Professional Documents
Culture Documents
2
Angiosperms: Monocotyledons
Garrett E. Crow and C. Barre Hellquist
The University
of Wisconsin Press
Figures 60. 63.75.112. 118. 136. 147. 193.216.259.261. 297. The University of Wisconsin Press
316.317.318.319.324.330.419.421.585. and 586 are reprinted 2537 Daniels Street
in part or in whole. with permission. from The New Britton and Brown Madison. Wisconsin 53718
Illustrated Flora oj the Northeastern United States and Adjacent Canada. by
Henry A. Gleason. Copyright 1952 by The New York Botanical 3 Henrietta Street
Garden. London WC2E 8LU. England
Figures 56. 68. 75. 77. 80. 83.97.98. 103. 104. 108. 124. 128.
129.165.166.170.171.174.175.176.177.178.179.180.181.
183.184.198.201.204.213.224.225.238.245.246.247.248.
249.250.253.255.270.274.275.276.278.280.281.283.286.
287.288.289.338.342.360.362.382.388.396.403.404.407.
408.409.410.412.415.416.417.420.422.42~426.42~428.
430.433.441.443.446.449.453.454.455.456.458.464.470.
471.472.473.474.475.476.477.478.479.480.481.482.488.
491.492.494.495.496.497.498.499.500.501.502.503.505.
508.509.510.511.512.516.517.520.521.525.527.532.533.
534. 535. 536. 552. 553. 554. 561. 568. 585. 590. 592. 594. 595.
597. 598. and 60S are reprinted in part or in whole from Aquatic and
Wetland Plants oj the Southeastern United States by Robert K. Godfrey
and Jean W. Wooten. by permission of the University of Georgia Press.
Copyright 1979. 1981 by the University of Georgia Press.
Contents
v
Preface
The first edition of Fassett's Manual of Aquatic Plants ap- Newfoundland. The coverage of Virginia in the first edi-
peared in 1940 and quickly became a classic in its field. tion was necessarily incomplete, and that gap has been
Its success was, in large part, due to Dr. Fassett's aim to closed in this treatment. Fassett's treatment of 752 taxa
provide a manual which would make possible the identi- (plus 95 named forms) has been greatly expanded in this
fication of an aquatic plant in sterile as well as reproduc- edition to include 1186 taxa.
tive conditions. He made an all-out effort to construct his Dr. Fassett's decision to exclude bog species in the first
keys as simply as possible. coupled with flowering or fruit- edition left many aquatic biologists and students in
ing characters where essential for proper identification. northern regions without a comparable guide to the iden-
He copiously cross-referenced characters in his keys to la- tification of species occupying those wet, boggy habitats.
beled illustrations to facilitate the identification process Thus, we have included those species which typically
further. grow in saturated peat. Likewise, we have attempted to
Shortly before his death in 1954, Dr. Fassett invited Dr. meet the needs of aquatic biologists along the Atlantic
Eugene Ogden to prepare a revised edition. This effort, coast by including species of salt marshes and tidal wa-
which appeared in 1957, resulted in a 22-page appendix ters, a study of which began Dr. Fassett's career as a be-
aimed at bringing the nomenclature of the original text ginning student in Professor Fernald's Botany 7 class at
up to date. It also provided a much improved supplemen- Harvard University and later became the focal point of
tary key to the genus Potamogeton. Both Drs. Fassett and his doctoral dissertation.
Ogden readily acknowledged that the addition of an ap- In the original edition Dr. Fassett also included a few of
pendix, though extremely useful, fell far short of the need the more common examples of various groups of algae
for a complete revision. and some common mosses and liverworts. There are ex-
This new edition provides the badly needed updating, cellent references now available for the identification of
yet attempts to retain the features which made Fassett's these groups of plants. We feel that to give adequate at-
work a classic. The region covered by the first edition, tention even to just the aquatic bryophytes would be be-
Minnesota to Missouri, eastward to the Gulf of st. Law- yond the scope of our work.
rence and Virginia, has been extended slightly to include
vii
Permission graciously provided by the New York State
Museum is acknowledged for use of illustrations pub-
lished in several of their bulletins (NYS Museum); Rich-
ard S. Mitchell and E. O. Beal (1979), Magnoliaceae through
Ceratophyllaceae of New York State; Richard S. Mitchell
and J. Kenneth Dean (1978), Polygonaceae (Buckwheat
Family) of New York State; Richard S. Mitchell and J. Ken-
neth Dean (1982), Ranunculaceae (Crowfoot Family) of
New York State; Richard S. Mitchell and Charles J. Sheviak
(1982), Rare Plants of New York State; Eugene C. Ogden
(1981), Field Guide to Northeastern Ferns; and Charles J.
Sheviak (1982), Biosystematic Study of the Spiranthes
cernua Complex.
x Acknowledgments
lum), George W. Argus (Salicaceae), Christopher S. Camp- ported research for Crow each year and the University of
bell (Poaceae), Paul M. Catling (Orchidaceae), Theodore New Hampshire Central University Research Fund sup-
S. Cochrane (Juncaceae, Cyperaceae, and other groups), ported fieldwork in Newfoundland during the summer
C. D. K. Cook (Ranunculaceae, Sparganiaceae), William ofl982.
D. Countryman (Trapaceae), Robert K. Godfrey (Lentibu- Financial support for the botanical illustrations pre-
lariaceae, Utricularia, preliminary checklist of taxa), Rob- pared by Pamela Bruns Brayton was provided by the New
ert R. Haynes (families of the Alismatidae), Hugh H. Iltis Hampshire Agricultural Experiment Station.
(Rubiaceae, Galium), Walter S. Judd (Ericaceae), Emmet J. We are also thankful for support services provided by
Judziewcz (Poaceae), Robert Kral (Cyperaceae, especially the Massachusetts College of Liberal Arts while Hellquist
Rhynchospora, Eriocaulaceae, Melastomataceae, Xyrid- coordinated the compilation of figures from various
aceae), Roger Laushman (Juncaginaceae, Triglochin), sources and prepared the draft copies of the plates. So too
Donald H. Les (Ceratophyllaceae), Richard S. Mitchell are we appreciative of the computing services at the Uni-
(Polygonaceae, preliminary checklist of taxa), C. Thomas versity of New Hampshire.
Philbrick (Callitrichaceae, Podostemaceae), Anthony A. We are especially grateful to Dr. Gordon C. Tucker for
Reznicek (Cyperaceae, especially Carex), George B. Ross- officiating the coin toss which determined the order of ap-
bach (Lentibulariaceae, Utricularia), A. Ernie Schuyler pearance of the authors' names on the title page. Prepa-
(Cyperaceae, especially Scirpus), Charles J. Sheviak (Or- ration of this manual was an equally shared endeavor. the
chidaceae), S. Galen Smith (Cyperaceae, especially Scir- coauthors working over and revising every aspect of the
pus, Typhaceae), Harold St. John (Hydrocharitaceae), manuscript together.
Ronald L. Stuckey (Brassicaceae, preliminary checklist of To our families-Charlyn, Megan, and Jason Crow,
taxa), Janice Coffey Swab (Juncaceae), w. Carl Taylor Lindsay Crow Ackerman, and Marion, Eric, and Paul
(Pteridophytes. especially Isoetes) , Gordon C. Tucker (Cy- Hellquist-we owe a special debt of gratitude for their
peraceae, especially Cyperus, Eleocharis, Kyllinga, Lipocar- love, patience, and endurance during the preparation of
pha, Elatinaceae), Edward G. Voss (preliminary checklist this work. During the early stages of this project they
of taxa), John H. Wiersema (Nymphaeaceae), Jean W. cheerfully participated in two extended botanical expedi-
Wooten (Alismataceae). tions, camping in Newfoundland and enduring the ele-
Support for fieldwork is gratefully acknowledged: the ments of the predictably unpredictable weather of a land
University of Michigan Biological Station supported work we have all come to love. Eric proved to be an exceedingly
for Hellquist during the summers of 1977, 1983, and able field assistant with a keen eye for plants, and has the
1985; the University of Oklahoma Biological Station sup- distinction of finding the first plants of the elusive Curly-
ported work for Hellquist during the summer of 1 981 : grass Fern, Schizaea pusilla, in a raised bog in western
the New Hampshire Agricultural Experiment Station sup- Newfoundland.
Introduction
The aim of this work is to aid in the identification of vas- and species are then arranged in the order in which they
cular plants which are native or have become naturalized key out. Infraspecific taxa considered important from an
in aquatic and wetland habitats of the Northeast. As a aquatic standpoint have been included in the keys. Be-
taxonomic-floristic work. it is designed to be of value to cause of the considerable variation in vegetative morphol-
biologists. students of biology. conservationists. environ- ogy exhibited by many aquatic plants. formal recognition
mental consultants. personnel of local. state. and federal of the rank Jorma has not been applied here. All too often
agencies. and any individuals with general ecological in- a single plant can change from one morphological form
terests. Recognizing that the prospective users will vary to another as the season progresses and water levels
with respect to experience in plant identification. we have change in aquatic and wetland habitats. This morpholog-
attempted to utilize less technical language wherever pos- ical variability has. however. been taken into account in
sible. Glossaries are included to clarify terminology. Illus- constructing the keys; thus some species may key out in
trations are also provided to aid the users; the 606 plates more than one place.
include figures of 1087 taxa. with 92 percent of the taxa The generic descriptions are intended to be diagnostic.
fully or partly illustrated. To facilitate the identification providing additional information which may not be uti-
process further. references to the figures are included in lized in the constrnction of the keys but which may be
the keys. useful in the identification process. We have tried to in-
The geographical range covered by this work includes clude as many illustrations as possible to aid also in the
the region from Newfoundland west along the 50th paral- process of identifying specimens. and have especially at-
lel to southeastern Manitoba and Minnesota. south to Vir- tempted to provide figures that show the important fea-
ginia and Missouri. Thus. on the southern edge. our range tures. Because of spatial constraints we have elected to
abuts the range covered by Drs. Godfrey and Wooten in include only selected family descriptions. because they
their Aquatic and Wetland Plants oj the Southeastern United are readily available in numerous works.
States. We have attempted to utilize vegetative features as
In order to increase the usefulness of our manual we much as possible to facilitate the identification of plants
have expanded our coverage to include plants of wetland in vegetative condition. however. it must be recognized
habitats as well as trnly aquatic habitats. Although we that for many species a reliable determination of the iden-
have included plants of bogs and salt marsh habitats. we tity of a plant will require flowering and/or fruiting mate-
have not included terrestrial plants which may occur typ- rial. This is especially true of some families. such as the
ically on flood plains. in woodland springs. or along ver- Cyperaceae. Orchidaceae. Poaceae. and Sparganiaceae.
nal woodland or seasonal alpine brooks. Admittedly it is In general. it is much more difficult to identify aquatic
sometimes difficult to determine the outer boundary of a and wetland plants early in the growing season. Ideally.
wetland habitat. therefore the decision to include or ex- in inventory work one should be able to make observa-
clude certain species has been necessarily somewhat sub- tions and collect specimens over the entire growing sea-
jective. Our tendency has been toward inclusion rather son. Thus. one can observe the relationship of growth
than exclusion. Some plants which are only occasionally forms in aquatics such as ScIerolepis uniflora whereby the
found in the water may be mentioned in the text but plants remain vegetative in the submersed aquatic form.
omitted from the keys. but as the water level drops, the stems of the aquatic form
The keys in the manual treat a total of 1139 species stranded along the newly exposed shore give rise to up-
(1186 taxa) representing 295 genera in 109 families of right, seemingly unrelated fertile shoots. While the
vascular plants. The families presented follow the classi- aquatic form of this species will key out in Key 3 of the
fication system of Tryon and Tryon (1982) for Pterido- General Keys. keying out the species in the family key of
phytes. the classification of Cronquist. Takhtajan. and the Asteraceae requires fertile material.
