure 206. Eupatorium serotinum ... 285
xii
Figure 208. Heterotheca subaxillaris ... 286
Figure 209. Hieracium gronovii ... 286
Figure 210. Hypochaeris radicata ... 287
Figure 211. Iva frutescens ... 287
Figure 212. Krigia virginica ... 288
Figure 213. Lactuca canadensis ... 288
Figure 214. Liatris pilosa ... 289
Figure 215. Mikania scandens ... 289
Figure 216. Pityopsis graminifolia ... 290
Figure 217. Pluchea foetida ... 290
Figure 218. Pluchea odorata ... 291
Figure 219. Prenanthes serpentaria ... 291
Figure 220. Pyrrhopappus carolinianus ... 292
Figure 221. Solidago altissima ... 292
Figure 222. Solidago erecta ... 293
Figure 223. Solidago fistulosa ... 293
Figure 224. Solidago odora ... 294
Figure 225. Solidago sempervirens ... 294
Figure 226. Sonchus asper ... 295
Figure 227. Symphyotrichum dumosum ... 295
Figure 228. Symphyotrichum patens ... 296
Figure 229. Symphyotrichumnovi-belgii ... 296
Figure 230. Symphyotrichum undulatum ... 297
Figure 231. Taraxacum officinale ... 297
Figure 232. Verbesina occidentalis ... 298
Figure 233. Carpinus caroliniana ... 298
Figure 234. Ostrya virginiana ... 299
Figure 235. Bignonia capreolata ... 299
Figure 236. Campsis radicans ... 300
Figure 237. Cardamine hirsuta ... 300
Figure 238. Cardamine parviflora var. arenicola ... 301
Figure 239. Cardamine pensylvanica ... 301
Figure 240. Lepidium virginicum ... 302
Figure 241. Triodanis perfoliata ... 302
xiii
Figure 243. Lonicera sempervirens ... 303
Figure 244. Cerastium fontanum ... 304
Figure 245. Cerastium glomeratum ... 304
Figure 246. Scleranthus annuus ... 305
Figure 247. Silene antirrhina ... 305
Figure 248. Silene caroliniana var. pensylvanica ... 306
Figure 249. Stellaria media ... 306
Figure 250. Euonymus americanus ... 307
Figure 251. Crocanthemum canadense ... 307
Figure 252. Hudsonia tomentosa ... 308
Figure 253. Lechea maritima var. virginica ... 308
Figure 254. Lechea mucronata ... 309
Figure 255. Calystegia sepium ... 309
Figure 256. Ipomoea pandurata ... 310
Figure 257. Cornus florida ... 310
Figure 258. Cornus stricta ... 311
Figure 259. Melothria pendula ... 311
Figure 260. Drosera intermedia ... 312
Figure 261. Diospyros virginiana ... 312
Figure 262. Chimaphila maculata ... 313
Figure 263. Gaylussacia frondosa ... 313
Figure 264. Lyonia ligustrina ... 314
Figure 265. Lyonia lucida ... 314
Figure 266. Monotropa uniflora ... 315
Figure 267. Oxydendrum arboreum ... 315
Figure 268. Vaccinium fuscatum ... 316
Figure 269. Vaccinium pallidum ... 316
Figure 270. Acalypha gracilens ... 317
Figure 271. Cnidoscolus stimulosus ... 317
Figure 272. Euphorbia polygonifolia ... 318
Figure 273. Albizia julibrissin ... 318
Figure 274. Apios americana ... 319
Figure 275. Centrosema virginianum ... 319
Figure 276. Cercis canadensis ... 320
xiv
Figure 278. Clitoria mariana ... 321
Figure 279. Crotalaria rotundifolia ... 321
Figure 280. Desmodium glabellum ... 322
Figure 281. Desmodium paniculatum ... 322
Figure 282. Galactia volubilis ... 323
Figure 283. Hylodesmum nudiflorum ... 323
Figure 284. Lespedeza stuevei ... 324
Figure 285. Lotus corniculatus ... 324
Figure 286. Melilotus officinalis ... 325
Figure 287. Rhynchosia difformis ... 325
Figure 288. Robinia pseudo-acacia ... 326
Figure 289. Strophostyles helvola ... 326
Figure 290. Trifolium arvense ... 327
Figure 291. Trifolium campestre ... 327
Figure 292. Trifolium pratense ... 328
Figure 293. Trifolium repens ... 328
Figure 294. Vicia sativa ssp. nigra ... 329
Figure 295. Castanea pumila ... 329
Figure 296. Fagus grandifolia ... 330
Figure 297. Quercus alba ... 330
Figure 298. Quercus falcata ... 331
Figure 299. Quercus laurifolia ... 331
Figure 300. Quercus michauxii ... 332
Figure 301. Quercus nigra ... 332
Figure 302. Quercus virginiana ... 333
Figure 303. Corydalis halei ... 333
Figure 304. Gelsemium sempervirens ... 334
Figure 305. Sabatia calycina ... 334
Figure 306. Sabatia dodecandra ... 335
Figure 307. Geranium carolinianum ... 335
Figure 308. Myriophyllumheterophyllum ... 336
Figure 309. Myriophyllum spicatum ... 336
Figure 310. Proserpinaca palustris ... 337
Figure 311. Hamamelis virginiana ... 337
xv
Figure 313. Hypericum hypericoides ... 338
Figure 314. Hypericum mutilum ... 339
Figure 315. Triadenum walteri ... 339
Figure 316. Itea virginica ... 340
Figure 317. Carya glabra ... 340
Figure 318. Carya pallida ... 341
Figure 319. Callicarpa americana ... 341
Figure 320. Monarda punctata ... 342
Figure 321. Teucrium canadense ... 342
Figure 322. Trichostema dichotomum ... 343
Figure 323. Persea palustris ... 343
Figure 324. Sassafras albidum ... 344
Figure 325. Utricularia gibba ... 344
Figure 326. Mitreola petiolata ... 345
Figure 327. Decodon verticillatus ... 345
Figure 328. Lythrum lineare ... 346
Figure 329. Magnolia virginiana ... 346
Figure 330. Hibiscus moscheutos ... 347
Figure 331. Kosteletzkya virginica ... 347
Figure 332. Mollugo verticillata ... 348
Figure 333. Morus rubra ... 348
Figure 334. Morella cerifera ... 349
Figure 335. Nelumbo lutea ... 349
Figure 336. Nyssa biflora ... 350
Figure 337. Nyssa sylvatica ... 350
Figure 338. Fraxinus caroliniana ... 351
Figure 339. Fraxinus pennsylvanica ... 351
Figure 340. Fraxinus profunda ... 352
Figure 341. Ludwigia brevipes ... 352
