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Aquatic and Standing Water Plants
of the Central Midwest
Books in the Aquatic and Standing Water Plants
of the Central Midwest Series by Robert H. Mohlenbrock
Cyperaceae: Sedges
Filicineae, Gymnospermae, and Other Monocots, Excluding Cyperaceae:
Ferns, Conifers, and Other Monocots, Excluding Sedges
Acanthaceae to Myricaceae: Water Willows to Wax Myrtles
Nelumbonaceae to Vitaceae: Water Lotuses to Grapes

Other Southern Illinois University Press Books

by Robert H. Mohlenbrock
Guide to the Vascular Flora of Illinois, revised and enlarged edition
Distribution of Illinois Vascular Plants, with Douglas M. Ladd
A Flora of Southern Illinois, with John W. Voigt

In the Illustrated Flora of Illinois Series

Ferns, 2nd edition
Flowering Plants: Basswoods to Spurges
Flowering Plants: Flowering Rush to Rushes
Flowering Plants: Hollies to Loasas
Flowering Plants: Lilies to Orchids
Flowering Plants: Magnolias to Pitcher Plants
Flowering Plants: Nightshades to Mistletoe
Flowering Plants: Pokeweeds, Four-o’clocks, Carpetweeds,
Cacti, Purslanes, Goosefoots, Pigweeds, and Pinks
Flowering Plants: Smartweeds to Hazelnuts
Flowering Plants: Willows to Mustards
Grasses: Bromus to Paspalum, 2nd edition
Grasses: Panicum to Danthonia, 2nd edition
Sedges: Carex
Sedges: Cyperus to Scleria, 2nd edition
Aquatic and Standing Water Plants of the Central Midwest

Nelumbonaceae to Vitaceae
Water Lotuses to Grapes

Robert H. Mohlenbrock

Southern
Illinois
University Press
Carbondale and
Edwardsville
Copyright © 2010 by the Board of Trustees,
Southern Illinois University
All rights reserved
Printed in the United States of America
13 12 11 10 4 3 2 1

Cover Illustration by Mark W. Mohlenbrock

Library of Congress Cataloging-in-Publication Data
Mohlenbrock, Robert H., 1931–
Nelumbonaceae to Vitaceae : water lotuses to grapes / Robert H. Mohlenbrock.
p. cm. — (Aquatic and standing water plants of the central Midwest)
Includes bibliographical references and index.
ISBN-13: 978-0-8093-2894-9 (alk. paper)
ISBN-10: 0-8093-2894-1 (alk. paper)
1. Angiosperms—Middle West—Identification. 2. Angiosperms—Middle West—Pictorial
works. I. Title. II. Series: Mohlenbrock, Robert H., 1931– Aquatic and standing water plants
of the central Midwest
QK128.M645 2010
581.977—dc22 2009025824

Printed on recycled paper.

The paper used in this publication

meets the minimum requirements
of American National Standard
for Information Sciences—
Permanence of Paper for
Printed Library Materials,
ANSI Z39.48-1992. '
 This book is dedicated to Grenville Lucas, former director of Kew Gardens and former

chairman of the Species Survival Commission of the International Union for the

Conservation of Nature, who selected me to chair the North American Plant Group of

the Species Survival Commission, a position I held for fourteen years.