Zimmermann (1966) for Gymnosperms. and the system The geographical distributions and habitat information
of Cronquist (1981) for Angiosperms. As in Cronquist's presented for the taxa in this work have been drawn
system. the families of the Dicotyledons (class Magnoliop- widely from the published literature, such as regional
sida) which constitute Volume 1 precede those of the Mo- floras and manuals, monographs and revisions. and flo-
nocotyledons (class Liliopsida) in Volume 2. Families are ristic papers, from herbarium specimens documenting lo-
then arranged by subclasses within the 2 classes. Genera cations not cited in the literature, and from our own field
xi
xii Introduction
experience. Generally. ranges are written from east to junct populations in Nova Scotia. Occurrence in the West
west. then southward to give an overview of the distribu- Indies. Central America. and South America is noted.
tion in North America. For instance. a northern species Taxa introduced into our flora from the Old World are so
may be encountered from Newfoundland westward to noted. However. we have not attempted to include natu-
Alaska and southward to New England. northern New ral or adventive occurrences of our native species in the
Jersey. Pennsylvania. northern Ohio. northern Indiana. Old World.
northern Illinois. Minnesota. Saskatchewan. Montana. In bringing nomenclature up to date we have at-
Idaho. and Washington. Sometimes a northern species tempted to provide a partial listing of some of the more
may also extend southward along the Appalachians as significant synonyms to aid the user in comparing taxa
far as North Carolina. or in the Rocky Mountains south- treated here with those in other botanical references. Us-
ward to Colorado and in the Cascades southward to ers desiring a more comprehensive compilation of syn-
northern California. A species which chiefly occurs on onyms we refer to A Synonymized Checklist of the Vascular
the Atlantic and Gulf coastal plain may range from Flora of the United States. Canada. and Greenland. by Kartesz
southeastern Massachusetts on Cape Cod southward to (1994). Although we have not always followed the syn-
Long Island. New York. southern New Jersey. Delaware. onymy as presented in that work. it remains a valuable
Maryland. Virginia. the Carolinas. Georgia. and Florida. guide to synonymy. Abbreviations of authors of scientific
then westward along the Gulf Coast to Alabama. Missis- names follow those of Fernald (1950).
sippi. Louisiana. and eastern Texas. and not infrequently As a guide to literature pertaining to aquatic and wet-
extend from the Gulf northward along the Mississippi em- land taxa which might be helpful to the user of the man-
bayment as far as southwestern Illinois and southeastern ual. selected references have been provided. Full citations
Missouri. Some coastal plain species also occur as dis- are found in the References section.
xiii
Nuisance Aquatic Plants
of the Northeast
Aquatic plants are becoming an increasingly common quist, 1993) and have a negative impact on the biodiver-
nuisance in lakes. ponds. and streams of the northeastern sity of aquatic habitats of a region which has a much
United States and southeastern Canada. Areas in the greater species richness than previously known (Crow.
northern portion of this manual's range tend to be rela- 1993).
tively free of some of the more severe weed problems af- In the Northeast the genus causing the greatest con-
fecting more southern areas. Yet. fishermen. boaters. cern is Myriophyllum. the Water-milfoils. Three species
swimmers. water-skiers. and landowners find aquatic are of particular concern: M. heterophyllum. M. spicatum.
weeds to be locally problematic when they increase at a and M. aquaticum. Myriophyllum heterophyllum. a mainly
great rate. In some cases weeds eventually cover areas of southern and midwestern species. has become established
open water and prevent its recreational use. and is particularly problematic in many of the acid lakes
Once the weeds become established. various costly and ponds of New England. especially those of eastern
methods may be attempted to eradicate or control the Massachusetts. central New Hampshire. and southern
plants. Some of these methods are: spraying with chemi- Maine. This species typically occurs in waters of higher
cal herbicides. harvesting. dredging. using growth barri- alkalinity in other portions of its range. such as in Michi-
ers. and introducing various natural predators. Some of gan and Oklahoma. In these areas it is usually not as
the natural predators used for biological control are: weedy. Myriophyllum heterophyllum can form extensive
fungi. insects. and the triploid amur. or grass carp. Con- stands. with individual plants becoming extremely ro-
trol methods often require application by licensed profes- bust. It is not certain whether this species is actually na-
sionals and/or state permits. tive in the more northern portion of its range.
Boaters and fishermen who move boats from one body Eurasian Water-milfoil. Myriophyllum spicatum. has
of water to another are often responsible for the introduc- now spread throughout the United States. but only since
tion of plants into new sites. and weedy species most fre- about 1970 has it become a species of concern. Now it is
quently appear for the first time in lakes near public ac- frequently the dominant weedy species in highly alkaline
cess sites. Therefore when a boat is removed from a lake. waters of our region. It has replaced much of the native
it is important for all weeds to be discarded from inside Northern Water-milfoil. Myriophyllum sibiricum (M. exal-
the boat and cleaned from the propeller before moving bescens). Recently it has also shown up in the acid and
the boat to another location. brackish waters of eastern New England. causing con-
Aquatic plants can easily become established in a re- cern regarding its adaptability to a broader range of water
gion. because most of them reproduce both vegetatively alkalinity. It now occurs in acid waters with M. heterophyl-
and by seed (Philbrick and Les. 1996). For some species ev- lum. Throughout the Northeast probably more money is
ery fragment can develop roots and form a new plant. thus spent attempting to control these two species than for the
rapidly building up a population. Many produce winter control of any others. In more southern areas Myriophyl-
buds. or turions. that serve as a means for vegetative lum aquaticum. the Parrot Feather. may be problematic. It
propagation. dispersal. or overwintering without forming is a South American species that can be obtained at pet
seeds. shops and water gardens. and is often used as an attrac-
Although some of our native aquatic species can be- tive shallow-water species for aquatic gardens. It readily
come problematic weeds. many of our nuisance aquatics escapes from cultivation and becomes weedy. It often
have migrated from foreign countries over time and have takes on a terrestrial habit when water levels become low.
become permanently established as components of the thus making it less susceptible to control by drawdowns.
North American flora. and often out-competing the in- Presently it occurs as far north as Long Island. New York.
digenous species (Stuckey. 1993). In studying aquatic but might be expected to spread northward. especially to
and wetland habitats for various management policies northeast coastal regions. where the maritime climate is
and strategies. correct identification of aquatic plants is less harsh in the winter.
very important. Although control of weeds is highly de- Waterweed. Egeria densa. is a tropical and subtropical
sirable. misidentifications can lead to possible inadvertent plant which has become established at a few sites in the
eradication of important or rare indigenous species (Hell- Northeast. This species is known to be a problem in the
xv
xvi Nuisance Aquatic Plants of the Northeast
states of Alabama, Georgia, Florida, Louisiana, North States, with the greatest concentrations in the Midwest.
Carolina, South Carolina, and Oklahoma, and it occurs Only recently has it started moving eastward. It has now
as far north as coastal northeastern Massachusetts. This been found in alkaline waters in western Vermont, Mas-
aquatic plant is commonly sold in the pet trade and by sachusetts, and southeastern New Hampshire (Padgett
biological supply companies, and is erroneously labeled and Crow, 1993a). Since it is a species of alkaline waters,
as Elodea canadensis or Anacharis. and most of New England's waters are acid, further
Hydrilla, Hydrilla verticillata, is a common aquatic plant spreading in the region is likely to be slow. Because this
throughout tropical and subtropical Eurasia and Africa species is an annual, reproducing abundantly by seed, in-
and occurs as far north as boreal Siberia. This species was festations often appear to develop later in the summer.
introduced into the southern United States, southern Cali- During late summer it also fragments readily and may
fornia, and the American tropics. Hydrilla is considered by spread from one body of water to another.
many wetland biologists as potentially the most dreaded The European Curly-leaved Pondweed, Potamogeton
species of aquatic weed in warm temperate areas, such as crispus, has occurred in the United States for many years.
the southeastern United States. Presently it occurs in scat- Documentation in eastern New England goes back over
tered sites as far north as Maryland, Delaware, and south- 100 years. Presently it is found in waters that are highly
ern Connecticut (Les et aI., 1997). This species is similar to polluted or are extremely alkaline (Hellquist, 1980). Pota-
our native Elodea canadensis and is occasionally sold as Elo- mogeton crispus is found in every northeastern state ex-
dea. Every effort must be made to prevent the introduction cept Maine. This species has an interesting life cycle, be-
and spread ofthis species into the northeast region. IfHy- cause it reaches its maximum development rather early
drilla were to become established, it would have the poten- in June in the Northeast. At this time fruits are produced,
tial to be one ofthe most difficult species to eradicate. although fruiting is sparse. By early July winter buds
European Frogbit, Hydrocharis morsus-ranae, was intro- have formed, and they begin to dislodge from the plants
duced into North America in the early 1930s near Ot- as the main portion of the plant dies off. Surprisingly, by
tawa, Ontario, Canada (Catling and Dore, 1982; Roberts, mid-August the winter buds start to develop and small
Stuckey, and Mitchell, 1981). Since then it has become plants appear. These plants remain small and the species
locally established and is a nuisance species in southeast- overwinters in this form. In the following spring the max-
ern Canada and northern New York, and was recently imum growth occurs, leading to the robust plants seen
discovered in Lake Champlain, Vermont. Frogbit is ag- in June.
gressive in its vegetative reproduction and overwinters by The Water-chestnut, Trapa natans, native to Asia, is a
forming winter buds. It has been theorized by some that rooted annual aquatic. Upon reaching the surface, plants
the advent and spread of Frogbit in the Northeast prob- spread rapidly by the production of many floating ro-
ably arose from its introduction by sportsmen's clubs to settes, which can choke out large areas of open water.
provide food and cover for waterfowl. However, as with This species has been established in the Northeast for
many aquatic plants, it may have inadvertently been in- many years. Presently it is known in the Champlain Val-
troduced by people dumping out the contents of aquaria ley of Vermont and New York, the Mohawk and Hudson
into a local body of water. river systems of New York, throughout Massachusetts,
One of the most difficult weeds to eradicate in the the Connecticut River in Connecticut, and the Chesa-
Northeast is Fanwort, Cabomba caroliniana. This species is peake Bay region (Trudeau, 1982). In 1998 it was dis-
native to the southern United States. However, it is a very covered for the first time in the Nashua River in southern
popular aquarium plant and can be purchased at almost New Hampshire (pers. comm., Kenneth Warren). Al-
any pet store or water garden in the Northeast. Cabomba though this species can occupy very large areas, it is one
is hardy and extremely aggressive. Once established in a that can be effectively controlled by persistent mechanical
pond it is almost impossible to eradicate. It is resistant to harvesting. Since the plant is an annual, the floating ro-
various herbicides and is not a favored food of the amur, settes should be harvested early in the summer before the
or grass carp, a fish species often used to control aquatic plants produce their large caltrop nuts. This species can
weeds. This species is very common in acid-water lakes of rival the tropical Water-hyacinth, Eichhornia crassipes, in
eastern New England (Hellquist and Crow, 1986). In the its ability to choke out a water surface.
Northeast the Fanwort probably ranks second to the One species of Water-fern, Marsilea quadrifolia, occa-
Water-milfoils for money spent on weed control. This spe- sionally becomes aggressive. There is some question as to
cies is prohibited from import into Canada. whether this Eurasian species might also be native in
The Minor Naiad, Najas minor, was discovered in the some areas of North America. It is extremely hardy when
Hudson River, New York, in 1934 (Meriliiinen, 1968). compared with the many tropical species that may be
Since its discovery, this species has spread quickly west- purchased, such as Marsilea mutica from Australia. Care
ward, occurring widely throughout much of the United should be taken not to introduce any species of Marsilea.
Nuisance Aquatic Plants of the Northeast xvii
The subtropical species planted in a shallow pond can Yellow Waterlilies, Nuphar spp., can all spread and be-
grow during the summer at a phenomenal rate-up to a come weedy. Pickerel-weed, Pontederia cordata, and Wild-
foot a week. Marsilea quadrifolia grows at a much slower rice, Zizania spp., often form near monocultures in shal-
rate, but still can become weedy. low water. Equisetum fluviatile, Water Horsetail, readily
The Yellow Floating-heart, Nymphoides peltata, is an- invades and colonizes shallow waters. Ceratophyllum de-
other aggressive Eurasian aquatic that is not yet abun- mersum, Coon tail, a free-floating submersed species, also
dant in the northeast region, but has the potential to be- often chokes out the middle and upper layers of a body
come a real nuisance. The earliest known herbarium of water.
specimen for this species was collected in 1882 from east- The various native cattails, Typha spp., can also become
ern Massachusetts (Stuckey, 1974). This species has the invasive in a wetland or along the margins of lakes and
potential to colonize an area quickly. The plants produce ponds. Care should be exercised not to introduce any
many blossoms, which, in turn, produce many seeds. plants to a new body of water. A potential problem is on
Plants also spread rapidly by fast-growing shallow rhi- the increase. Various water-garden nurseries throughout
zomes. Although typically a plant of warmer regions, it is the country are offering exotic species of cattails for sale.
hardy even in the Northeast. More concern should be These various species are all hardy and may lead to weed
paid to this species. problems in the future. They include: Typha minima, T.
A common weedy marsh plant throughout the north- laxmannii, and T. gracilis.
eastern United States is Purple Loosestrife, Lythrum salica- Probably one of the most invasive wetland plants in the
ria. This tall aggressive Eurasian marsh plant was re- Northeast is the Reed, Phragmites australis, a plant with a
ported adventive as early as 1824, growing in meadows nearly worldwide distribution. This large grass (often up
in New England and Canada (Stuckey, 1980a). This spe- to 4 meters tall) is extremely aggressive and readily colo-
cies very likely escaped from flower gardens, because it nizes almost any area where drainage has been impeded.
has been grown for many years as an attractive orna- Roadside ditches are common places to observe this spe-
mental. Purple Loosestrife may presently be purchased at cies. Probably the largest population of this species occurs
many retail nurseries. However, it is best not to purchase in the northern New Jersey meadowlands just west of
this plant unless you are sure it is a sterile hybrid. But New York City. Eradication of this species should take
even these hybrids have the ability to spread vigorously place when new sites are discovered, because it will
from rhizomes. Throughout the Northeast, Purple Loose- quickly outcompete the present vegetation.
strife has colonized hundreds of acres of land, especially Care should be taken in planting the showy Lotus spe-
seasonally wet sites that tend to dry by late summer. The cies, both our native Yellow Water Lotus, NeIumbo lutea,
attractive magenta-purple flower produces a spectacular and the Indian Lotus, N. nucifera. There are large aggres-
display during midsummer. Of greatest concern is the sive populations of both species in scattered localities in
tendency of this species to crowd out and replace much the East. They often are planted for horticultural pur-
of the native vegetation. poses because of the large showy colorful blossoms.