Figure 342. Ludwigia decurrens ... 353
Figure 343. Ludwigia repens ... 353
Figure 344. Oenothera biennis ... 354
Figure 345. Oenothera laciniata ... 354
Figure 346. Agalinis purpurea ... 355
xvi
Figure 348. Conopholis americana ... 356
Figure 349. Epifagus virginiana ... 356
Figure 350. Oxalis dillenii ... 357
Figure 351. Passiflora lutea ... 357
Figure 352. Bacopa monnieri ... 358
Figure 353. Nuttallanthus canadensis ... 358
Figure 354. Plantago lanceolata ... 359
Figure 355. Plantago major ... 359
Figure 356. Plantago virginica ... 360
Figure 357. Veronica officinalis ... 360
Figure 358. Persicaria hydropiper ... 361
Figure 359. Persicaria hydropiperoides ... 361
Figure 360. Persicaria pensylvanica ... 362
Figure 361. Persicaria punctata ... 362
Figure 362. Persicaria setacea ... 363
Figure 363. Rumex acetosella ... 363
Figure 364. Rumex conglomeratus ... 364
Figure 365. Rumex crispus ... 364
Figure 366. Rumex floridanus ... 365
Figure 367. Rumex hastatulus ... 365
Figure 368. Hottonia inflata ... 366
Figure 369. Samolus parviflorus ... 366
Figure 370. Ranunculus bulbosus ... 367
Figure 371. Ranunculus sardous ... 367
Figure 372. Ranunculus sceleratus ... 368
Figure 373. Berchemia scandens ... 368
Figure 374. Amelanchier canadensis ... 369
Figure 375. Aronia arbutifolia ... 369
Figure 376. Geum canadense ... 370
Figure 377. Potentilla indica ... 370
Figure 378. Prunus angustifolia ... 371
Figure 379. Rosa palustris ... 371
Figure 380. Rubus trivialis ... 372
Figure 381. Cephalanthus occidentalis ... 372
xvii
Figure 383. Diodia virginiana ... 373
Figure 384. Galium circaezans ... 374
Figure 385. Galium obtusum ... 374
Figure 386. Galium tinctorium ... 375
Figure 387. Houstonia pusilla ... 375
Figure 388. Mitchella repens ... 376
Figure 389. Zanthoxylum clava-herculis ... 376
Figure 390. Salix caroliniana ... 377
Figure 391. Salix sericea ... 377
Figure 392. Acer rubrum ... 378
Figure 393. Saururus cernuus ... 378
Figure 394. Verbascum thapsus ... 379
Figure 395. Solanum carolinense ... 379
Figure 396. Symplocos tinctoria ... 380
Figure 397. Polypremum procumbens ... 380
Figure 398. Ulmus americana ... 381
Figure 399. Boehmeria cylindrica ... 381
Figure 400. Valerianella radiata ... 382
Figure 401. Phyla lanceolata ... 382
Figure 402. Phyla nodiflora ... 383
Figure 403. Viola lanceolata ... 383
Figure 404. Viola sororia ... 384
Figure 405. Phoradendron serotinum ... 384
Figure 406. Ampelopsis arborea ... 385
Figure 407. Parthenocissus quinquefolia ... 385
Figure 408. Vitis aestivalis ... 386
1 INTRODUCTION
Kitty Hawk Woods (KHW) is one of ten reserves comprising the North Carolina Coastal Reserves system. Its unique geography and plant communities make it an interesting component of North Carolina‘s coastal
fringe. Located on Bodie Island along the Outer Banks, KHW is situated on a series of parallel, north-south running, relict beach dunes. The reserve is comprised of 760 ha of forested wetlands, sandy wooded ridges, and brackish marshes, bordered on the west by Currituck Sound and located a mile from the Atlantic Ocean on the east (Fig. 1). Designated as a national significant natural area (NCNHP 2009), KHW hosts several globally rare plant communities: Maritime Deciduous Forest (S1 G1), Maritime Swamp Forest (S1S2 G2), and Interdunal Ponds (S1S2 G2?; Schafale & Weakley 1990). Maritime Swamp Forest dominated by Taxodium distichum (L.) Rich. occurs in only two locations on the Outer Banks and is developed to its greatest extent at KHW
(Wentworth et al. 1992). Maritime Deciduous Forest occurs in only two additional localities in North Carolina: Nags Head Woods and Southern Shores Cypress Swamp (Schafale & Weakley 1990). Eight plant species documented from KHW are state listed as Significantly Rareand fifteen more are on the NC Natural Heritage Program‘s Rare Plant Watch List (Buchanan & Finnegan 2010; Table 1). The forested wetlands and brackish marshes serve as critical wildlife habitat for a variety of amphibians, reptiles and birds, including the bald eagle (Haliaeetus leucocephalus L.) (B. Dame, pers. comm.).
In the last twenty years, Dare County has seen a 60% increase in population (U.S. Census Bureau) and mounting development pressure on its natural areas. In order to effectively manage and preserve the biodiversity of KHW, comprehensive resource inventory and monitoring were identified as top priority
management goals in the Kitty Hawk Woods management plan (KHW Mgmt Plan 1998). The objectives of the present work were to contribute to these goals by developing: (1) a checklist of the vascular flora of Kitty Hawk Woods based on a thorough floristic inventory and survey of historic collections and (2) an illustrated guide based on the checklist to facilitate monitoring, research, and precision management.
METHODS
Checklist. An initial species list was compiled of all plant taxa reported from the Outer Banks by Burk (1961) and Stalter and Lamont (1997), as well as those mapped for Dare County by the SE Flora Atlas (NCU). This list was used to search the collections at Duke University (DUKE), North Carolina State University (NCSC), and University of North Carolina–Chapel Hill (NCU) for historic voucher specimens from KHW and Bodie Island.