Contents

List of Illustrations / IX

Series Preface / XIII

Map of the Central Midwest / XVIII

Descriptions and Illustrations / 1

Glossary / 443

Illustration Credits / 449

Index to Genera and Species / 451

Index to Common Names / 461

Illustrations

1. Nelumbo lutea 12 46. Phlox paniculata 68

2. Nuphar advena 14 47. Polemonium reptans 69
3. Nuphar variegatum 15 48. Polygala cruciata 70
4. Nymphaea odorata 16 49. Polygala sanguinea 71
5. Nymphaea tuberosa 17 50. Brunnichia ovata 73
6. Nyssa aquatica 18 51. Fallopia cristata 75
7. Nyssa biflora 19 52. Fallopia scandens 76
8. Nyssa sylvatica 20 53. Persicaria amphibium 79
9. Forestiera acuminata 22 54. Persicaria amphibium 80
10. Fraxinus lanceolata 24 55. Persicaria bicornis 82
11. Fraxinus nigra 25 56. Persicaria careyi 83
12. Fraxinus pennsylvanica 26 57. Persicaria coccinea 85
13. Fraxinus profunda 27 58. Persicaria glabra 86
14. Chamerion angustifolium 29 59. Persicaria hydropiper 87
15. Epilobium ciliatum 30 60. Persicaria hydropiperoides 89
16. Epilobium coloratum 32 61. Persicaria lapathifolia 90
17. Epilobium glandulosum 32 62. Persicaria longiseta 92
18. Epilobium hirsutum 33 63. Persicaria maculosa 94
19. Epilobium leptophyllum 34 64. Persicaria opelousana 95
20. Epilobium strictum 35 65. Persicaria pensylvanica 97
21. Gaurella canescens 36 66. Persicaria punctata 98
22. Ludwigia alternifolia 38 67. Persicaria robustior 100
23. Ludwigia decurrens 39 68. Persicaria setacea 101
24. Ludwigia glandulosa 40 69. Polygonum ramosissimum 103
25. Ludwigia hirtella 41 70. Rumex altissimus 105
26. Ludwigia leptocarpa 43 71. Rumex britannica 106
27. Ludwigia microcarpa 44 72. Rumex conglomeratus 107
28. Ludwigia palustris 44 73. Rumex crispus 108
29. Ludwigia peploides 46 74. Rumex crispus 109
30. Ludwigia polycarpa 47 75. Rumex fueginus 110
31. Ludwigia repens 48 76. Rumex obtusifolius 112
32. Ludwigia sphaerocarpa 49 77. Rumex obtusifolius 113
33. Ludwigia uruguayensis 50 78. Rumex occidentalis 114
34. Oenothera fruticosa 52 79. Rumex stenophyllus 115
35. Oenothera perennis 53 80. Rumex triangulivalvis 116
36. Oenothera pilosella 54 81. Rumex verticillatus 118
37. Oenothera tetragona 55 82. Tracaulon arifolium 119
38. Parnassia glauca 56 83. Tracaulon sagittatum 121
39. Parnassia grandiflora 57 84. Anagallis minima 123
40. Plantago cordata 59 85. Hottonia inflata 125
41. Platanus occidentalis 61 86. Lysimachia X commixta 127
42. Podostemum ceratophyllum 62 87. Lysimachia fraseri 128
43. Phlox carolina 64 88. Lysimachia nummularia 129
44. Phlox glaberrima 65 89. Lysimachia terrestris 131
45. Phlox maculata 66 90. Naumbergia thyrsiflora 132