The Flowering-rush, Butomus umbellatus, has become Three tropical and subtropical weedy species are: Alli-
well established along the st. Lawrence River drainage gator Weed, Alternanthera philoxeroides, Water Lettuce,
and the shores of Lake Champlain. At these locations it Pistia stratiotes, and Water-hyacinth, Eichhornia crassipes
has become a major component of the emerged shoreline (Aurand, 1982). These species are presently reported in
and marsh vegetation. Since about 1970 this species has our range from southeastern Virginia, where they are
been spreading westward, and may now be found at most likely short-lived. Although they are very problem-
widely scattered locations across the northern United atic in the South, these should not become a problem in
States and southern Canada. It appears that some popu- the Northeast.
lations have been introduced from Europe, whereas oth- There are various other exotic species that have the po-
ers have had an Asian origin. tential to become a nuisance. Some of these are: Money-
Although many of our aquatic weeds are aliens, there wort, Lysimachia nummularia, Water Cress, Nasturtium of-
is a potential for many native aquatic plants in the North- ficinale, Great Hairy Willow Herb, Epilobium hirsutum, and
east to become aggressive if introduced into new areas. Forget-me-not, Myosotis scorpioides.
They may also become more weedy if present aquatic A major source of introductions of non-native wetland
habitats become greatly disturbed. Such plants include plants is in the replication of wetlands. Manmade wet-
Waterweed, Elodea canadensis and E. nuttallii, and various lands are recommended or required in many states when
species of Potamogeton, for example, Potamogeton illi- an existing wetland is destroyed or a portion of a wetland
noensis, P. nodosus, and the various narrow-leaved Pota- is filled. Often the plants that are purchased for the newly
mogetons. Likewise, the White Waterlilies Nymphaea odor- constructed wetland are obtained from nurseries in the
ata ssp. odorata and N. odorata spp. tuberosa, and the western or Plains states. This creates a philosophical
xviii Nuisance Aquatic Plants of the Northeast
xix
Key 1 Plants woody, trees, shrubs, or subshrubs
24. Leaves strictly opposite; flowers and fruits in axillary pairs or in much-branched
cymes; fruit succulent, a berry or drupe . . . . . . . . . . . . . . . . . . . . . Caprifoliaceae, 1:372
16. Leaves alternate.
26. Plants viny, often climbing, becoming woody toward base, herbaceous toward tip (Solanum) . . . . . . . . Solanaceae, 1;276
26. Plants trees, shrubs, or subshrubs.
27. Leaves lobed; bark of older stems peeling in strips (Physocarpus) . . . . . . . . . . . . . . . . . . . . . . . . Rosaceae, 1:179
27. Leaves not lobed; bark not peeling.
28. Leaf margins entire or very minutely serrulate or crenulate so as to appear entire.
29. Trees; stem pith diaphragmed (with transverse septae) in longitudinal section.
30. Leaves leathery, with stipules, somewhat evergreen (especially southward); flowers very large,
showy, petals 6-9; fruit an aggregate of follicles (Magnolia) . . . . . . . . . . . . . . . . Magnoliaceae, 1:35
30. Leaves not leathery, lacking stipules, deciduous; flowers small, petals 5 or often absent:
fruit a drupe (Nyssa) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nyssaceae, 1:230
29. Trees, shrubs, or subshrubs; stem pith continuous (not diaphragmed) in longitudinal section.
31. Leaves mucronate at apex (margins sometimes very minutely toothed, yet appearing entire),
glabrous; corolla of separate petals; fruit a red drupe, rarely yellow, with 4 or 5 seed-like nutlets
(Nemopanthus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aquifoliaceae, 1:237
31. Leaves acute to obuse or rounded at apex, but not mucronate, either glabrous or pubescent;
corolla united (if petals separate, then densely tomentose beneath, Ledum); fruit a capsule, a
red, blue to black, many-seeded berry or a black drupe with 10 seed-like nutlets .... Ericaceae, 1:158
28. Leaf margins serrate, dentate, or crenate.
32. Leaves at least 3 times as long as wide; lateral buds covered by a single bud scale (Salix). Salicaceae, 1:129
32. Leaves less than twice as long as wide; lateral buds covered by 2 or more bud scales.
33. Flowers greatly reduced, borne in catkins (fig. li,j).
34. Plants monoecious; fruit a nutlet; pistillate catkins persisting; trees or shrubs ... Betulaceae, 1: 72
34. Plants dioecious; fruit a capsule; pistillate catkins falling off early; trees (Populus) Salicaceae, 1: 129
33. Flowers not reduced, borne variously but not in catkins.
35. Leaf bases rounded, usually oblique (Planera) . . . . . . . . . . . . . . . . . . . . . . . . . Ulmaceae, 1:68
35. Leaf bases tapering, acute, or if occasionally rounded, then not oblique.
36. Flowers with hypanthium present (free or adnate to ovary); stamens 15 or more;
fruit a cluster of follicles (Spiraea) or a pome (Aronia) . . . . . . . . . . . . . . .. Rosaceae, 1: 179
36. Flowers lacking hypanthium; stamens 10 or fewer; fruit a capsule, berry, or drupe.
37. Twigs with dense stellate pubescence; fruit a 3-valved capsule (Clethra) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Clethraceae, 1:153
37. Twigs glabrous or pubescent, but hairs not stellate; fruit a 5-valved capsule,
berry, or drupe.
38. Corolla united, urceolate; fruit a capsule or a many-seeded berry (Leu-
cothoe, Lyonia, Vaccinium) . . . . . . . . . . . . . . . . . . . . . . . . . . . Ericaceae, 1:158
38. Corolla only slightly united at base (petals appearing separate) or absent;
fruit a 3-10-seeded drupe.
39. Corolla present; drupes with 5-10 seeds (stones), reddish, or if black,
then leaves evergreen and leathery (Ilex) . . . . . . . . . . . Aquifoliaceae, 1:237
39. Corolla absent; drupes with 3 seeds (stones), black (Rhamnus) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhamnaceae, 1:243
f
Fig. 1. a. Nyssa sylvatica, portion of branch (F); b. Larix laricina, portion of branch (Sargent); c. Taxodiurn
distich urn, branchlet (deciduous as a unit) (F); d. CephaIanthus, opposite leaves (whorled not shown) (F);
e. Thuja occidentalis, portion of branch (Sargent); f. Fraxinus, compound leaf (F); g. Forestiera, opposite
leaves (F); h. Alnus incana, leaf (F); i. Alnus, staminate catkins (F); j. Alnus, pistillate catkins (F).
xxii
Key 2 Plants free-floating on the surface
1. Plants with rosette of inflated branches sub tending flower scapes (fig. 2i); bladders present on submersed portion of plant (Utricularia)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lentibulariaceae, 1:352
1. Plants lacking rosette of inflated branches (petioles appearing inflated in Eichhornia, fig. 2b, and Trapa, fig. 2c); bladders absent.
2. Plants dichotomously forked or lobed.
3. Plants with overlapping leaves, upper surface velvety (fig. 31a,f) (AzolIa) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salviniaceae, 1:14
3. Plants lacking leaves, thalloid, upper surface glabrous (Riccia, fig. 2g, Ricciocarpus, fig. 2h, non-vascular plants, a family
of liverworts) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ricciaceae
2. Plants not dichotomously forked.
4. Plants not distinctly differentiated into stems and leaves, small to minute (fig. 2d,e,f,k) . . . . . . . . . . . . . . . . . . . . . . Lemnaceae, 2:78
4. Plants differentiated into stems and leaves, small or larger.
S. Plants small, not forming rosettes; leaves lacking petioles, 0.5-15 mm long; plants reproducing by spores borne in
sporocarps.
6. Leaves appearing opposite (actually whorled, 2 leaves floating, 1 submersed), not overlapping, upper surface covered with
erect forked hairs (fig. 29g) (Salvinia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Salviniaceae, 1:14
6. Leaves alternate, overlapping, upper surface velvety (fig. 30a) (AzolIa) . . . . . . . . . . . . . . . . . . . . . . . . . . . Salviniaceae, 1:14
S. Plants larger, forming rosettes; leaves petiolate (fig. 2a,b,c,j), blades more than 20 mm long; plants reproducing by flowers
and seeds (detached Nymphoides, fig. 2a, and Trapa, fig. 2c, normally rooted, might key out here).
7. Leaves with petioles appearing greatly inflated (fig. 2b), blades not spongy; flowers showy, bisexual (Eichhornia) ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Pontederiaceae, 2:312
7. Leaves with petioles not inflated, but blades spongy (fig. 2j); flowers not showy, unisexual (Limnobium, Hydrocharis) .. ,
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Hydrocharitaceae, 2: 11
xxiii
a
Fig. 2. a. Nymphoides, upper portion of plant (F); b. Eichhornia crassipes, habit (Beal); c. Trapa, upper portion of
plant (NHAES); d, e. Wolffia, habit (F); f. Wolffiella, habit (F); g. Riccia, habit (F); h. Ricciocarpus, habit
(F); i. Utricularia radiata, upper portion of plant (F); j. Limnobium, floating plant (F); k. Lemna, habit (F).
xxiv
Key 3 Stems and leaves, or their divisions, limp, thread-like or ribbon-like, more than 10 times as long as wide
xxv
xxvi Key 3
stem
a
Fig. 3. a. Scattered simple thread-like leaves on a branched stem, generalized; b. CeratophyIlum, leaf; c. flat,
opposite, ribbon-like leaves on stem, generalized; d. Ranunculus longirostris; e. Isoetes, generalized;
f. ribbon-like leaves arising from one point, generalized; g. thread-like leaves or stems arising in clusters
along a rhizome (F).
xxvii
bladder
free} stipule
stem
Fig. 4. a. Potamogeton, diagrammatic; b. Nt:c]'m;; leaf b~ses generalized; c. Zannichellia, slender leaf bases, gener-
alized; d. Megalodonta, section of stem; e. Ranuncu!us, sheathing leaf bases, generalized; f. Cabomba;
g. Utricularia, generalized; h. Stuckenia pectinata, leaf diagrammatic; i. Myriophyllum, generalized; j. Rup-
pia, leaf, generalized; k. Neobeckia, section of stem with leaf (F).
xxviii
e
sheath
b c d f
II
11M-
Fig. 5. Sections of leaves. about natural size: a. Poaceae (grasses). generalized; b. Vallisneria; c. Sagitta ria;
d. Sagittaria; e. Pontederia cordata. submersed form; f. Sparganium. generalized (F).
xxix
d
b stipule
Fig. 6. a. Sparganium. leaf. generalized; b-d. Potamogeton. leaves. each with 2 stipules at base; e. Zosterella dubia.
leaf with 2 stipules at base; f. Najas. axillary leaf fascicles; g. portion of plant with scattered ribbon-like
leaves. diagrammatic; h. Didiplis diandra. aquatic form. section of stem with leaves (F).
xxx
Key 4 Leaves compound, cut into several Oat IeaOets
1. Plants distinctly fern-like. reproducing by spores (spores never in sporocarps) (figs. 8a. 22).
2. Sterile and fertile fronds similar. sporangia borne on lower surface of frond.
3. Fronds from upright rhizomes (appearing clumped) (Dryopteris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dryopteridaceae. 1:3
3. Fronds from creeping rhizomes.
4. Sori round. sparsely or densely spaced along center of pinnules (fig. 22d); rhizome 1-2 mm in diameter; stipes black. lacking
scales (The1ypteris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thelypteridaceae. 1:3
4. Sori elongate. appearing continuous along midrib of pinnae and pinnules (fig. 8a); rhizome 6-10 mm in diameter; stipes
purplish-brown. with scales. deciduous at base (Woodwardia virginica) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Blechnaceae. 1:8
2. Sterile and fertile fronds different (fig. 20b.c). or fronds divided into sterile and fertile portions (fig. 19) (Osmunda) ..... Osmundaceae. 1:3
1. Plants not fern-like. reproducing by flowers and seeds. or by spores borne in sporocarps.
5. Leaves compound. with 4 leaflets. cross-shaped (fig. 29a.b) (similar to 4-leaved clover); plants reproducing by spores borne in
sporocarps (Marsilea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Marsileaceae. 1:8
5. Leaves various. but not with 4 leaflets; plants reproducing by flowers and seeds.
6. Plants not rooted. usually free-floating below surface in tangled mats; plants thalloid. consisting of branched. long-stalked
fronds. giving compound appearance (fig. 402f.g) (Lemna trisulca) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lemnaceae.2:78
6. Plants rooted. not free-floating; plants with distinct stems and leaves.
7. Divisions ofleaves fine. less than 1 mm wide (fig. 7a.c).
8. Leaves crowded at upper portion of stem (figs. 7a. 140a) (Hottonia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Primulaceae.l:168
8. Leaves scattered along most of stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haloragaceae.l:194
7. Divisions of leaves broad. greater than 1 mm wide (fig. 7d-i). or if less than 1 mm. then more than once compound
(figs. 205b.d. 206b).