2 Nomenclature and taxonomic concepts generally follow the cited Flora of North America treatments unless they were not yet available, in which case Weakley (2010) and Radford et al. (1968) were followed. For
Galactia P. Browne nomenclature and taxonomic concepts follow Wilbur (1963). Plant community types were identified using Schafale and Weakley (1990).
Keys. The keys were developed based on a checklist compiled from the author‘s collections, vouchers of historical collections, and literature reports from Burk (1961) and Stalter and Lamont (1997). Species excluded from this list were ones known exclusively from habitats not present at KHW, such as foredunes. Additional keys were developed to species not exclusively reported or vouchered from KHW when common congeners both occurred on Bodie Island and their presence would be expected in the reserve. The most conservative approach was taken for Cyperus L., Eleocharis R. Br., Fimbristylis Vahl, and Rhynchospora Vahl for which all recorded taxa from Bodie Island were included in the keys. When available, keys were primarily adapted from the cited treatments in the Flora of North America, or secondarily Weakley (2010) and Radford et al. (1968).
Format. For each taxon, basionym, synonymy in Radford et al. (1968), habitat, reproductive phenology, and abundance are provided. Abundance was estimated following Palmer et al. (1995; Table 2). Taxa are designated exotic if they are not native to North America. State listed rare or watch list taxa are noted. Relevant vouchers from Kitty Hawk and then Bodie Island and nearby Colington Island are cited. The following symbols after the specimen vouchers indicate literature reports of the species specifically from Kitty Hawk Woods: ♣ (Burk 1961) and ♦ (Stalter & Lamont 1997).
Illustrations were taken from the following public domain works: Britton and Brown (1913), Hitchcock (1950), and USDA NRCS (2010).
STUDY AREA
Setting. Kitty Hawk Woods is located on Bodie Island, off the coast of Dare County, NC. The reserve is composed of several non-contiguous tracts, which are bordered by Currituck Sound on the west and urban development on the east (Fig. 2). The reserve is bounded by US 158 and Colleton Ave to the north and Kitty Hawk Bay to the south. Located south of Kitty Hawk are the Kitty Hawk Bay Islands comprising ca. 174 ha (23%) of the total reserve. The islands have remained undeveloped, Juncus roemerianus Scheele-dominated brackish marsh. Bisecting the western portion of the reserve is Jean Guite Creek, which flows north to south and stretches from Kitty Hawk Bay to Currituck Sound. Two major roads and associated housing developments separate the remaining components of the reserve.
Due to Jean Guite Creek‘s connection to the sound, the salinity in the creek and marsh ranges 0–5 ppt
3 In between the ridges of KHW are numerous freshwater swamps and ponds, which can range in depth from zero to two feet depending on climatic conditions. The ionic concentrations of these ponds and swamps are slightly higher than inland wetlands due to the proximity of salt spray (Odum 1988). This supports an abundance of freshwater vegetation which builds up on the bottom of the ponds. Odum (1988) suggested that ponds do not eventually fill in because of the periodic drying and exposure to oxygen which accelerates decomposition.
Geomorphology. The formation of the Outer Banks is relatively recent in geologic time, occurring after sea levels rose at the end of the last glaciation of the Pleistocene (12,000 B.P.). North Carolina‘s Coastal Plain
represents the accumulation of fluvial deposited sediments from the weathering of the Appalachians and Piedmont as well as marine deposited sediments from the higher sea level (Stuckey 1965). These mostly unconsolidated sediments reach depths of several thousand feet at Cape Hatteras. After the last glaciation, sea levels initially rose rapidly and coastal sediments were left largely untouched until sea level rise slowed. The slowing of the sea level allowed for the introduction of sediments from the land to the already sediment loaded ocean (Davis 1994). This ample supply of unconsolidated sediments serves as the basis for the formation of barrier islands.
In general, barrier islands are formed from the deposition of large amounts of sediments and are dynamic systems as sediments are deposited and eroded by wave and wind actions (Davidson–Arnott 2010). Over 80% of the barrier islands that form North Carolina‘s Outer Banks are landward migrating, as sediment is eroded
from the seaward side and washed over to the landward side. The sediments from the mainland across the estuaries also accumulate on the landward side of the barrier island, contributing to its slow landward movement (Moslow & Heron 1994).
The portion of Bodie Island that KHW occupies however is actually a seaward migrating barrier island. The morphological characteristic of these islands is a series of shore-parallel beach ridges that document past shorelines. Only Bogue Banks, and portions of Bodie and Hatteras Islands exhibit this type of barrier island growth (Moslow & Heron 1994). Only at Kitty Hawk do the ridges run north to south; at the other two sites, the ridges run east to west. Abundant sediment supply results in the formation of a dune, behind which forms a low-lying area close to the water table, commonly called a swale. Beach grasses and herbs colonize and stablilize the foredune and eventually maritime forests develop on these stable relict ridges (Davidson-Arnott 2010). With a high sediment supply, successive dunes are formed seaward, thus the oldest ridges are the ones closest to land.
4 periods of time in which the shoreline advanced rapidly without the formation of dunes/beach ridges. The swamps and creek were portions of the beach that was left behind when the shoreline progressed.
The islands in Kitty Hawk Bay are the result of sediment deposition at a past inlet south of the town of Kitty Hawk, where Colington Island is now. The islands were formed as sediment flowing from the Atlantic Ocean through the relict inlet was deposited, much like deltas at the mouth of rivers (Fisher 1967).
Soils. Reflecting the relative recent formation of the barrier islands, the dominant soils at KHW are
Entisols—soils predominantly composed of sand with minimal profile development. In low-lying areas that are saturated at least seasonally, there is an accumulation of organic material and the development of Inceptisols and Histosols (Fig. 3). The soils are generally acidic.
Histosols
Currituck (CuA) mucky peat (sandy or sandy skeletal, mixed, euic thermic Terric Haplosaprists)
Very poorly drained soils on nearly level surfaces. The organic layer has a depth to 100 cm on top of sand, which exceeds depths of 165 cm. The soil is frequently flooded with a seasonal high water table ranging between 30 cm above to 30 cm below the surface (Tant 1992). This is the soil map unit of the Kitty Hawk Bay Islands and the brackish marshes of the southern portion of the reserve.