IX
 91. Samolus ebracteatus 134 139. Argentina anserina 197
92. Samolus parviflorus 135 140. Comarum palustre 198
93. Steironema ciliatum 136 141. Crataegus acutifolia 202
94. Steironema hybridum 138 142. Crataegus apiomorpha 203
95. Steironema lanceolatum 140 143. Crataegus coccinioides 204
96. Steironema quadriflorum 141 144. Crataegus collina 204
97. Steironema radicans 142 145. Crataegus crus-galli 206
98. Anemone canadensis 144 146. Crataegus disjuncta 207
99. Caltha palustris 146 147. Crataegus disperma 207
100. Clematis crispa 147 148. Crataegus dispessa 208
101. Clematis viorna 148 149. Crataegus fecunda 209
102. Coptis groenlandica 149 150. Crataegus hannibalensis 210
103. Myosurus minimus 151 151. Crataegus incaedua 210
104. Ranunculus abortivus 153 152. Crataegus laxiflora 211
105. Ranunculus ambigens 154 153. Crataegus lettermanii 212
106. Ranunculus carolinianus 156 154. Crataegus locuples 212
107. Ranunculus cymbalaria 157 155. Crataegus lucorum 213
108. Ranunculus flabellaris 158 156. Crataegus marshallii 214
109. Ranunculus gmelinii 160 157. Crataegus mollis 215
110. Ranunculus laxicaulis 161 158. Crataegus neobushii 216
111. Ranunculus longirostris 162 159. Crataegus nitida 217
112. Ranunculus macounii 163 160. Crataegus noelensis 218
113. Ranunculus pensylvanicus 164 161. Crataegus ovata 218
114. Ranunculus pusillus 165 162. Crataegus palmeri 219
115. Ranunculus recurvatus 167 163. Crataegus pedicellata 220
116. Ranunculus repens 168 164. Crataegus peoriensis 221
117. Ranunculus sardous 169 165. Crataegus populnea 221
118. Ranunculus sceleratus 171 166. Crataegus pringlei 222
119. Ranunculus septentrionalis 172 167. Crataegus putnamiana 223
120. Ranunculus septentrionalis var. 168. Crataegus regalis 223
caricetorum 173 169. Crataegus reverchonii 224
121. Ranunculus subrigidus 174 170. Crataegus simulata 225
122. Ranunculus trichophyllus 175 171. Crataegus spathulata 225
123. Thalictrum dasycarpum 177 172. Crataegus vailiae 226
124. Thalictrum dasycarpum var. 173. Crataegus vallicola 227
hypoglaucum 178 174. Crataegus verruculosa 228
125. Thalictrum pubescens 179 175. Crataegus viridis 229
126. Thalictrum revolutum 180 176. Dalibarda repens 230
127. Trautvetteria carolinensis 181 177. Filipendula rubra 231
128. Berchemia scandens 183 178. Fragaria virginiana 232
129. Frangula alnus 184 179. Geum aleppicum 233
130. Rhamnus alnifolia 186 180. Geum canadense 234
131. Rhamnus cathartica 187 181. Geum laciniatum 235
132. Rhamnus lanceolata 188 182. Geum rivale 236
133. Agrimonia parviflora 191 183. Malus angustifolia 237
134. Agrimonia pubescens 192 184. Malus coronaria 238
135. Amelanchier alnifolia 193 185. Malus ioensis 239
136. Amelanchier interior 194 186. Pentaphylloides floribunda 241
137. Amelanchier laevis 195 187. Photinia floribunda 242
138. Amelanchier sanguinea 196 188. Photinia melanocarpa 243

X / ILLUSTRATIONS
189. Photinia pyrifolia 244 239. Salix discolor 304
190. Physocarpus opulifolius 246 240. Salix eriocephala 306
191. Potentilla millegrana 247 241. Salix fragilis 307
192. Potentilla paradoxa 248 242. Salix glaucophylloides var.
193. Potentilla pentandra 249 glaucophylla 309
194. Potentilla rivalis 249 243. Salix interior 310
195. Prunus angustifolia 251 244. Salix lucida 312
196. Prunus hortulana 252 245. Salix nigra 313
197. Prunus munsoniana 253 246. Salix pedicellaris 315
198. Prunus serotina 254 247. Salix pentandra 316
199. Rosa multiflora 255 248. Salix petiolaris 318
200. Rosa palustris 256 249. Salix purpurea 319
201. Rosa setigera 257 250. Salix rigida 321
202. Rubus hispidus 259 251. Salix sericea 322
203. Rubus occidentalis 260 252. Salix serissima 323
204. Rubus odoratus 261 253. Salix X subsericea 325
205. Rubus pubescens 261 254. Comandra umbellata 326
206. Rubus strigosus 262 255. Cardiospermum halicacabum 328
207. Rubus trivialis 263 256. Sideroxylon lycioides 329
208. Sanguisorba canadensis 264 257. Sarracenia purpurea 331
209. Spiraea alba 265 258. Anemopsis californica 333
210. Spiraea tomentosa 266 259. Saururus cernuus 333
211. Cephalanthis occidentalis 268 260. Chrysosplenium americanum 335
212. Diodia teres 269 261. Chrysospenium iowense 336
213. Diodia virginiana 270 262. Mitella nuda 336
214. Callum aparine 272 263. Penthorum sedoides 338
215. Galium asprellum 273 264. Saxifraga pensylvanica 339
216. Galium boreale 274 265. Agalinis besseyana 342
217. Galium labradoricum 275 266. Agalinis paupercula 342
218. Galium obtusum 276 267. Agalinis purpurea 343
219. Galium tinctorium 277 268. Agalinis skinneriana 344
220. Galium trifidum 278 269. Agalinis tenuifolia 345
221. Galium triflorum 279 270. Bacopa caroliniana 346
222. Houstonia caerulea 280 271. Bacopa rotundifolia 347
223. Houstonia purpurea 281 272. Buchnera americana 348
224. Houstonia pusilla 281 273. Castilleja coccinea 349
225. Mitchella repens 283 274. Chelone glabra 350
226. Oldenlandia boscii 283 275. Chelone obliqua 351
227. Oldenlandia uniflora 284 276. Gratiola aurea 352
228. Spermacoce glabra 285 277. Gratiola neglecta 353
229. Populus balsamifera 287 278. Gratiola virginiana 354
230. Populus deltoides 289 279. Graiola viscidula 355
231. Populus heterophylla 291 280. Leucospora multifida 356
232. Populus tremuloides 292 281. Limosella aquatica 357
233. Salix alba 295 282. Lindernia anagallidea 358
234. Salix amygdaloides 297 283. Lindernia dubia 359
235. Salix babylonica 298 284. Mazus pumilus 360
236. Salix bebbiana 300 285. Mecardonia acuminata 361
237. Salix candida 301 286. Melampyrum lineare 362
238. Salix caroliniana 303 287. Mimulus alatus 364