9. Leaves opposite (Bidens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asteraceae.l:375
9. Leaves alternate.
10. Leaves with stipules (figs. 152b. 153f). leaflets toothed; sepals. petals. and stamens borne on a shallow hypan-
thium (Geum. PotentilIa. Sanguisorba) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rosaceae. 1:179
10. Leaves lacking stipules. or if present (Fabaceae). then leaflets entire; sepals. petals. and stamens not borne on
a hypanthium.
11. Base of petiole expanded into a sheath (fig. 7d.e).
12. Stem completely encircled by base of petiole (figs. 202a. 21Oa) . . . . . . . . . . . . . . . . . . . . . Apiaceae. 1:245
12. Stem not encircled by base of petiole.
13. Leaves pinnately compound (fig. 227b).
14. Leaflets usually 15 or more; corolla blue; stamens 5 (Polemonium) ......... Polemoniaceae.l:285
14. Leaflets usually 5; corolla white; stamens 3 (Valeriana) . . . . . . . . . . . . . . . . . Valerianaceae.l:375
13. Leaves palmately compound (figs. 64d. 223a).
15. Corolla white. united. lobes conspicuously fringed; flowers bisexual. with 1 pistil and 5
stamens (Menyanthes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Menyanthaceae. 1:276
IS. Corolla yellow. not united. petals never fringed. or corolla absent; flowers bisexual or unisex-
ual. with several to many pistils and/or stamens . . . . . . . . . . . . . . . . . . . . . Ranunculaceae. 1:50
11. Base of petiole not expanded into a sheath.
16. Flowers borne in dense heads. subtended by involucral bracts (Eupatorium) ............ Asteraceae. 1:375
16. Flowers borne in few- to many-flowered racemes.
17. Flowers radially symmetrical. 4-merous. cross-shaped; fruit a silique; terminal leaflet usually the
largest (fig. 7h) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brassicaceae. 1:143
17. Flowers bilaterally symmetrical. 5-merous; fruit a legume (fig. 154d) or loment (fig. 154e); termi-
nalleaflet about the same size as lateral leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . .. Fabaceae. 1: 191
xxxi
Fig. 7. a. Hottonia, crowded feather-like leaves; b. Proserpinaca, leaf variation on a single plant; c. Myriophyllum,
generalized, with 2 leaves scattered and 4 whorled leaves; d. Sium, emersed leaf; e. Sium, submersed
leaf; f. Bidens frondosus, leaf; g. Potentilla palustris, leaf with stipule adnate to petiole; h. Nasturtium, leaf;
i. Rorippa palustris, leaf (F).
xxxii
Key 5 Leaves simple, deeply lobed, or basal lobes extending laterally or forward
xxxiii
b
Fig. 8. a. Woodwardia virginica. portion offrond (NYS Museum); b. c. Neobeckia. submersed leaves (F); d. Ranun-
culus. leaf. generalized (F); e. Ranunculus sceleratus; leaf (F); f. Hydrocotyle. leaf. generalized (F); g. Rorippa
palustris. basal leaves (F); h. Bidens connatus. leaf (F); i. Onoc1ea. leaf and fertile fronds (NYS Museum);
j. Proserpinacapalustris. habit (F).
xxxiv
Key 6 Leaves with basal lobes extending below junction of blade and petiole, or leaves peltate
xxxv
xxxvi Key 6
21. Petals violet-blue, in a dense, erect spike; fruit a utricle, crested with toothed ridges
(Pontederia) .......................................... " Pontederiaceae, 2: 312
21. Petals white, in a dense nodding spike-like raceme, or in whorls on a raceme; fruit a globose
cluster of 3-5 nutlets, or numerous achenes.
22. Leaves net-veined, with a midvein; flowers in a dense spike-like raceme, nodding at tip
(fig. 43a); fruit a globose cluster of 3-5 nutlets (Saururus) ............. Saururaceae, 1:35
22. Leaves parallel-veined, lacking a midvein; flowers in whorls in a raceme; fruit a globose
cluster of numerous achenes (Echinodorus cordifolius) ................ Alismataceae, 2:3
Fig. 9. a. Nelumbo. portion of peitate leaf; b. Heteranthera reni/ormis; c. Pontederia cordata; d. Limnobium. leaf; e.
Calla. leaf; f. Nymphoides. leaf; g. Hibiscus laevis. hastate leaf; h. Caltha. leaf; i. Nuphar; leaf; j. Sagitta ria
lati/olia; k. Saururus cernuus. leaf (F).
xxxvii
Key 7 Leaves basal or cauline, more than 10 times as long as wide; or stems naked or appearing naked, with
leaves reduced to bladeless sheaths
1. Stems (root-like) in soil with tiny bladders; upright stems leafless. with yellow or purple spurred flowers (fig. 289a) (Utricularia)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lentibulariaceae. 1:352
1. Stems or roots in soil lacking bladders; upright stems naked or with leaves. flowers various.
2. Leaves greatly reduced. appearing as scales (figs. 74b.d. 215a.f) or bumps (fig. 100 along stems. stems thus appearing naked.
3. Plants distinctly succulent; plants of saline sites. usually coastal (Salicornia) . . . . . . . . . . . . . . . . . . . . . . . . . Chenopodiaceae. 1:79
3. Plants not succulent; plants of non-saline sites.
4. Stems several. arising along a creeping rhizome; leaves rounded at apex. appearing as minute bumps (fig. 100 (Myriophyllum
tenellum) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haloragaceae. 1: 194
4. Stems single; leaves acute at apex.
5. Flowers 3-merous. perianth tube 3-angled or 3-winged; capsule 3-winged (fig. 592i) (Burmannia) . . . . . . Burmanniaceae. 2:328
5. Flowers 4-merous. perianth tube round; capsule round in cross-section (Bartonia) . . . . . . . . . . . . . . . . . Gentianaceae. 1:267
2. Leaves not reduced to scales or bumps along stems (although sometimes reduced to bladeless sheaths).
6. Leaves basal (fig. lla.e.g); or stems naked (or appearing naked). all arising from one point (figs. lOa. l3b). or arising scattered
along a rhizome (fig. 12b).
7. Plants covered with conspicuous. sticky glandular hairs (fig. 108c) (Droserafiliformis) . . . . . . . . . . . . . . . . . . Droseraceae. 1:120
7. Plants lacking glandular hairs.
8. Leaves or naked stems usually 15 cm or more long.
9. Erect structures (usually leaves) flat in cross-section.
10. Base of plant triangular in cross-section (sterile plants) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae. 2:121
10. Base of plant not triangular.
11. Leaf with a ligule at junction of blade and sheath (fig. llb) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poaceae.2:252
11. Leaf without a ligule. sheath present or absent.
12. Flowers bilaterally symmetrical. with 1 larger petal forming a distinct lip or labellum (figs. 593b. 595a);
stamens and pistils united. modified to form a column (figs. 602d. 605f) ............. Orchidaceae.2:328
12. Flowers radially symmetrical. without a lip; stamens and pistils appearing normal. not specialized.
13. Leaf with a keeled center (fig. llf.h.i). or a thick. raised midrib.
14. Leaf base. when crushed. with an aromatic odor; flowers and fruits borne in a spadix. ap-
pearing lateral on the leaf-like axis (fig. 398a.b) (Acorus) . . . . . . . . . . . . . . . . . . . Acoraceae. 2:76
14. Leaf base. when crushed. lacking an aromatic odor; flowers and fruits borne in globose heads
(fig. 569) (Sparganium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparganiaceae. 2:296
13. Leaf lacking a keeled center or a raised midrib.
15. Junction of sheath and blade abrupt. summit of sheath truncate (fig. llc) (Typha domingensis.
sheath tapering to blade. but plants usually 2-3 m tall. fig. lId); flowers borne in a dense
cylindrical spike (Typha) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Typhaceae.2:306
15. Junction of sheath and blade transitional. sheath gradually tapering to blade. or lacking;
flowers not borne in a dense. cylindrical spike.
16. Perianth wooly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haemodoraceae. 2:317
16. Perianth glabrous.
17. Leaves 5 mm or more wide.
18. Plants with a conspicuous rhizome; flowers 6 cm or more wide (Iris)
.................................................. Iridaceae. 2:323
18. Plants tufted. lacking a rhizome (fig. lIe); flowers 1 cm or less wide (Xyris) ..
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Xyridaceae.2:85
17. Leaves less than 5 mm wide.
19. Leaves arising from a rhizome.
20. Flowers borne in an umbel.
21. Flowers pink. carpels 6. separate; stalk of inflorescence tall. up to 1.5
m; rhizome short. thick (fig. lla) (Butomus) . . . . . . . . . . . Butomaceae.2:3
21. Flowers white. carpels 2. united; stalk of inflorescence short (near the
ground). 3-10 mm; rhizome elongate. slender (Lilaeopsis) ... Apiaceae. 1:245
20. Flowers borne in a raceme or spike . . . . . . . . . . . . . . . . . . . . Liliaceae.2:320
19. Leaves arising in a dense tuft. lacking a rhizome (fig. lle). base
sometimes bulbous (Xyris) . . . . . . . . . . . . . . . . . . . . Xyridaceae.2:85
9. Erect structures (leaves or stems) round to oval. U-shaped. square. or triangular in cross-section.
22. Ascending structures consisting of leaves.
23. Leaves triangular in cross-section; flowers and fruits borne in globose heads (fig. 569) (Sparganium)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sparganiaceae. 2:296
23. Leaves round to oval or U-shaped in cross-section. or tubular (fig. 107b); flowers solitary or borne in a spike.
or if flowers not produced. then plants reproducing by spores in sporangia embedded in leaf bases.
24. Plants with a hard. bulbous base (fig. 12e); plants reproducing by spores borne in sporangia embedded
in leaf bases (fig. 36b) (Isoetes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isoetaceae.1:25
xxxviii
Key 7 xxxix
24. Plants lacking a hard. bulbous base; plants reproducing by flowers and seeds.
25. Leaves tubular. open at upper end (water accumulating in tubular portion). not succulent (fig.
107b) (Sarracenia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sarraceniaceae. 1:120
25. Leaves not tubular. somewhat succulent.
26. Leaves sheathing at base (fig. 367b); pistils 3-6 (Triglochin) .............. Juncaginaceae. 2:33
26. Leaves clasping at very base. but not sheathing; pistils 1 (Plantago) ........ Plantaginaceae. 1:323
22. Ascending structures consisting of stems (usually with basal. bladeless sheaths).
27. Plants vegetative.
28. Stems 1 m or more long (Scirpus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae. 2:121
28. Stems less than 1 m long.
29. Stems 3-angled or 4-8-angled in cross-section.
30. Stems 3-angled (Scirpus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae, 2:121
30. Stems 4-8-angled (Eleocharis elIiptica. E. quadrangulata. E. tenuis. E. verrucosa) . .... Cyperaceae. 2: 121
29. Stems nearly round in cross-section.
3l. Sheaths open, edges often overlapping, but not united to form a tube around stem (fig. 416d)
(funcus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juncaceae, 2:97
3l. Sheaths closed, forming a tube around stem (figs. 496d. 516a).
32. Stems single or few, arising at intervals along a rhizome (figs 10c. 12b. 494b) (Eleocharis)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae.2:121
32. Stems tufted, lacking a creeping rhizome (Scirpus. Eleocharis; these genera are difficult to
distinguish in vegetative condition) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae.2:121
27. Plants with flowers/fruits. borne at or near tips of stems (figs. 10a,g. 13b,d.f).
33. Flowers reduced. hidden by overlapping scales (fig. 498b); perianth absent (fig. 498f.g) or reduced to
bristles (figs. 498c.e, 521d.h); fruit an achene.
34. Inflorescence (spikelet) terminal. with no involucral bract extending beyond (fig. 13a.i).
35. Achene with a tubercle formed by the persistent style base (fig. 490) (Eleocharis) ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... Cyperaceae. 2: 121
35. Achene lacking a tubercle (involucral bract so short as to appear absent; Scirpus cespitosus,
S. rufus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae. 2:121
34. Inflorescence appearing somewhat lateral, with a single involucral bract extending beyond (fig.
l3d,f) (Scirpus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae, 2:121
33. Flowers small, but readily visible; perianth parts 6, in 2 whorls, scale-like, brownish (fig. l3e,g); fruit
a capsule (funcus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .... Juncaceae. 2:97
8. Leaves or naked stems less than 15 cm long.
36. Flowers irregular, with 1 large petal forming a distinct lip (fig. 606b); stamens and pistil united. modified to form a
column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orchidaceae, 2:328
36. Flowers regular or irregular; stamens and pistils appearing normal, not specialized; or plants without flowers, repro-
ducing by spores.