Inceptisols
Conaby (CnA) muck (coarse-loamy, mixed, non-acid, thermic Histic Humaquepts)
Very poorly drained soils on nearly level surfaces. The organic layer of muck is 35 cm thick, overlying sand which exceeds 150 cm in depth. The seasonal high water table is at or near the surface. These soils are in the swales and low flats between relict beach ridges (Tant 1992). They are more common in the areas along Jean Guite Creek and the portion of the reserve on the east side of Woods Rd.
Entisols
Fripp (FrD) fine sand (Thermic, uncoated Typic Quartzipsamments)
Excessively drained soils on 2–30% slopes. The soil texture is sand to at least a depth of 200 cm. This soil makes up the relict beach ridges and is generally protected from salt spray (Tant 1992). This soil map unit forms the ridges in the portion of the reserve north of Twiford Rd.
Osier (OsA) fine sand (Siliceous, thermic Typic Psammaquents)
5 is found in swales and shallow interdunal ponds in the western portion of the reserve, near the boundary at Currituck Sound.
Ousley (OuB) fine sand (Thermic, uncoated Aquic Quartzipsamments)
Moderately well drained soils with 0–5% slopes. The soil texture is sand to a depth of at least 200 cm and the seasonal high water table is 45 to 92 cm below the surface, areas only subjected to flooding in extreme climatic events (Tant 1992). These are the soils of low dunes along the sound and are found along the ridges extending through the marshes in the southern portion of the reserve.
Psamments (Psb)
These are found in areas where the natural soils have been dredged or disturbed. In order to build housing along low-lying areas, fill is used to elevate the surface layer to prepare it for more intensive use. These soils are typically very sandy and soil properties vary depending on the origin of the fill material (Tant 1992). They are moderately well drained soils, only rarely flooded and found near the housing development at the very southern of the reserve.
Climate. The climate at KHW is warm temperate, with hot summers and adequate rainfall throughout the year (Peel et al. 2007). In general, the Outer Banks experience milder weather than points further inland, with less extremes in temperature (Burk 1961; Stalter & Lamont 1997). The nearest weather station to Kitty Hawk is in Kill Devil Hills (36°01‘82‖ N, 75°67‘13‖ W) at four meters above sea level, however this station has only
been operational from January 2004 to the present. The mean annual precipitation from 2004–2010 was 694 mm, reflecting the periodic droughts over this period. The next closest weather station with observations from the last forty-four years is in Manteo (35°91‘72‖ N, 75°7‘0‖ W) on Roanoke Island, 4 meters above sea level. Average annual temperature over the last forty-four years recorded there was 16.7°C,with mean annual precipitation of 1299 mm. Average daily maximum temperature is 21.21°C and average daily minimum temperature is 15.51°C (Fig. 4). Typically, there is precipitation received every month, with a peak in the summer in August. Winds generally blow from the north or northeast from September to March and from the south or southwest from April to August. Cape Hatteras to the south experiences similar climatic conditions, but with higher mean annual precipitation (1448 mm). Bodie Island has a growing season between 240–300 days annually with average annual amount of sunlight between 2400–2800 hours (Krings 2010).
Land Use History. The following description of the history of KHW was adapted from the KHW Management Plan (1998).
6 today was part of a much larger forested system that covered Colington Island to the south and Southern Shores to the north. Native Americans lived on Bodie Island and likely used the woods for hunting and the gathering of plants prior to the arrival of European colonialists. There were English settlements in the Kitty Hawk area but no evidence of residences within the woods themselves. Mature trees were harvested for shipbuilding but serious logging operations did not begin until 1910 and continued until the early 1930s.
The area remained undisturbed until the late 1980s, when a housing developer, who held 566 ha of land within KHW made plans for the residential and commercial development of the woods. In 1992, the developer exchanged the land to the state for permits to fill wetlands located elsewhere. The town of Kitty Hawk also recognized the importance of the woods and purchased the 186 ha tract east of Woods Road. Further land acquisition has been ongoing as State funds are available, a process contributing to the piecemeal nature of the reserve. The area east of Woods Rd has suffered very little intrusion and remains one of the best examples of Maritime Swamp Forest on the Outer Banks (Lopazanski et al. 1998).
PLANT COMMUNITIES
The ecological importance of KHW lies in the diversity of plant communities it encompasses. Six different natural plant communities can be described following the concepts of Schafale and Weakley (1990), in addition to one anthropogenic community along a powerline right-of-way. However, drawing boundaries around plant communities in KHW is difficult due to the complex mosiac of closely spaced wetland swales and upland ridges. Species composition appears to be less affected by differences along topographic moisture gradients than sites found elsewhere on the Outer Banks or inland (Schafale & Weakley 1990). Species typically found in wet or saturated soils, such as Persea palustris (Raf.) Sarg. (FACW) and Quercus michauxii Nutt. (FACW-) are present on the sandy ridges at KHW. One explanation for this could be the generally low nature of the beach ridges in KHW, on average rising no more than three meters above the water table. Even growing on the tops of these ridges, plants with extensive roots systems are not far from adequate moisture. In contrast, sites that have more pronounced elevational differences between ridges and swales would be expected to exhibit more distinct segregation between wetland and upland species.
Maritime Deciduous Forest. This is a globally rare plant community (S1 G1) found along the well-protected relict beach ridges of KHW. The predominant soil series is Fripp (Typic Quartzipsamments). The species assemblages of this community are very similar to mesic deciduous forests found inland, with species not generally associated with harsh maritime conditions being common, such as Fagus grandiflora Ehrh.,
Conopholis americana (L.) Wallr., or Botrypus virginianus (L.) Michx.