ILLUSTRATIONS / XI
288. Mimulus glabratus 364 318. Phyla cuneifolia 404
289. Mimulus guttatus 365 319. Phyla lanceolata 405
290. Mimulus ringens 366 320. Phyla nodiflora 406
291. Pedicularis lanceolata 367 321. Verbena hastata 407
292. Penstemon alluviorum 368 322. Verbena urticifolia 408
293. Veronica americana 370 323. Viola affinis 411
294. Veronica anagallis-aquatica 371 324. Viola blanda 412
295. Veronica comosa 372 325. Viola canadensis var. rugulosa 413
296. Veronica peregrina 373 326. Viola conspersa 414
297. Veronica scutellata 374 327. Viola cucullata 416
298. Veronicastrum virginicum 375 328. Viola lanceolata 417
299. Solanum dulcamara 377 329. Viola macloskeyi ssp. pallens 419
300. Styrax americanus 379 330. Viola missouriensis 420
301. Tilia heterophylla 380 331. Viola nephrophylla 421
302. Celtis laevigata 382 332. Viola pratincola 422
303. Celtis occidentalis 383 333. Viola primulifolia 424
304. Planera aquatica 385 334. Viola sagittata 425
305. Ulmus alata 387 335. Viola sororia 426
306. Ulmus americana 388 336. Viola striata 428
307. Boehmeria cylindrica 390 337. Ampelopsis arborea 430
308. Laportea canadensis 391 338. Ampelopsis cordata 430
309. Pilea fontana 393 339. Parthenocissus inserta 432
310. Pilea pumila 394 340. Parthenocissus quinquefolia 432
311. Urtica chamaedryoides 396 341. Vitis aestivalis 434
312. Urtica dioica 397 342. Vitis cinerea 435
313. Valeriana ciliata 398 343. Vitis palmata 436
314. Valeriana uliginosa 399 344. Vitis riparia 437
315. Valerianella chenopodiifolia 400 345. Viola rotundifolia 438
316. Valerianella radiata 402 346. Vitis vulpina 439
317. Valerianella umbilicata 402