37. Leaves in cross-section appearing like 2 united tubes (fig. 12d) (Lobelia dortmanna) .......... Campanulaceae, 1:365
37. Leaves in cross-section not appearing as 2 united tubes.
38. Plants with a bulbous. hard base (figs. 12e. 405a).
39. Leaves flat, 2-ranked; plants reproducing by flowers and seeds (Xyris torta) ............ Xyridaceae, 2:85
39. Leaves round in cross-section, not 2-ranked (fig. 12e); plants reproducing by spores borne in sporangia
embedded in leaf bases (fig. 36b) (Isoetes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isoetaceae, 1:25
38. Plants lacking a bulbous, hard base.
40. Leaves 10 mm or more wide at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alismataceae, 2:3
40. Leaves less than 10 mm wide at base.
4l. Tufts of leaves connected by slender rhizomes or runners (figs. 10c, 12fJ.
42. Erect green leaves or stems of essentially the same width throughout, or somewhat expanded
into a narrow blade toward apex (fig. 18e).
43. Tufts ofleaves connected by green runners (fig. lSe); petals yellow (Ranunculus reptans)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ranunculaceae, 1:50
43. Tufts of leaves, solitary leaves, or stems connected by rhizomes (often white); petals
white to purplish, or absent.
44. Leaves usually borne singly along a rhizome (fig. 199c) (Lilaeopsis) ...... Apiaceae, 1:245
44. Leaves or stems in clumps along a rhizome.
45. Erect green structures (leaves or stems) pointed at apex; flowers borne in
spikes. racemes, or whorled clusters.
46. Erect green structures stems; flowers in terminal spikes, petals lacking
(Eleocharis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae, 2:121
46. Erect green structures leaves; flowers in racemes or whorled clusters. pet-
als white.
47. Roots septate (fig. 34Sb); pistils numerous; fruit a globose head of
achenes (fig. 34Sd) (Sagittaria) . . . . . . . . . . . . . . . . . . . . Alismataceae. 2:3
47. Roots not septate; pistils 1; fruit a capsule (Tofieldia) . ...... Liliaceae. 2:320
45. Erect green structures (leaves) rounded at apex; flowers solitary, on recurved
stalks (Limosella) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scrophulariaceae, 1:327
42. Erect green leaves or stems widest at base, tapering toward apex.
48. Leaves with minute cross-lines when held to light (hand lens needed); flowers in spikes
or clusters, bisexual.
49. Sheaths open, edges overlapping but not united to form a tube around stem; flow-
ers visible, with perianth of 3 scale-like sepals and 3 scale-like petals (fig. 13g);
fruit a capsule (funeus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Juncaceae, 2:97
49. Sheaths closed, forming a tube around stem; flowers hidden by overlapping scales,
with perianth absent or reduced to slender bristles, petals lacking; fruit an achene
(Eleoeharis, especially E. acieularis) . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae, 2:121
48. Leaves lacking cross-lines when held to light; flowers solitary, unisexual (Littorella) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plantaginaceae, 1:323
41. Tufts of leaves not connected by slender rhizomes.
50. Leaves dimorphic; vegetative leaves short, curly, grass-like; fertile leaves erect, pinnatifid,
appearing I-sided (fig. 21); plants reproducing by spores (SehizaeapusilIa) ...... Schizaeaceae, 1:3
50. Leaves all similar; plants reproducing by flowers and seeds.
51. Leaves slightly to decidedly succulent.
52. Leaves sheathing at base; pistils 3-6 (Trigloehin) . . . . . . . . . . . . . . Juncaginaceae, 2:33
52. Leaves clasping at very base, but not sheathing; pistils 1 (Plantago maritima) ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plantaginaceae, 1:323
51. Leaves not succulent.
53. Leaves widest at base and tapering toward apex.
54. Roots septate (fig. 12i) (Erioeaulon) . . . . . . . . . . . . . . . . . . . . . Eriocaulaceae, 2:92
54. Roots not septate.
55. Base of plant with persistent old leaves; flowers numerous, borne in a
dense, button-like head (LaehnoeauIon) . . . . . . . . . . . . . . . Eriocaulaceae, 2:92
55. Base of plant lacking persistent old leaves; flowers few, borne in a raceme
(fig. 12h) (Subularia) . . . . . . . . . . . . . . . . . . . . . . . . . . . Brassicaceae, 1:143
53. Leaves widest near or above middle.
56. Leaves broadly rounded at apex (fig. 12j); flowers solitary; pistils and stamens
numerous (Myosurus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ranunculaceae, 1:50
56. Leaves pointed (sometimes slightly twisted) at apex (fig. lIe); flowers borne in
a capitate spike (fig. lle); pistils 1, stamens 3 (Xyris) ........... Xyridaceae, 2:85
6. Leaves scattered along stem; stems never appearing naked.
57. Leaves whorled, (2)3-12 per node.
58. Leaves (2) 3 or 4 per whorl; stems not hollow (small lacunar canals present).
59. Leaves (2) 3 per whorl, not tipped by a tiny callus (Elodea) . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrocharitaceae, 2:11
59. Leaves (3) 4-6 per whorl, tipped by a tiny callus (Sc1erolepis) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asteraceae, 1: 3 75
58. Leaves 6-12 per whorl; stems hollow (Hippuris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hippuridaceae, 1:315
57. Leaves alternate or opposite.
60. Flowers greatly reduced, hidden in axils of overlapping scales, forming spikes or spikelets; perianth absent or reduced
to bristles.
61. Stems solid (except Duliehium, hollow including nodes; Scirpus eyperinus, sometimes with pith breaking down),
triangular in cross-section, angles sometimes rounded, but pith angular; leaves 3-ranked; sheaths closed; flowers/
fruits each subtended by a single bract (fig. 432) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cyperaceae, 2:121
61. Stems hollow, solid at nodes, round in cross-section; leaves 2-ranked; sheaths open (except Glyceria); flowers/fruits
each enclosed by 2 bracts (lemma and palea) (fig. 537) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Poaceae, 2:252
60. Flowers not reduced (although sometimes small), readily visible, solitary, or forming various inflorescences; perianth
present, or if absent, then surrounded by perianth-Iike spathe.
62. Leaves not sheathing at base.
63. Flowers in dense heads (often simulating a single flower), subtended by involucral bracts (Aster. Euthamia,
Solidago) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asteraceae, 1:375
63. Flowers borne variously, not in heads, lacking involucral bracts.
64. Leaves and stems scabrous (fig. 13h) (Campanula aparinoides) . . . . . . . . . . . . . . . . . . . Campanulaceae, 1:365
64. Leaves and stems not scabrous.
65. Leaves alternate.
66. Flowers bilaterally symmetrical, blue (Lobelia) . . . . . . . . . . . . . . . . . . . . . . . Campanulaceae, 1:365
66. Flowers radially symmetrical, white, yellow, or green.
67. Leaves succulent; flowers 5-merous; ovary superior.
68. Flowers white with yellow eye; inflorescence a stalked spike, terminal spikes often
paired, axillary spikes single (Heliotropium eurassavicum) . .......... Boraginaceae, 1:285
68. Flowers green (petals absent); inflorescence a sessile compact cluster of mostly 3
flowers, axillary (Suaeda) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Chenopodiaceae, 1:79
Key 7 xli
67. Leaves not succulent; flowers 4-merous; ovary inferior (fig. 174c.d,e) (Ludwigia linearis)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onagraceae, 1:210
65. Leaves opposite.
69. Corolla of 4 separate petals, borne at summit of floral tube; stamens separate from petals;
ovary inferior (Epilobium palustre) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onagraceae, 1:210
69. Corolla united, lobes 4 or 5, floral tube absent; stamens inserted on corolla tube; ovary su-
perior.
70. Petals deep blue or purple; corolla lobes fringed or toothed at apex (Gentianopsis) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gentianaceae, 1:267
70. Petals blue, pink, lavender, or white; corolla lobes not fringed or toothed at apex .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Scrophulariaceae, 1:327
62. Leaves sheathing at base.
71. Flowers in dense heads.
72. Flower/fruit heads axillary or supra-axillary and terminal; leaves keeled (Sparganium) .... Sparganiaceae, 2:296
72. Flower/fruit heads terminal; leaves not keeled (Eryngium aquaticum) . . . . . . . . . . . . . . . . . . . Apiaceae, 1:245
71. Flowers borne variously, not in heads.
73. Leaf margins spinulose to conspicuously toothed, or if margin entire, at least leaf sheath splnulose
(Najas) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Najadaceae, 2:68
73. Leaf margins entire.
74. Flowers bilaterally symmetrical, with 1 larger petal forming a distinct lip (fig. 598c,d); stamens and
pistils united, modified to form a column . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orchidaceae, 2:328
74. Flowers radially symmetrical, lacking a lip; stamens and pistils appearing norma\' not specialized.
75. Leaves acute to blunt at apex, lacking a terminal pore; perianth yellow; fruit a capsule (2os-
terella) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pontederiaceae, 2:312
75. Leaves blunt at apex, with a terminal pore (fig. 365f); perianth greenish; fruit an inflated
follicle (fig. 365e) (Scheuchzeria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scheuchzeriaceae, 2:33
c
-leaf
creeping
rhizome
Fig. 10. a. Scirpus acutus or related bulrushes; b. Dulichium. portion of stem with scattered leaves; c. Eleocharis
acicularis or related species; d, e. rosette plants (such as Sagittaria, Eriocaulon), generalized; f. Myrio-
phyllum tenellum; g. Cyperaceae (sedges), generalized (F).
xlii
b
g
sheath
c d
Fig. 11. a. Butamus; b. Poaceae (grasses), leaf. generalized; c. Typha, leaf. diagrammatic; d. leaf with tapered
sheath, diagrammatic; c. Xyris, generalized; f. Sparganium, portion of leaf, generalized; g. Cyperaceae
(sedges), triangular cross-section, plant generalized; h, i. Sparganium or Acarus, leaf sections, diagram-
matic (F).
xliii
erect
stem
rhizome
b
septate
roots
Fig. 12. a. Ranunculus reptans var. ovalis; b. Eleocharis acicularis; c. Xyris, leaf, generalized; d. Lobelia dortmanna,
leaf section, diagrammatic; e. Isoetes, generalized; f. funcus pelocarpus, submersed form; g. Lobelia dort-
manna; h. Subularia; i. Eriocaulon, showing some roots enlarged; j. Myosurus, leaf, generalized (F).
xliv
-+--blade
\\JI1!'I!r---ligule
a
~ ~
>- sheath
h
d e
Fig. 13. a. Eleocharis quadrangulata, showing inflorescence and square stem cross-section; b. Eleocharis, general-
ized; c. Poaceae (grasses), leaf. generalized; d. Scirpus pungens or related species. showing inflorescence
and triangular stem cross-section; e. funcus effusus. inflorescence; f. Scirpus hotarui or related species,
inflorescence and ascending involucral bract; g. funcus, flower. generalized; h. Campanula aparinoides.
basal portion ofleaf; i. Eleocharis smallii or related species. upper portion (F).
xlv
Key 8 Leaves basal or cauline, less than 10 times as long as wide
1. Leaves many-ranked.linear-subulate; plants reproducing by spores. sporangia borne in a strobilus (fig. 35) ........... Lycopodiaceae. 1:23
1. Leaves alternate. opposite. or whorled; plants reproducing by flowers and seeds.
2. Stamens and pistils united. modified to form a column (orchids unlikely to be collected in vegetative condition); flowers bilaterally
symmetrical. with 1 larger petal forming a distinct lip (figs. 596f.g. 605e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Orchidaceae.2:328
2. Stamens and pistils appearing normal. not specialized; flowers radially or bilaterally symmetrical (lip present in Pinguicula and
Thalia).
3. Leaves basal; flowering scape naked or bearing a small leaf or stem so short that leaves appear to arise mostly from one point.