The canopy is dominated by a mixture of Pinus taeda L. and deciduous hardwoods such as Carya glabra
(Mill.) Sweet, Fagusgrandifolia, Juniperus virginiana L., Liquidambar styraciflua L., Nyssa sylvatica
7 Walter, Cornus florida L., Ilex opaca Aiton, Ostrya virginiana (Mill.) K. Koch, Oxydendrum arboreum (L.) DC., Persea palustris, and Sassafras albidum (Nutt.) Nees. Within this community are pockets of Quercus laurifolia Michx., Q. michauxii, and Symplocos tinctoria (L.) L‘Her. These pockets are only found in the western part of the reserve and are associated with the ecotone between ridges and swales. The shrub layer consists of Callicarpa americana L., Gaylussacia frondosa (L.) Torr. & A. Gray, Hamamelis virginiana L.,
Hypericum hypericoides (L.) Crantz, Morella cerifera (L.) Small, and Vaccinium pallidum Aiton. Common vines include Bignonia capreolata L., Lonicera sempervirens L., Parthenocissus quinquefolia (L.) Planch.,
Toxicodendron radicans (L.) Kuntze, Vitis aestivalis Michx., V. rotundifolia Michx., Smilax bona-nox L., and S. smallii Morong. A common subshrub or creeping vine is Mitchella repens L. The herb layer consists of
Chasmanthium laxum (L.) Yates, Desmodium Desv. spp., Dichanthelium (Hitchc. & Chase) Gould spp., Galium hispidulum Michx., Elephantopusnudatus A. Gray, E. tomentosus L., Pteridium aquilinum (L.) Kuhn, and
Tipularia discolor (Pursh) Nutt. Solidago L. spp. and Symphyotrichum Nees spp. are associated with clearings within this community type.
Maritime Evergreen Forest. This state rare community type (S1 G2G3) differs from Maritime Deciduous Forest by the lack of deciduous hardwoods in the canopy, consisting only of combinations of Pinus taeda,
Quercus hemisphaerica Bartr. ex Willd. and Q. virginiana Mill. Associated soil series are Fripp (Typic Quartzipsamment) and Ousley (Aquic Quartzipsamments). In KHW, this community type is predominately found along low ridges, particularly on the ridges that extend southward and are surrounded by brackish marsh. Understory trees include Cornus florida, Ilex opaca, Juniperusvirginiana, Ostrya virginiana, Persea palustris, and Sassafras albidum. Shrubs include Callicarpa americana, Hypericum hypericoides, Ilex vomitoria Aiton, and Morella cerifera, Typical vines include Berchemia scandens (Hill) K. Koch, Bignonia capreolata,
Parthenocissus quinquefolia, Smilax bona-nox, S. glauca Walter, Toxicodendron radicans, and Vitis rotundifolia. Mitchella repens is the predominant forest floor covering. The herb layer consists of
Dichanthelium spp., Elephantopusnudatus, E. tomentosus, and Clitoria mariana L.
Maritime Swamp Forest. This community type is globally rare (S1 G1) and the KHW portion represents the most extensive and well preserved in the state (Lopazanski et al. 1988). The community is typically associated with mucky soils of the Conaby series (Histic Humaquept), in which almost permanent saturation leads to the development of a thick organic layer. The canopy is dominated by Acer rubrum L., Liquidambar styracifllua,
Nyssa biflora Walter, Taxodium distichum, and Ulmus americana L. Common understory trees and shrubs include Carpinus caroliniana, Fraxinus pennsylvanica Marshall, Lyonia lucida (Lam.) K. Koch, Persea palustris, Quercus nigra, and Vaccinium fuscatum Aiton. Woody vines include Berchemia scandens, Smilax
8 L.spp., Glyceria septentrionalis Hitchc., Osmundaregalis L. var. spectabilis (Willd.) A. Gray, Osmundastrum cinnamomeum (L.) C. Presl, Saururus cernuus L., Thelypteris palustris Schott, Woodwardia aerolata (L.) Moore, and W. virginica (L.) Sm.
Maritime Shrub Swamp (Willow Subtype). In contrast to Maritime Swamp Forest, Maritime Shrub Swamp (S1 G1) is characterized by the dominance of shrubs or small trees. Originally only two variants, Dogwood and Red Bay, were recognized for this community type with examples found only in Buxton Woods and Nags Head Woods (Schafale & Weakley 1990). While not included in the 3rd Approximation, the Willow Subtype is recognized in the 4th Approximation (Schafale, in prep.) as being dominated by Salix caroliniana Michx. instead of Perseapalustris or Cornus stricta Lam. The associated soil series is the saturated mucky Conaby (Histic Humaquept). This community type forms ―islands‖ within larger interdunal swales and may represent a
successional stage between Interdunal Ponds and Maritime Swamp Forest (Schafale & Weakley 1990). Other shrubs and small trees may include Acer rubrum, Cephalanthus occidentalis L., Cornus stricta,and Morella cerifera. Common vines include Berchemia scandens and Smilax spp. The understory consists of Carex spp.,
Sagittaria lancifolia L., S. latifolia Willd., Saururus cernuus, Thelypteris palustris, Woodwardiaareolata,and
W. virginica.
Interdune Ponds. This state rare community type (S1S2 G2?) occurs in the deeper swales and depression between beach ridges. The soil series associated with the interdunal ponds are Conaby (Histic Humaquept) and Osier (Aquic Quartzipsamment). Ponds at KHW range from deep with open water to shallow and mostly covered with vegetation. Pond margins typically exhibit a variety of shrubs and small trees, including Acer rubrum, Cephalanthusoccidentalis, Cornus stricta, Morella cerifera, Rosa palustris Marshall, Salix
caroliniana, Symplocos tinctoria and Vaccinium fuscatum. Shallow ponds frequently may be dominated by a single taxon, such as Carex hyalinolepis Steud., Decodon verticillatus (L.) Elliott, Saururus cernuus,
Sparganium americanum Nutt., or Typha spp. Intrusions of Alternanthera philoxeroides (Mart.) Griseb. and
Phragmites australis (Cav.) Trin. ex Steud. are starting to take over some ponds. Common herbs include
Boehmeria cylindrica (L.) Sw., Carex comosa Boott, Echinochloa crus-galli (L.) P. Beauv., Ludwigia spp.,
Peltandravirginica (L.) Schott, Persicaria spp., and Triadenum walteri (J.F. Gmel.) Gleason. Floating or submerged herbs include Ceratophyllumechinatum A. Gray, Hydrocotyle ranunculoides L. f., Limnobium spongia (Bosc) Rich. ex. Steud., and Myriophyllum heterophyllum Michx.
9
roemerianus, with Cladium jamaicense Crantz, Spartina cynosuroides (L.) Roth and Spartina patens (Aiton) Muhl. found along edges. There may be inclusions of Phragmites australis.