XII / ILLUSTRATIONS
Series Preface

The purpose of the four books in the Aquatic and Standing Water Plants of the
Central Midwest series is to provide illustrated guides to the plants of the central
Midwest that may live in standing or running water at least three months a year,
though a particular species may not necessarily live in standing or running water
during a given year. The states covered by these guides include Iowa, Illinois, Indi-
ana, Ohio, Kansas, Kentucky, Missouri, and Nebraska, except for the Cumberland
Mountain region of eastern Kentucky, which is in a different biological province.
Since 1990, I have taught weeklong wetland plant identification courses in all of
these states on several occasions.
The most difficult task has been to decide what plants to include and what plants
to exclude from these books. Three groups of plants are within the guidelines of
the manuals. One group includes those aquatic plants that spend their entire life
with their vegetative parts either completely submerged or at least floating on the
water’s surface.
This group includes obvious submerged aquatics such as Ceratophyllum, the
Najadaceae, the Potamogetonaceae, Elodea, Cabomba, Brasenia, Nymphaea, some
species of Ranunculus, Utricularia, and a few others.
Plants in a second group are called emergents. These plants typically are rooted
under water, with their vegetative parts standing above the water surface. Many of
these plants can live for a long period of time, even their entire life, out of the water.
Included in this group are Sagittaria, Alisma, Peltandra, Pontederia, Saururus, Justicia,
and several others.
The most difficult group of plants that I had to consider is made up of those
wetland plants that live most or all of their lives out of the water, but which on
occasion can live at least three months in water. I concluded that I would include
within these books only those species that I personally have observed in standing
water during the year, or which have been reported in the literature as living in
water. In this last group, for example, I have included Poa annua, since Yatskievych,
in his Steyermark’s Flora of Missouri (1999), indicates that this species may occur
in standing water, even though I have not observed this myself.
In these books, I have included most plants that live in lakes, ponds, rivers, and
streams, as well as plants that live in marshes, bogs, fens, wet meadows, sedge
meadows, wet prairies, swampy woods, and temporary depressions in woods, on
cliffs, and in barrens.
Swink and Wilhelm (1999) consider a marsh as a transition between aquatic
communities and drier communities, or in large flats which are regularly inun-
dated by shallow surface water for much of the growing season. Cattails are often
frequent in marshes. Trees are generally uncommon in a typical marsh.
Wet meadows, as treated here, are similar to marshes, but without pools of
water. Woody plants are generally absent, and cattails do not dominate and are
often not present.

XIII
 Sedge meadows are similar, except that the overwhelming majority of species is
composed of members of the sedge family (Cyperaceae). Wet prairies are also simi-
lar, but the dominant vegetation consists of grasses. Ladd (1995) recognizes dry,
moist, and wet prairies, and only the last of these is included in these manuals.
Fens are wetlands where the underlying groundwater is rich with calcium or
magnesium carbonates. Fens may or may not have woody plants present. Swink
and Wilhelm (1999) also recognize marly fens that occur on open prairie slopes
and hillside fens that are wooded seeps on steep bluffs.
Bogs are habitats typified by acidic, usually organic, substrates. However, bogs
that have been influenced by carbonate-rich water have been called alkaline bogs
by Swink and Wilhelm. Where minerotrophic water is insignificant, acid bogs
develop. Floating sedge mats are acidic and often develop in sand flats or basins that
rise and fall with the water table.
Swampy woods are forested wetlands in poorly drained flats or basins. In areas
around Lake Michigan, swamps occur in wet sandy flats and on the moraine in wet
depressions and in large flats behind the high dunes. In the southern part of the
central Midwest, swampy woods may have standing water throughout the year.
Occasional depressions that fill with water harbor species that are included
in these books. These depressions may occur in various kinds of woodlands, on
exposed sandstone blufftops, and, in Kentucky, even on rocky barrens. Seeps may
occur on the faces of cliffs, and the plants that live in this constantly wet habitat are
also included in these books.
It is likely that I failed to include a few plants that should have been included, but
that I had not observed myself.
The nomenclature that I have used in these manuals reflects my own opinion
as to what I believe the scientific names should be. If these names differ from those
used by the United States Fish and Wildlife Service, I have indicated this. A partial
list of synonymy is included for each species, particularly accounting for synonyms
that have been in use for several decades.
After the description of each plant, I have indicated the habitats in which
the plant may be found, followed by the states in which the plant occurs. I have
indicated the U.S. Fish and Wildlife Service wetland designation for each species for
the states in which each occurs. In 1988, the National Wetland Inventory Section
of the United States Fish and Wildlife Service attempted to give a wetland designa-
tion for every plant occurring in the wild in the United States. The states covered
by these aquatic manuals occur in three regions of the Fish and Wildlife Service.
Kentucky and Ohio are in region 1; Illinois, Indiana, Iowa, and Missouri are in
region 3; and Kansas and Nebraska are in region 5. Definitions of the Fish and
Wildlife Service wetland categories are:
OBL (Obligate Wetland). Occur almost always under natural conditions in wet-
lands, at least 99% of the time.
FACW (Facultative Wetland). Usually occur in wetlands 67–99% of the time,
but occasionally found in non-wetlands.
FAC (Facultative). Equally likely to occur in wetlands or non-wetlands 34–66%
of the time.