4. Leaves covered with sticky glands.
5. Leaves petiolate. blades spatulate. oval. or linear. glands distinctly stalked (fig. 108f); corolla white. of separate petals.
lacking a spur; flowers numerous. borne in a I-sided inflorescence (scorpioid cyme) (Drosera) . . . . . . . . . . . Droseraceae. 1:120
5. Leaves lacking a petiole. broad. spatulate to elliptic. glands not stalked; corolla lavender. united. with a long spur; flowers
solitary borne, on a scape (Pinguicula) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Lentibulariaceae. 1:352
4. Leaves lacking sticky glands.
6. Leaves tubular. open at upper end (water accumulating in tubular portion) (fig. 107a) (Sarracenia) . . . . . . . Sarraceniaceae. 1:120
6. Leaves flat.
7. Leaves with 3 or more veins running more or less parallel from base toward apex (fig. 14e).
8. Leaves with stipules at base of petiole (fig. 18c) (Potamogeton; short-stemmed forms of normally long-stemmed
plants which have been stranded in the mud. especially common in P. gramineus. fig. 385c. P. natans. P. pulcher. and
P. oblongus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Potamogetonaceae, 2:36
8. Leaves lacking stipules.
9. Plants usually more than 5 cm tall (few plants of Pamassia parviflora shorter).
10. Inflorescence an elongate golden-yellow spadix (fig. 398d) (Orontium) . . . . . . . . . . . . . . . . . . . . . Araceae.2:76
10. Inflorescence various or flowers solitary.
11. Flowers 4- or 5-merous.
12. Flowers solitary; scape with a single leaf near middle (Pamassia) . . . . . . . . . . . . . Saxifragaceae.l:177
12. Flowers in an inflorescence; scape lacking leaves.
13. Petals lavender. 5-merous. borne in a panicle (Limonium) . . . . . . . . . . . . Plumbaginaceae. 1:101
13. Petals whitish (scarious). 4-merous. borne in a spike (Plantago cordata) . . . . . Plantaginaceae. 1:323
11. Flowers 3-merous.
14. Perianth pink. in a compact inflorescence; leaves evergreen (Helonias) . . . . . . . . . . . . . Liliaceae. 2:320
14. Perianth white. solitary or in an open inflorescence; leaves not evergreen.
15. Flowers in an inflorescence. spathes absent; leaf blades much longer than wide . . Alismataceae. 2: 3
15. Flowers solitary. arising from a spathe; leaf blades nearly round to slightly longer than wide
(fig. 360) (Hydrocharis, Limnobium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hydrocharitaceae. 2:11
9. Plants 5 cm or less tall.
16. Rosettes of leaves connected by runners as thick as the petioles (fig. 18e) (Ranunculus reptans var. ovalis,
common) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ranunculaceae. 1:50
16. Rosettes connected by white thread-like runners (Limosella aquatica, rare) ........... Scrophulariaceae. 1:327
7. Leaves with 1 major vein running from base toward apex. usually with lateral veins branching from it (fig. 14d.g).
17. Leaves with a strong skunk-like odor when crushed; flowers borne in a globose spadix. subtended and surrounded
by a spathe (fig. 40lc) (Symplocarpus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Araceae. 2:76
17. Leaves lacking a skunk-like odor; flowers solitary, or inflorescence various. not borne in a spadix.
18. Leaves floating, margins coarsely toothed. petiole inflated; fruit a 4-spined nut (fig. 167c) (Trapa) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trapaceae. 1:206
18. Leaves emersed. margins entire to finely toothed or crenate; fruit a capsule, achene, or utricle.
19. Plants large. 1-3 m tall; blades 30-50 cm long (Thalia) . . . . . . . . . . . . . . . . . . . . . . . . Marantaceae. 2:310
19. Plants smaller, less than 1 m tall; blades 20 cm or less long.
20. Flowers solitary. bilaterally symmetrical. spurred (Viola) . . . . . . . . . . . . . . . . . . . . . . . Violaceae, 1:125
20. Flowers more than 1. borne in an inflorescence. radially symmetrical. not spurred (SaxiJraga) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Saxifragaceae. 1: 1 77
3. Leaves cauline or cauline and basal; flowers not borne on a scape.
21. Plants (at least leaves) distinctly succulent; plants mostly of coastal or of inland saline or alkaline sites.
22. Cauline leaves opposite (fig. 15a).
23. Leaves extremely minute. scale-like. stems appearing leafless (fig. 74) (Salicomia) . . . . . . . . . . . . . . Chenopodiaceae. 1:79
23. Leaves larger. not scale-like.
24. Leaves sessile; flowers variously colored. but not green.
25. Leaves with dots (hand lens needed); flowers weakly bilaterally symmetrical (Bacopa) .. Scrophulariaceae. 1:327
25. Leaves lacking dots; flowers radially symmetrical.
26. Leaf bases connate (fig. 16d); perianth parts separate; pistils 3 or 4; fruit a follicle (Crassula) ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Crassulaceae. 1: 1 77
26. Leaf bases not connate; perianth parts united; pistils 1; fruit a capsule (Glaux) . . . . . . Primulaceae.1:168
24. Leaves petiolate (often some leaves hastate); flowers greenish (Atriplex) . . . . . . . . . . . . . . . Chenopodiaceae. 1:79
22. Cauline leaves alternate (fig. 15c). at least upper leaves.
xlvi
Key 8 xlvii
Fig. 14. a. Rumex. leaf; b. Rumex. leaves basal. arising from one point; c. Potamogeton. terrestrial form; d. Trapa.
leaf; e. Orontium. leaf; f. Limnobium. leaf; g. Viola lanceolata. leaf; h. Alismataceae (water-plantains).
leaf. generalized (F).
Ii
g
Fig. 15. a. Opposite leaves, diagrammatic; b. whorled leaves, diagrammatic; c. alternate leaves, diagrammatic;
d. Bidens, leaf; e. Scrophulariaceae (figworts), leaf, diagrammatic; f. Callitriche, showing floating rosette
of uppermost leaves and fruits in leafaxils; g. Ludwigia palustris, showing an axillary fruit; h. Decodon,
leaf; i. Lysimachia, lower portion of leaf, generalized; j. Eupatorium perfoliatum, pair of leaves, bases
connate (F).
Iii
d
@, I. .
o
e ~h
Fig. 16. a. Elodea; b. Callitriche, generalized, with some axillary fruits; c. Hippuris, emersed upper portion;
d. Tillaea, upper portion; e. Elatine triandm, submersed form; f. Elatine, capsule, generalized; g. Elodea;
h. Didiplis diandm, terrestrial form; i. Ammannia Xcoccinea, leaf and leaf base, showing truncate leaf
base and ridge running down stem from leaf base (F).
liii
d
Fig. 17. a. Penthorum, leaf with stipules at base; b. petiole with sheathing leaf base; c. Trapa, leaf with swollen
petiole; d. Rumex, section of stem with leaf, showing stipular sheath; e. Rorippa palustris var. glabrata,
leaf; f. Lobelia cardinalis, portion of leaf showing gland-like callus at vein tips along margin; g. Proserpi-
naca palustris, showing leaf variation on a single plant (F).
liv
c
Fig. 18. a. Myriophyllum tenellum; b. Lythrum alatum, section of stem with leaf, showing ridges running down
stem from leaf base; c. Potamogeton, leaf with pair of stipules at base, generalized; d. Nasturtium offici-
nale, dwarfed form; e. Ranunculus reptans var. ovalis; f. Ludwigiapolycarpa, leaf; g. Potamogeton, general-
ized, stems reaching and partly floating on surface; h. Ranunculus laxicaulis, leaf; i. Polygonum, section
of stem with leaf, showing stipular sheath, generalized (F).
Iv
Aquatic
and Wetland Plants
of Northeastern
North America
2
Angiosperms: Monocotyledons
Angiosperms
Class Liliopsida
Monocotyledons
Subclass Alismatidae
Butomaceae / Flowering-rush Family
1. Butomus (Flowering-rush)
Perennials, arising from fleshy rhizomes; leaves erect or floating; flowers pink, bisexual, numerous, borne in an umbel; carpels 6,
separate, connate at very base; fruit a whorl of follicles with long beaks.
REFERENCES: Anderson, Zeis, and Alam, 1974; Core, 1941; Gaiser, 1949; Roberts, 1972; Staniforth and Frego, 1980; Stuckey, 1968;
Zenkert, 1960.
1. B. umbellatus L. Fig. 339 in North America is the result of introductions from two differ-
Marshes and shores. S. Me., St. Lawrence R. system in Que., ent geographical regions of the Old World: the populations of
L. Champlain valley of vt. and N.Y., Connecticut R. valley of the Great Lakes and more westerly localities are related to Euro-
Conn., w. along the Great Lakes to N.Y., Ohio, Mich., Wisc., pean plants; the plants of the St. Lawrence River region are
Minn., and widely scattered sites in c. Ohio, N.D., S.D.. Mont .. more like those of Asia, which are treated by Soviet botanists
Ida., and Man.; natzd. from Eur. Anderson and his colleagues (Fedchenko, 1934; Czerepanov, 1981) as B. junceus Turcz. (=
(1974) note that the presence of Butomus umbellatus sensu lato B. umbellatus var. junceus (Turcz.) Micheli).
1. Pistils and achenes in a single flat-topped ring (fig. 340c); flowers all bisexual, stamens 6 ......................... 1. Alisma
1. Pistils and achenes in a dense globose head (figs. 342f, 355d); flowers bisexual or unisexual, stamens more than 6.
2. All flowers bisexual; achenes plump, conspicuously ridged, not winged (fig. 342c,g); leaf blade never sagittate; roots not
septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. Echinodorus
2. Upper flowers unisexual, usually staminate, lower flowers pistillate or sometimes bisexual (petals then bearing a basal spot,
S. calycinal, or plants sometimes dioecious (S. latifolia); achenes flattened, winged, not ridged (fig. 355e); leaf blade often
sagittate; roots septate (figs. 345a, 355a) ..................................................... 3. Sagittaria
1. Alisma (Water-plantain)
Perennial herbs, arising from corm-like rhizomes; leaves all basal, ribbon-like, ovate, or elliptic; flowers bisexual, borne in a panicle;
petals white; fruit achenes, borne in a circle on a flattened receptacle.
1. Leaves submersed and ribbon-like (fig. 340b) or, if emersed, then lanceolate to narrowly elliptic, 0.5-2 cm wide (fig. 340a);
achenes with 2 dorsal grooves and a central ridge (fig. 340e) 1. A. gramineum
3
rhizome
Fig. 339. Butomus umbellatus: a. habit, submersed plant; b. habit, emersed plant; c. plant base; d. flower;
e. cluster of follicles (NHAES).
4
Alismataceae / Water-plantain Family 5
1. Leaves emersed with ovate to elliptic blades (fig. H1a), up to 15 cm wide, occasionally submersed or floating (fig. 341b);
achenes with single dorsal groove (fig. Hlf).
2. Flowers 7-13 mm wide; sepals at anthesis 3-4 mm long; petals 3.5-6 mm long; fruiting heads 4-7 mm in diameter;
achenes 2.2-3 mm long ................................................................. 2. A. triviale
2. Flowers 3-3.5 mm wide; sepals at anthesis 2-2.5 mm long; petals 1-2 mm long; fruiting heads 3-4 mm in diameter;
achenes 1.5-2 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3. A. subcordatum
2. Echinodorus (Burhead)
Annual or perennial herbs, often spreading by creeping rhizomes; leaves basal; submersed leaves lanceolate-Iinear, phyllodial or
ribbon-like; emersed leaves petiolate with ovate, lanceolate-truncate, to cordate blades; flowers bisexual. borne in umbellate to
whorled to paniculate inflorescences; petals white; fruit achenes, in globose heads.
1. Plants small, seldom reaching 15 cm tall; leaves lanceolate-Iinear (fig. 342a); stamens 9 ....................... 1. E. tenellus
1. Plants large, more than 15 cm tall; leaves ovate or broadly lanceolate-truncate, occasionally cordate; stamens 12-22.
2. Flowering stems erect; flowers 8-10 mm wide; stamens 12; veins of sepals smooth ......................... 2. E. berteroi
2. Flowering stems arching, often rooting at nodes (fig. 342d); flowers 12-20 mm wide; stamens 20-22; veins of sepals
papillate-ridged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. E. cordifolius
1. E. tenellus (Mart.) Buchenau Fig. 342 the East; e. Del. w. to Ohio, Ill., Iowa, Neb., and Calif., s. to Fla.,
Sandy shores, pond margins, and wet depressions. Local, Mass. e. Tex., and Mex.; w.1. and S.Am. (E. rostratus (Nutt.) Engeim.;
w. to Ont., Mich., and Minn., s. to c. Fla., s. Tex., and Mex.; w.1., E. berteroi var. IanceoIatus (Engelm.) Fassett)
C.Am., and S.Am. (E. tenellus var. parvuIus (Engelm.) Fassett,
E. parvuIus Engelm.) 3. E. cordifoIius (1.) Griseb. Fig. 342
Swamps, ponds, streams, and ditches. Md. and Va.; w. Ky., sw.
2. E. berteroi (Spreng.) Fassett Fig. 343 Ind., and s. Ill. w. to Mo. and Kans., s. to nw. Fla., e. Okla., and
Shallow ponds, muddy shores, marshes, and ditches. Local in e. Tex.; w.1. and S.Am. (E. radicans (Nutt.) Engelm.)
Adams and Godfrey, 1961; Beal, Wooten, and Kaul, 1982; Bogin, 1955; Haynes and HeIlquist, 1996; Smith, 1895;
REFERENCES:
Wooten, 1970, 1971. 1973a, 1973b.
1. Petals with a green or purple basal spot; sepals of mature flowers appressed (fig. 345d); lower flowers bisexual; stamens 9-15.