Along the edges or scattered within the marsh occur Amaranthus cannabinus (L.) J.D. Sauer, Apios americana Medik., Asclepias lanceolata Walter, Baccharis halimifolia L., Bolboschoenus robustus (Pursh) Sojáak, Calystegiasepium (L.) R. Br., Cyperus spp., Distichlis spicata (L.) Green, Fimbristyliscaroliniana
(Lam.) Fernald, Fimbristylis castanea (Michx.) Vahl, Hibiscus moscheutos L., Ipomoea sagittata Poir., Iva frutescens L., Kosteletzkya virginica (L.) C. Presl, Lythrumlineare L., Persicaria spp., Pluchea odorata (L.) Cass., Phyla lanceolata (Michx.) Greene, Sabatia dodecandra (L.) Britton, Sterns & Poggenb., Sagittaria lancifolia, Samolusparviflorus Raf., Schoenoplectus spp., and Teucrium canadense L.
Powerline Right-of-Way. The eastern boundary of the reserve is marked by a powerline right-of-way that runs north to south along a relict beach dune ridge. The associated soil series is Fripp (Typic
Quartzipsamment). Many areas along the powerline right-of-way are sparse in vegetation and are simply unconsolidated sands. The absence of a canopy coupled with a high disturbance regime creates an environment favoring ruderal and weedy species. The highest percentage of exotics is found along this cut. There are extensive vine tangles of Smilax spp. and Vitisrotundifolia. Common herb species include Achillea millefolium
L., Andropogon L. spp., Cenchrusspinifex Cav., Cenchrus tribuloides L., Conyza canadensis (L.) Cronquist,
Clitoria mariana, Digitaria Haller spp., Elymus glabriflorus (Vasey) Scribn. & C.R. Ball, Eragrostis Wolf spp.,
Eupatorium capillifolium (Lam.) Small, Lechea mucronata Raf., Nuttallanthus canadensis (L.) D.A. Sutton,
Monarda punctata L., Pityopsis graminifolia (Michx.) Nutt., Pteridium aquilinum, Rubusdiscolor Weihe & Nees, Rubus trivialis Michx., Solidago odora Aiton, and Vulpia C.C. Gmel. spp. Hudsonia tomentosa Nutt. (S1 G4), Lechea maritima Legg. ex Britton, Sterns & Poggenb var. virginica Hodgon (S1 G5T3Q) and Solidago villosicarpa (S1 G1) are also present in this community. Some of the wetter portions of the powerline right-of-way support Carex longii Mack., Cyperus retrofractus (L.) Torr., Dichanthelium scoparium (Lam.) Gould,
Eupatorium pilosum Walter, Juncus effusus L., Rhexia nashii Small, and Scirpus cyperinus (L.) Kunth. During the end of the season in 2010, the powerline right-of-way was significantly widened, cutting into more of the woods and exposing more wet depressions. Future seasons could see a change in the species composition.
FLORISTIC SUMMARY AND DISCUSSION
10 habit is herbs (323 spp.), followed by trees and shrubs (71 spp.), and vines (35 spp.). The largest families of herbs are the Poaceae (63 spp.), Asteraceae (45 spp.), Cyperaceae (32 spp.), Fabaceae (13 spp.) and
Polygonaceae (10 spp.). The Fagaceae is the largest family of trees (9 spp.), followed by the Ericaceae (6 spp.) and Rosaceae (4 spp.). The three largest families of vines are Fabaceae (9 spp.), Smilacaceae (5 spp.), and Vitaceae (4 spp.; Fig. 5).
Species-area analysis was used to compare species richness of KHW to that of six other sites along the Outer Banks (Au 1969; Dickerson 1978; Hill 1986; Stalter & Lamont 1999; Kelly 2006; Krings 2010; Fig. 6). There was a significant relationship in the regression of log10 species richness on log10 area (p < 0.01, r2 = 0.46).
Likely due to its diversity in natural communities, KHW is species rich for its size, especially when compared to other sites from the coastal fringe, which lack the diversity of habitats. Most barrier islands share two similar plant communities–the dune grass communities on the seaward side and the brackish to saltwater marsh communities on the sound side of the island. Only islands that are wide enough or have dunes high enough to protect plants from salt-spray develop shrub or forest communities (Burk 1961). Islands also must have relict beach ridges and swales to develop freshwater interdunal ponds. These conditions are met only in a few sites along the Outer Banks (Nags Head Woods, Kitty Hawk Woods, and Buxton Woods) and provide a possible explanation for the species richness found at Nags Head Woods and Kitty Hawk Woods.
Both Nags Head Woods (NHW) and KHW contain similar plant communities, yet NHW hosts a higher number of species (559 spp.) over a smaller area. However, the proportion of species rich families in the different habit classes is very similar between the two reserves (Fig. 7). Both sites exhibit the same top three families of trees and shrubs and vines, and essentially the same top five families of herbs. A noticeable exception to the later are the Orchidaceae, which are represented by fourteen species at NHW, but only four at KHW. It remains unclear whether this is a sampling artifact or a natural phenomenon. Lamont and Stalter (2007) described the area encompassing NHW and KHW as a ―hotspot‖ of orchid diversity due to the range of habitats they cover.
A factor contributing to species differences between KHW and NHW may be topographical. The
topography of NHW, while similar to KHW, has been affected by drifting dunes perpendicular to the shoreline creating deep, almost circular, ponds as opposed to the broad shallow parallel swales of KHW (Odum et al. 1988). On visual inspection of aerial photographs and soil maps, it appears that a greater percentage of KHW is covered in forested wetlands. The shallower swales might allow for the greater establishment of woody wetland species. There are several native wetland associated species present such as Fraxinus pennsylvanica
11
velutina Lam. Though both sites have dry sandy ridges, there is a greater percentage of sandy, excessively drained soils at NHW (Tant 1992). In addition, two of the most species rich areas of NHW are a maintained powerline cut that runs through the reserve and bisects high ridges and low swales and a freshwater seepage marsh bounded by the live sand dunes of Run Hill State Natural Area (Krings 2010). The absence of canopy cover, coupled with both wet and dry habitats, and repeated disturbance, allows for the development and maintenance of a diverse herb layer. Species associated with these areas include Xyris L. spp., Impatiens capensis Meerb., Persicaria sagittata (L.) H. Gross. In contrast, the powerline cut at KHW runs along a ridge—and thus at a relatively even altitude and drainage class, and remains in many places just bare sand. In contrast to Nags Head Woods, KHW also lacks a sandy beach along Currituck Sound; instead the edges of the site are dominated by vegetation such Juncus roemerianus or Spartina spp., and sound shore species such as
Cakile edentula (Bigelow) Hook. are absent.