XIV / SERIES PREFACE

 FACU (Facultative Upland). Usually occur in non-wetlands 67–99% of the time,
but occasionally found in wetlands.
UPL (Upland). Occur in uplands at least 99% of the time, but under natural
conditions not found in wetlands.
NI (Not Indicated). Due to insufficient information.
A plus or minus sign (+ or –) may appear after FACW, FAC, and FACU. The plus
means leaning toward a wetter condition; the minus means leaning toward a drier
condition.
Although the Fish and Wildlife Service made changes to the wetland status of
several species in an updated version in 1997, this later list has never been approved
by the Congress of the United States.
Following this is one or more common names currently employed in the central
Midwest. A brief discussion of distinguishing characteristics and nomenclatural
notes is often included. Illustrations accompany each species, showing the diag-
nostic characteristics. In some of the illustrations, a gap in the stem signifies that a
portion of the stem has been omitted due to space limitations.
The sequence of families in these aquatic manuals is as follows:

1. Azollaceae 28. Potamogetonaceae 55. Caprifoliaceae

2. Blechnaceae 29. Ruppiaceae 56. Caryophyllaceae
3. Equisetaceae 30. Scheuchzeriaceae 57. Ceratophyllaceae
4. Isoetaceae 31. Sparganiaceae 58. Convolvulaceae
5. Lycopodiaceae 32. Typhaceae 59. Cornaceae
6. Marsileaceae 33. Xyridaceae 60. Corylaceae
7. Onocleaceae 34. Zannichelliaceae 61. Cuscutaceae
8. Osmundaceae 35. Acanthaceae 62. Droseraceae
9. Thelypteridaceae 36. Aceraceae 63. Elatinaceae
10. Pinaceae 37. Amaranthaceae 64. Ericaceae
11. Taxodiaceae 38. Anacardiaceae 65. Escalloniaceae
12. Acoraceae 39. Apiaceae 66. Euphorbiaceae
13. Alismataceae 40. Apocynaceae 67. Fabaceae
14. Araceae 41. Aquifoliaceae 68. Fagaceae
15. Butomaceae 42. Araliaceae 69. Gentianaceae
16. Cyperaceae 43. Aristolochiaceae 70. Grossulariaceae
17. Eriocaulaceae 44. Asclepiadaceae 71. Haloragidaceae
18. Hydrocharitaceae 45. Asteraceae 72. Hamamelidaceae
19. Iridaceae 46. Balsaminaceae 73. Hippuridaceae
20. Juncaceae 47. Betulaceae 74. Hydrophyllaceae
21. Juncaginaceae 48. Bignoniaceae 75. Hypericaceae
22. Lemnaceae 49. Boraginaceae 76. Juglandaceae
23. Maranthaceae 50. Brassicaceae 77. Lamiaceae
24. Najadaceae 51. Cabombaceae 78. Lauraceae
25. Orchidaceae 52. Caesalpiniaceae 79. Leitneriaceae
26. Poaceae 53. Callitrichaceae 80. Lentibulariaceae
27. Pontederiaceae 54. Campanulaceae 81. Limnanthaceae

SERIES PREFACE / XV
82. Linaceae 95. Plantaginaceae 107. Sarraceniaceae
83. Lythraceae 96. Platanaceae 108. Saururaceae
84. Malvaceae 97. Podostemaceae 109. Saxifragaceae
85. Melastomaceae 98. Polemoniaceae 110. Scrophulariaceae
86. Menyanthaceae 99. Polygalaceae 111. Solanaceae
87. Molluginaceae 100. Polygonaceae 112. Styracaceae
88. Myricaceae 101. Primulaceae 113. Ulmaceae
89. Nelumbonaceae 102. Ranunculaceae 114. Urticaceae
90. Nymphaeaceae 103. Rhamnaceae 115. Valerianaceae
91. Nyssaceae 104. Rosaceae 116. Verbenaceae
92. Oleaceae 105. Rubiaceae 117. Violaceae
93. Onagraceae 106. Salicaceae 118. Vitaceae
94. Parnassiaceae

This volume consists of families 89 to 118. Volume 3 has the remaining dicots,
from families 35–88. The first volume in the series was devoted exclusively to the
Cyperaceae, family 16. The second volume included the ferns, gymnosperms, and
monocots, families 1–15 and 17–34.

XVI / SERIES PREFACE

Aquatic and Standing Water Plants
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