2. Petals with a green basal spot; leaf blades present. not spongy, broadly sagittate or ovate (fig. H4b); plants of freshwater
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1a. S. montevidensis ssp. calycina
dorsal
grooves
central
ridge
Fig. 340. Alisma gramineum: a. habit, emersed plant; b. habit, submersed plant; c. fruiting head; d. achene,
side view; e. achene, dorsal view (NHAES).
6
dorsal
groove
e f
S~
T c
Fig. 341. Alisma subcordatum: a. habit. emersed plant; b. habit. submersed plant; c. flower; d. fruiting head;
e. achene. side view; f. achene. dorsal view (NHAES).
Alisma triviale: g. flower (NHAES).
7
b
c
, e
Fig. 342. Echinodorus tenellus: a. habit; b. nodding flower; c. achene, oblique, side, and dorsal views (NHAES).
Echinodorus cordifolius: d. habit; e. flower; f. fruiting head; g. achene; h. seed (G&W).
8
d
Fig. 343. Echinodorus berteroi: a. habit, emersed plant; b. habit, submersed plant; c. portion of inflorescence;
d. flower; e. stamen; f. portion of infructescence; g. achene (Mason).
9
10 ANGIOSPERMS
2. Petals with a purple basal spot; leaf blades absent, or if present, spongy, lanceolate to obovate, or rarely sagittate (fig. 345);
plants of tidal marshes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1b. S. montevidensis ssp. spongiosa
1. Petals all white; sepals of mature flowers reflexed or spreading (fig. 355b,d); lower flowers pistillate, or all flowers pistillate or
staminate; stamens more than 15.
3. Pedicels of pistillate flowers recurved in fruit (fig. 346a); leaf blades unlobed.
4. Plants emersed, leaves erect, with blades, 5-20 em long, narrow to broadly ovate, occasionally with 1 or 2 basal projec-
tions; filaments pubescent .......................................................... 2. S. platyphylla
4. Plants submersed, leaves linear, often recurved, to ribbon-like, bladeless. or sometimes expanding into narrow blades
up to 4 em long; filaments glabrous.
5. Leaves linear-awl-shaped. often firm, recurved, up to 30 em long (fig. 347c); plants usually estuarine ...... 3. S. subulata
5. Leaves flexuous. ribbon-like. over 30 em long (fig. 347a). sometimes developing elliptic. oval, or ovate floating leaf
blades; plants often in fast-moving freshwater ............................................. 4. S. filiJormis
3. Pedicels of pistillate flowers divergent, ascending; sessile or subsessile in fruit (fig. 34Sa,b); leaf blades lobed or unlobed.
6. Stamens with pubescent filaments (figs. 34Sc. 350c); blades unlobed (rarely lobed in S. rigida).
7. Emersed and submersed leaves round in cross-section and quill-like (fig. 34S) .......................... 5. S. teres
7. Emersed leaves flattened or with petiole and distinct elliptic blade; submersed leaves flattened. not quill-like.
S. Pistillate flower and fruit heads sessile or subsessile (fig. 34ge); scape often bent (fig. 349a) ............. 6. S. rigida
S. Pistillate flower and fruit heads pedicelled (fig. 351a); scape straight. upright.
9. Stamen filaments linear ....................................................... 7. S. lanciJolia
9. Stamen filaments dilated at base.
10. Achene 1.5-2.5 mm long; beak of achene less than a fourth as long as width of achene; anthers as long
as or longer than filaments.
11. Phyllodia 1 em or less wide, linear-Ianceolate, acute at apex; pedicels of mature pistillate heads 1-3.5
cm long ................................................. Sa. S. graminea ssp. graminea
11. Phyllodia more than 1 em wide, linear. rounded at apex; pedicels of mature pistillate heads 4-6.5
em long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 8b. S. graminea ssp. weatherbiana
10. Achene 2.5-3 mm long; beak ofachene about half as long as width ofachene (fig. 353c); anthers shorter
than filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. S. cristata
6. Stamens with glabrous filaments; blades with basal lobes (except rarely lobed in S. ambigua).
12. Leaves linear-Ianceolate to elliptic, unlobed (rarely lobed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10. S. ambigua
12. Leaves sagittate (rarely unlobed, then mostly in juvenile plants).
13. Achenes with beak horizontal (fig. 355e); bracts 1.4 cm or less long; when plants staminate, flowers with
21-40 stamens ............................................................. 11. S. latiJolia
13. Achenes with beak ascending or erect (fig. 359d); bracts 0.7-4 cm long; when plants staminate (S. cuneata),
flowers with 10-24 stamens.
14. Achenes 1.8-2.6 mm long, with tiny erect beak 0.2-0.5 mm long (fig. 357e); submersed plants forming
ribbon-like leaves (fig. 357a) and/or floating lanceolate or sagittate leaves (fig. 357c); emersed plants
often with recurved petioles ................................................. 12. S. cuneata
14. Achenes 2.1-4 mm long. with erect or oblique beak 0.4-2.1 mm long; plants always emersed, petioles
ascending.
15. Achenes lacking facial resin ducts, facial wings not extending onto beaks; bracts firm, papery,
pointed at tip.
16. Petiole sharply 5-winged; fruiting heads 1-1.5 cm in diameter; pedicels of pistillate flowers 3-12
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13. S. australis
16. Petiole ridged, not winged; fruiting heads (1.2)1.7-2.2 cm in diameter; pedicels of pistillate
flowers 10-20 mm long ........................................... 14. S. brevirostra
15. Achenes with facial resin ducts (fig. 359d), facial wings extending onto beaks; bracts thick, herba-
ceous, blunt at tip ................................................ 15. S. engelmanniana
carpus spongiosus (Engelm.) J. G. Sm.; S. calycina var. spongiosa streams. Coastal plain, Va. s. to n. Fla. (S. weatherbiana Fern.; S.
Engelm.; S. spatulata 0. G. Sm.) Buchenau) graminea var. weatherbiana (Fern.) Bogin)
3. S. subulata (L.) Buchenau Fig. 347 10. S. ambigua J. G. Sm. Fig. 354
Tidal waters and shores of marshes, streams, and sloughs, in- Swamps and shores. Sw. Mo., Kans., and Okla.
land in fresh or brackish water. Coastal plain, Mass. s. to Fla.,
w. to Ala.; w.I. and n. S.Am. 11. S. latifolia Willd. Arrowhead, Wapato, Duck-potato
Figs. 355, 356
4. S. fili/ormis J. G. Sm. Fig. 347 Ponds, lakes, streams, sloughs, and marshes. Nfld. and N.B. w.
Ponds, lakes, and streams. Coastal plain, Me. and Mass. s. to e. to B.C., s. to n. Fla., Ala., La., Tex., Calif., and Mex.; C.Am. and
Pa., N.C., Fla., and Ala. This species occasionally flowers, but S.Am. The erect leaves are extremely variable, ranging from
has not been observed in fruit in the northern portion of its narrow bladeless leaves to narrowly sagittate to broadly sagit-
range. (S. stagnorum Small; S. subulata var. gracillima (S. Wats.) tate leaves. In the southern portion of the range, plants which
J. G. Sm.; S. subulata var. natans (Michx.) J. G. Sm.) are sparsely pubescent, their bracts and calyx often densely pu-
bescent, have been recognized as var. pubescens (Muhl.) J. G. Sm.
5. S. teres S. Wats. Dwarf Wapato Fig. 348 Several varieties and forms have been based on leaf shapes, but
Locally abundant in sandy and coastal plain freshwater ponds such leaf variation may all occur on a single plant. (S. latifolia
of se. Mass.; rare elsewhere. Mass. s. to R.I., s. N.J.. and e. Md. var. obtusa (Muhl. ex Willd.) Wieg.)
1. Plants floating on surface (or rooted when stranded), not submersed; leaves basal, with distinct petiole and blade (fig. 360).
2. Petals absent or, if present, then less than 1.5 times as long as sepals; anthers ca. 3.5 mm long, elongate; aerenchyma
spaces in central portion of floating leaf large, 0.3-1.7 mm in diameter (fig. 360b); stipules 1, adnate to petiole base; leaf
blade about twice as long as basal lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. Limnobiwn
2. Petals present, more than 1.5 times the length of sepals; anthers ca. 1 mm long, oval, as wide as long; aerenchyma spaces
in central portion of floating leaf small, 0.22-0.54 mm in diameter (fig. 360d); stipules 2, free from petiole; leaf blade about
1.5 times as long as basal lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. Hydrocharis
Fig. 344. Sagittaria montevidensis ssp. calycina: a. portion of inflorescence; b. leaf; c. flower; d. fruiting heads;
e. achene (C&C).
12
~'(j--- sepal
:
.\f:, .'.'.).'i.i'
. -' '.
. f
d
e
Fig. 345. Sagittaria montevidensis ssp. spongiosa: a. habit; b, c. leaf blade variations; d. flower; e. fruiting head;
f. achene (NHAES).
13
a
14
floating leaf
Fig. 348. Sagittaria teres: a. habit. partly submersed plant; b. habit. emersed plant; c. stamen. showing pubes-
cent filament; d. fruiting head; e. achene (NHAES).
16
e
. "i
\.r,i: '
i
'
,II",
1','/"
f
., '
'
'j C
Fig. 350. Sagitta ria lancifolia: a. habit (J. G. Smith, 1895); b. flower (C&C); c. stamen (J. G. Smith. 1895);
d. achene (J. G. Smith. 1895).
18
scape ------II
Fig. 351. Sagittaria graminea ssp. ,qraminea: a. habit. plant with lanceolate, tapering blades (NHAES); b. habit,
plant with linear phyllodia (NHAES); e. submersed sterile rosette (NHAES); d. achene (J. G. Smith,
1895).
19
Fig. 352. Sagitta ria graminea ssp. weatherbiana: a. habit. emersed plant; b. habit, submersed plant; c. achene
(PB).
20
fruiting
head
pedicel
Fig. 353. Sagittaria cristata: a. habit, emersed plant (PB); b. habit, submersed sterile rosette a. G. Smith, 1895);
c. achene (PB).
21
staminate
flowers
Fig. 354. Sagittaria ambigua: a. habit (PB); b. stamen G. G. Smith, 1895); c. achene (PB).
Sagittaria australis: d. habit; e. achene (Braun).
22
sepal
beak
tuber
a e
septate roots
Fig. 355. Sagittaria lati/olia: a. habit; b. pistillate flower; c. staminate flower; d. fruiting head; e. achene
(NHAES).
23
Fig. 356. Sagittaria Iatifolia: leaf variations (NHAES).
24
floating leaf
Fig. 357. Sagittaria cuneata: a. habit, submersed plant; b. sterile submersed rosette; c. habit, emersed plant;
d. fruiting heads; e. achene (NHAES).
25
Fig. 358. Sagitta ria brevi rostra: a. habit; b. leaf; c. flower; d. calyx; e. stamen; f. portion of infructescence;
g. achene (C&C).
26
Fig. 359. Sagi ttaria engelmanniana: a, b.
habit; c. fruiting head; d. ache ne
(NHAES).
27
28 ANGIOSPERMS
1. Plants submersed. rooted. or fragments free-floating beneath water surface; leaves basal or cauline. sessile.
3. Leaves basal. long. linear and ribbon-like (fig. 361) with a broad lacunae band (fig. 361g) ................. " 3. Vallisneria
3. Leaves cauline. short. opposite or whorled. lacking a lacunae band.
4. Leaves 1.2-4 cm long; flowers with nectaries .................................................. 4. Egeria
4. Leaves 0.6-l.7 cm long; flowers lacking nectaries.
5. Leaves in whorls of (2)4-6; leaf margins conspicuously toothed. midvein of lower surface often with spine-like teeth
(fig. 362c.d) (fresh specimens rough to the touch); spathe of staminate flowers spiny .................... 5. Hydrilla
5. Leaves in whorls of 3; leaf margins appearing entire to naked eye (minute teeth visible with hand lens). midvein of
lower surface lacking spine-like teeth (fresh specimens smooth to the touch); spathe of staminate flowers smooth.. 6. Elodea
Perennial herbs. stoloniferous; leaves basal. floating or emersed. with distinct blade and petiole; plants monoecious; staminate flowers
3-9 per spathe; pistillate flowers solitary in spathe; petals less than 1. 5 times the length of sepals; stamens 3-12; fruit an ovoid-
ellipsoid. many-seeded berry; seeds subglobose.
REFERENCES: Catling and Dore. 1982; Cook and Urmi-Konig. 1983; Lowden. 1992.
1. 1. spongia (Bosc) Steud. Fig. 360 Fla .. w. to e. Tex.; inland n. to se. Okla .. Mo .. and Ill. Our taxon
Shallow water or shores of ponds and streams. ditches. and is ssp. spongia.
swamps. Chiefly coastal plain. c. N.Y. s. N.J .. and e. Md. s. to
Perennial herbs. stoloniferous; leaves basal. floating or emersed. with distinct blade and petiole; plants monoecious; staminate flowers
1-4 per spathe; pistillate flowers solitary in spathe; petals greater than 1. 5 times the length of sepals; stamens 9-12; fruit an ellipsoid-
globose. many-seeded berry; seeds ellipsoidal.