12
K
EYS TO THE
M
AJOR
V
ASCULAR
P
LANT
G
ROUPS
1.Plant reproducing by spores ... Pteridophytes 1‘. Plant reproducing by seeds ... 2. 2. (1‘.) Leaves needlelike or scale like, < 3 mm wide; plant bearing cones ... Gymnosperms 2‘. Combination not as above ... 3. 3. (2‘.) Plant with 2 or more of the following characters: leaves parallel veined; floral parts in 3s; stem vascular
bundles scattered; cotyledon 1 ... Monocotyledons 3‘. Plant with 2 more more of the following characters: leaves not parallel veined; floral parts in 4s and 5s; stem
vascular bundles in a ring; cotyledons 2 ...Basal angiosperms, magnoliids, higher dicots
Pteridophytes
Key adapted from Radford et al. (1968) and Smith (1993).
1. Plant free-floating aquatic ... Azollaceae 1‘. Plant terrestrial or epiphytic ... 2.
2. (1‘.) Plant epiphytic ... Polypodiaceae 2‘. Plant rooted terrestrially ... 3. 3. (2‘.) Plants with scalelike leaves, borne on dichotomously branching stems ... Psilotaceae 3‘. Plants with expanded leaves, not on dichotomously branching stems ... 4. 4. (3‘.) Leaf divided into a sterile, photosynthetic portion and fertile, spore-bearing spike .. Ophioglossaceae 4‘. Leaves not as above ... 5. 5.(4‘.) Leaf bases with stipules or thin lateral wings ... Osmundaceae 5‘. Leaf bases without stipules ... 6.
13
A
SPLENIACEAE
1.
A
SPLENIUML.
Asplenium platyneuron(L.) Britton, Sterns & Poggenb. [= RAB]
Acrostichum platyneuron L.
Maritime woods. Infrequent. Apr–Oct. (Fig. 8)
Burk et al. b21-4 (NCU); Clark 180 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (Krings 450, NCSC).
REFERENCES
Wagner, W.H. Jr., R.C. Morgan, and C.R. Werth. 1993. Asplenium. In: Flora of North America, Flora North America Editorial Committee (eds.). Oxford Univ. Press, New York.
A
ZOLLACEAE
1.
A
ZOLLALam.
Azolla caroliniana Willd. [= RAB]
Interdunal ponds. Rare. Jun–Sep. (Fig. 9)
Burk (1961) reported this taxon from a pool near Kitty Hawk, citing a collection by Radford, however no voucher has been seen by the author and unfortunatley, the voucher for Azolla caroliniana, Clark 329, was damaged and destroyed.
Other Bodie Island specimens seen: Kill Devil Hills (Musselman 4104, NCU); Nags Head Woods (Atkinson & Radford 2495, NCU; Krings 1151, NCSC). ♣
REFERENCES
14
B
LECHNACEAE
1.
W
OODWARDIASm.
Key adapted from Cranfill (1993).1. Leaves strongly dimorphic; sterile blades pinnatifid, veins forming 2 or more rows of areoles between costae and margin, veins free only at margin... Woodwardia areolata
1‘. Leaves monomorphic; sterile blades pinnate, veins forming only one row of areoles between costae and
margin ...Woodwardia virginica
Woodwardia areolata (L.) Moore [= RAB]
Acrostichum areolatum L.
Maritime swamp forests and interdunal swales. Occasional. Jun–Sep. (Fig. 10)
Burk et al. b21-31 (NCU); Clark 24, 57, 217, 436 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (Krings 1228, 1360, 1361, NCSC).
Woodwardia virginica (L.) Sm. [= RAB]
Blechnum virginicum L.
Maritime swamp forests and interdunal swales. Infrequent. Jun–Sep. (Fig. 11)
Burk et al. b21-32 (NCU); Clark 442, 475 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (Lynch 113, NCSC).
REFERENCES
Cranfill, R.B. 1993. Woodwardia. In: Flora of North America, Flora North America Editorial Committee (eds.). Oxford Univ. Press, New York.
D
ENNSTAEDTIACEAE
1.
P
TERIDIUMGled. ex Scop.
Pteridium aquilinum (L.) Kuhn [= RAB]
Pteris aquilina L.
Maritime woods, sandy edges, and the powerline right of way. Frequent. Jul–Sep. (Fig. 12)
15 REFERENCES
Jacobs, C.A. and J.H. Peck. 1993. Pteridium. In: Flora of North America, Flora North America Editorial Committee (eds.). Oxford Univ. Press, New York.
O
PHIOGLOSSACEAE
Key adapted from Weakley (2010).
1. Fertile stalk joined to sterile leaf blade near the base of the sterile leaf blade, above the rhizome and well about the surface of the ground; leaves deciduous ... Botrypus virginianus
1‘. Fertile stalk joined to stalk of sterile leaf blade near the rhizome, at or below the surface of the ground;
leaves evergreen ... Sceptridium dissectum
1.
B
OTRYPUSMichx.
Botrypus virginianus (L.) Michx. [< Botrychium virginianum (L.) Sw. sensu RAB]
Osmunda virginiana L.
Moist forest edges. Rare. Arp–Jun. (Fig. 13)
Clark 469 (NCSC). No additional specimens seen from Bodie Island. There have been no other reports of this taxon on the Outer Banks.
2. S
CEPTRIDIUMLyon
Sceptridium dissectum (Spreng.) Lyon [= Botrychium dissectum Spreng. sensu RAB]
Botrychium dissectum Spreng.
Moist forest edges. Rare. Aug–Oct. (Fig. 14)
Clark 726 (NCSC). Other Bodie Island specimens seen: Duck (Burk et al. b34-1, NCU). Both Burk (1961) and Stalter and Lamont (1997) reported Sceptridium dissectum from Nags Heads Woods.
REFERENCES
16
O
SMUNDACEAE
Key adapted from Whetstone and Atkinson (1993) and Weakley (2010).
1. Fertile leaves with dimorphic pinnae, some spore-bearing, some not; tuft of hairs absent on abaxial surface of pinnae near base ... Osmunda
1‘. Fertile leaves with monomorphic pinnae, all spore-bearing; tuft of reddish hairs persistent on abaxial surface
of pinnae near base... Osmundastrum
1.
O
SMUNDAL.
Osmunda regalis L. var. spectabilis (Willd.) A. Gray [= RAB]
Osmunda spectabilis Willd.
Maritime swamp forests and interdunal swales. Occasional. Mar–Jun. (Fig. 15)
Clark 64, 449 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (Krings 538, 1357, 1358, NCSC).
2.
O
SMUNDASTRUMC.Presl
Osmundastrum cinnamomeum (L.) C.Presl [=Osmunda cinnamomea sensu RAB]
Osmunda cinnamomea L.
Maritime swamp forests and interdunal swales. Occasional. Mar–May. (Fig. 16)
Clark 489 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (Krings 534, 1356, NCSC).
REFERENCES
Whetstone, R.D. and T.A. Atkinson.. 1993. Osmundaceae. In: Flora of North America, Flora North America Editorial Committee (eds.). Oxford Univ. Press, New York.
P
OLYPODIACEAE
1.
P
LEOPELTISHumb. & Bonpl. ex Willd.
Pleopeltis polypodioides (L.) E.G. Andrews & Windham var. michauxiana (Weath.) E.G. Andrews & Windham [< Polypodium polypodiodes sensu RAB]
Polypodium polypodioides (L.) Watt var. michauxianum Weath.
Maritime evergreen and deciduous forests, epiphytic. Infrequent. Jun–Oct. (Fig. 17)
17 REFERENCES
Andrews, E.G. and M.D. Windham. 1993. Pleopeltis. In: Flora of North America, Flora North America Editorial Committee (eds.). Oxford Univ. Press, New York.
P
SILOTACEAE
1.
P
SILOTUMSw.
Psilotum nudum (L.) P. Beauv. [= RAB]
Lycopodium nudum L.
Maritime deciduous forest. Rare. Apr–Sep.
This is a county record for Dare County, NC; the only other documented occurrences of this taxon in North Carolina have been in Chowan County, NC and naturalized around a few buildings on the North Carolina State University campus in Wake County, NC (Perry & Musselman 1994). The Kitty Hawk Woods population is near a road originally intended to bisect a future housing development before the land was sold to the state. It is possible that in the course of the initial development of the area, individuals were introduced from building materials or mulch from further south. This species is on the North Carolina Heritage Program‘s rare plant watch list (S1 G5)
Clark 335 (NCSC). No additional specimens seen from Bodie Island.
REFERENCES
Perry, J.P. III and L.J. Musselman. 1994. Psilotum nudum new to North Carolina. Amer. Fern. J. 84:102-104.
Thieret, J.W. 1993. Psilotaceae. In: Flora of North America, Flora North America Editorial Committee (eds.). Oxford Univ. Press, New York.
T
HELYPTERIDACEAE
1.
T
HELYPTERISSchmidel
Thelypteris palustris Schott var. pubescens (G. Lawson) Fernald [< Thelypteris palustris Schott sensu RAB]
Lastrea thelypteris (L.) Bory var. pubescens G. Lawson
18
Clark 219, 471 (NCSC). Other Bodie Island specimens seen: Bodie Island (Godfrey & Fox 49568, NCSC); Nags Head Woods (Burk et al. b22-15, b58-11, NCU; Downs 332, NCSC; Krings 1354, 1362, 1363, NCSC).
REFERENCES
19
Gymnosperms
Key adapted from Eckenwalder and Thieret (1993).
1. Leaves needle-like or scale-like, not in fascicles; cones 1.5–4 cm ... Cupressaceae 1‘.Leaves needle-like, in fascicles; cones 4–12 cm ...Pinaceae
C
UPRESSACEAE
Key adapted from Watson (1993 ) and Weakley (2010).
1. Trees or shrubs evergreen; needles opposite or whorled; female cones fleshy and berry-like ... ... Juniperus virginiana
1‘. Trees deciduous; needles alternate; female cones woody ... Taxodium distichum
1. J
UNIPERUSL.
Juniperus virginiana
L. [= RAB]
Maritime woods, creek margins and upland edges of brackish marshes. Frequent. Jan–Feb; Oct–Nov. (Fig. 19)
Clark 18, 56 (NCSC). Other Bodie Island specimens seen: Duck (Burk et al. b20-4I, NCU); Nags Head Woods (Krings 373, NCSC); Southern Shores (Dunes of Dare Garden Club s.n., NCSC).
2. T
AXODIUMRich.
Taxodium distichum
(L.) Rich. [= RAB]
Cupressus disticha L.
Maritime swamps. Abundant. Mar–Apr; Oct. (Fig. 20)
In Kitty Hawk Woods, T. distichum occurs as a dominant canopy species in the nontidal wetlands of the eastern portion of the reserve.
Clark 88, 401, 413 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (McCall 3, NCSC).
REFERENCES
20
P
INACEAE
1. P
INUSL.
Key adapted from Kral (1993).1. Bark plates with evident resin pockets; leaves mostly two per fascicle, longer leaves mostly ≤13 cm; seed cones semi-persistant, 4–6 cm long when mature ... Pinus echinata
1‘. Bark plates without evident resin pockets; leaves mostly three per fascicle, longer leaves mostly ≥15 cm;
seed cones not persistant, 6–12 cm long when mature ... Pinus taeda
Pinus echinata Mill. [= RAB]
Dry, upland woods. Rare. Mar–Apr; Sep–Oct. (Fig. 21)
Stalter and Lamont (1997) reported this species only from Nags Head Woods and Burk (1961) reported it from there and Buxton Woods, though he theorized it was possibly planted at that site. There are only a few individuals occurring on the sandy ridges in the western part of Kitty Hawk Woods.
Clark 516 (NCSC). Other Bodie Island specimens seen: Nags Head Woods (Cooper 2816, NCSC).
Pinus taeda L. [= RAB]
Maritime evergreen forests, clearings and disturbed edges of maritime deciduous forests, creek and brackish marsh margins . Abundant. Mar–Apr; Oct–Nov. (Fig. 22)
Clark 14, 452 (NCSC). Other specimens seen: Bodie Island (Fox & Whitford 1087, NCSC); Nags Head Woods (Fox & Whitford 383, 387, NCSC; Krings 564, NCSC).
REFERENCES