REFERENCES: Catling and Dore. 1982; Catling and Porebski. 1995; Cook and Liiond. 1982; Dore. 1954. 1968; Roberts. Stuckey. and
Mitchell. 1981.
1. H. morsus-ranae L. European Fragbit Fig. 360 R. system in s. Que .. s. Ont.. and n. N.Y.. and L. Champlain. Vt.
Shallow water of marshes. ponds. and slow-moving streams. This species is similar to Limnobium. and the two are difficult to
This European introduction has spread along the St. Lawrence differentiate in the vegetative state.
Perennial herbs. submersed; leaves basal. long and ribbon-like; plants dioecious; pistillate flowers solitary. sessile. enclosed in a
tubular spathe. reaching surface by peduncle elongation; staminate flowers numerous. enclosed in spathes borne on short peduncles.
released and floating to surface; fruit elongate. cylindrical. peduncle recoiling after fertilization. submersing fruit.
4. Egeria (Waterweed)
Perennial herbs. submersed; leaves sessile. in whorls of 4( 6) along a round stem; plants dioecious; staminate flowers in spathes. borne
in upper leafaxils. raised to surface of water on a thread-like hypanthium; pistillate plants presently unknown in the United States.
1. E. densa Planch. Fig. 363 America. Populations are occasionally established at widely
Ponds. lakes. streams. and ditches. Vt. and e. Mass. w. to IlL. scattered areas. reproducing vegetatively. This species is more
Neb .. Kans .. and Okla .. s. to Fla. and Tex.; Oreg .. Calif.. and Ariz. common in the southern United States. (Elodea densa (Planch.)
This common aquarium plant has been introduced from South Casp.; Anacharis densa (Planch.) Vict.)
aerenchyma
aerenchyma blade
Fig. 360. Umnobium spongia: a. habit. floating plant; b. lower surface of floating leaf; c. habit. flowering and
fruiting stage (G&W).
Hydrocharis morsus-ranae: d. leaf (PB).
29
e
peduncle
Fig. 361. Vallisneria americana: a. habit, staminate plant; b. habit, pistillate plant; c. staminate spathe; d. stami-
nate flower; e. pistillate flower; f. fruit; g. section of leaf showing lacunae band (NHAES).
30
spine-like c
teeth
Fig. 362. Hydrilla verticillata: a. habit (G&W); b. habit (PB); c. lower surface of leaf (G&W); d. enlargement,
portion of lower leaf surface (G&W).
31
c
.~.r
~
..
Fig. 363. Egeria densa: a. habit; b. portion of plant with staminate flower; c. leaf (NHAES).
Elodea nuttallii: d. habit (NHAES); e. habit (F); f. leaf (NHAES); g. pistillate flower (NHAES); h. stami-
nate flower (NHAES); i. staminate bud within spathe (NHAES).
32
Juncaginaceae I Arrow-grass Family 33
5. Hydrilla (Hydrilla)
Perennial herbs, submersed; leaves in whorls of (2)4-8; leaf margins sharply serrate, midvein serrate on lower surface ofleaf; plants
monoecious or dioecious; staminate flowers short-stalked, detaching and floating to surface; pistillate flowers raised to surface on an
elongated hypanthium; fruit a soft, spiny, cylindrical or narrowly conic, few-seeded capsule.
l. H. verticillata (LJ.) Royle Fig. 362 tive to Uganda and Tanzania. this plant has spread Widely in
Ponds. lakes. streams. impoundments. and ditches. S. Conn .. many parts of the world and is considered by many to be one of
Md .. Del.. D.C., and N.C. s. to Fla .. w. to Tex.; Iowa (apparently the most troublesome weeds in the southern United States.
eradicated); w. Wash. and Calif.; C.Am. and S.Am.; unconfirmed Seeds seldom mature in North America.
report in s. N.Y. (Trudeau. 1982; Les et aI.. 1997). Believed na-
6. Elodea (Waterweed)
Perennial herbs. submersed; leaves along stem in whorls of 3 (some opposite); plants monoecious, dioecious. or polygamodioecious;
flowers with 3 sepals and 3 petals; staminate flowers with stamens monadelphous. flowers rising to surface by elongation at base of
hypanthium. or flowers sessile. but breaking loose at maturity. rising to surface where they open and float by divergent sepals;
pistillate flowers raised to surface by an elongated hypanthium; fruit a leathery. ovoid. few-seeded capsule.
REFERENCES: Cook and Urmi-Konig. 1985; Marie-Victorin, 1931; st. John. 1920. 1965.
1. Leaves generally 0.3-1.5 mm wide, acute at apex. not densely overlapping toward stem tip (fig. 363e); staminate flowers
enclosed in sessile spathes. released. floating to surface before anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. E. nuttaIlii
1. Leaves generally 1-5 mm wide. blunt at apex. densely overlapping at stem tip (fig. 364b); staminate flowers enclosed in stalked
spathes. reaching surface at anthesis by elongation at base of hypanthium ............................... 2. E. canadensis
1. E. nuttaIlii (Planch.) st. John Fig. 363 are rarely encountered and have leaves which tend to be longer
Acid to moderately alkaline streams. lakes. and ponds. P.B.I.. and more linear or lanceolate-oblong than the leaves of pistillate
N.B .. s. Que .. and n. Ont. w. to Ida .. s. to N.C .. Ala., Miss .. Okla .. plants. (Anacharis canadensis (Michx.) Planch.)
and N.M. (E. occidentalis (Pursh) st. John; Anacharis occidentalis
(Pursh) Vict.) . E. schweinitzii (Planch.) Casp.. a plant very similar to E. cana-
densis but with bisexual flowers. has been described and is
2. H. canadensis Michx. Fig. 364 known only from the vicinity of Bethlehem. Pennsylvania. It
Alkaline lakes. ponds. and streams. Que. w. to Sask. and Wash .. has not been collected since 1832 and is presumed extirpated.
s. to N.C., Ala .. n. Ark.. N.M., Ariz .. and Calif. Staminate plants
Perennial herbs. arising from creeping, jointed rhizomes; leaves linear, grass-like. with terminal pore at apex. broadly sheathing at
base; flowers borne in a loose raceme; fruit an inflated follicle.
l. S. palustris L. Fig. 365 americana (Fern.) Hult. The best diagnostic character for sterile
Sphagnum bogs. Nfld. w. to Man. and Wash .. s. to N.J.. Pa., n. plants is the terminal pore at the apex of the leaf. (S. palustris
Ohio. n. Ill.. n. Iowa. Neb .. N.M .. and Calif. Our taxon is ssp. var. americana Fern.)
Perennial herbs. arising from a short rhizome. some producing stolons; leaves basal. slender. slightly succulent, rush-like; flowers in
a spike-like raceme; fruit a cluster of 3-6 follicles separating from a persistent rachis.
REFERENCES: Crow and Hellquist. 1982; Fernald. 1903; Ford and Ball. 1988; LOve and Lieth. 1961; Love and LOve. 1958.
g
Fig. 364. Elodea canadensis: a. habit (Reed); b. tip of branch (F); c. staminate flower (Reed); d. staminate flower
(NHAES); e. pistillate flower (NHAES); f. capsules (Reed); g. seeds (Reed).
34
Fig. 365 . Scheuchzeria palustris: a.
habit. flowering plant;
plant; d. flower; e. fruit; b. habit. fruiting plant;
f. leaf apex wi th ter mi nal c. habit. sterile submerse
pore (NHAES). d
35
REFERENCES: Catling and Dobson, 1985; Fernald, 1932b; Hagstrm, 1916; Haynes, 1974, 1978, 1980, 1985; Haynes and Hellquist,
1996, 2000. Haynes and Williams, 1975; Hellquist, 1980, 1984; Hellquist and Crow, 1980, 1986; Hellquist and Hilton, 1983; Hell-
quist, Philbrick, and Hilton, 1988; Holub, 1997; Hunt and Lutz, 1959; Les, 1983; Les, Cleland, and Philbrick, 1995; Les and Haynes,
1996; Les and Sheridan, 1990a, 1990b; Moore, 1915; Muenscher, 1936; Ogden, 1943; Philbrick, 1983, 1988; Reznicek and Bob-
bette, 1976; Stuckey, 1979; Wehrmeister, 1978; Wehrmeister and Stuckey, 1992; Yeo, 1965.
c
Fig. 366. Triglochin gaspense: a. habit; b. leaf sheath; c. flowers; d. fruits (NHAES).
Triglochin striatum: e. habit; f. fruit, side view; g. fruit, top view (PB).
37
, .
scape
sheath
Fig. 367. l'riglochin maritimum: a. habit; b. leaf sheath; c. flower; d. fruit (NHAES)
Triglochin palusire: e. habit; f. leaf sheaths; g. flowers. two stages; h. fruit (NHAES).
38
Potamogetonaceae / Pondweed Family 39
3. Stipules on lower portion of stem loosely clasping. usually inflated. disintegrating with age; leaves 0.2-2.0 mm wide;
fruit not commonly maturing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2a. S. filifarmis ssp. accidentalis
3. Stipules on lower portion of stem tightly clasping or slightly inflated. persisting with age; leaves 0.2-1.0 mm wide; fruit
commonly maturing ........................................................ 2b. S. filifarmis ssp. alpina
2. Free portion of stipule barely forming a ligule, up to 1 mm long; summits of midstem stipular sheaths distinctly inflated,
twice the diameter of stem; fruit 3-3.5 mm long ............................................... 3. S. vaginata
1. S. pectinata (L.) Borner Sago Pondweed Fig. 368 formis ssp. occidentalis (Robbins) Les & Haynes; Potamogetonfili-
Calcareous, saline, and alkaline waters. Nfld. w. to Alask., s. to formis var. occidentalis (Robbins) Morong)
Fla., Calif., and Mex.; W.I .. C.Am., and S.Am. Leaves are variable
2b. S. filiformis ssp. alpina (Blytt) Haynes, Les, & Kral
in length and thickness, often coarser than illustrated in fig.
Fig. 369
368a. This species is easily recognized by the much-branched
Calcareous and brackish waters. Green!. and Lab. w. to Alask.,
stems and numerous filiform leaves spreading in fan-like fash-
s. to n. Me., n. N.H., n. Vt., e. Pa., n. Ohio, Mich., Neb., n. N.M.,
ion; tubers are frequently produced. (Caleageton pectinatus (L.)
Nev., and Calif. (Coleogeton filiformis ssp. alpinus (Blytt) Les &
Les & Haynes; Patamageton interruptus Kit.; P. pectinatus L.)
Haynes; Potamogetonfiliformis var. borealis (Raf.) st. John; P.fili-
formis var. alpinus (Blytt) Aschers. & Graebr.; P. filiformis var. ma-
2. S. filiformis (Pers.) Borner
counii Morang)
2a. S. filiformis ssp. occidentalis (Robbins) Haynes, Les, &
Knil Fig. 369 3. S. vagi nata (Turcz.) Holub Fig. 369
Cold, usually calcareous flowing waters. W. Nfld. and N.S. w. to Calcareous and brackish waters. N. Que. and sw. Ont. w. to
N.W.T., Yuk., and Alask., s. to n. Me., N.Y., N.J., Mich., Minn., n. Alask., s. to s. Wisc., ne. Minn., n. Colo., Utah, w. Mont., Ida.,
N.D., Colo.. N.M., and c. Oreg. Often confused with S. vaginata and c. Oreg. Most plants in our range, previously thought to
because of enlarged stipular sheaths, which deteriorate early belong to Stuckenia (Potamogeton) vaginata, are S. filiformis ssp.
and are therefore often lacking on older plants. (Coleogeton fili- occidentalis. (Coleogeton vaginatus (Turcz.) Les & Haynes; Potamo-
geton vaginatus Turcz.)
2. Potamogeton (Pondweed)
Perennial, or rarely annual, aquatic herbs, from rhizomes or winter buds; leaves alternate, submersed or submersed and floating,
variable. with a characteristic midvein, sometimes with distinct bands or rows of lacunae; flowers 4-merous, borne in spikes, axillary
or terminal, submersed or emergent; fruit drupe-like.
1. Leaves stiffish, conspicuously 2-ranked (fig. 371a), with basal lobe at junction with stipule (fig. 371b) ............. 1. P. robbinsii
1. Leaves lax, not conspicuously 2-ranked (or if so then with wide lacunae bands present, P. epihydrus), lacking basal lobes.
2. Stipules adnate to leaf base, at least partly so (fig. 370c).
3. Submersed leaves obtuse to acute at apex; floating leaves rounded at apex.
4. Submersed leaves obtuse at apex; fruit 1.3-2.2 mm wide, lateral keels scarcely developed (fig. 371g) or rarely well-
developed, beak absent ............................................................. 2. P. spirillus
4. Submersed leaves acute at apex; fruit 1-1.5 mm wide. lateral keels well-developed, minute beak present (fig. 370h)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3. P. diversifolius
3. Submersed leaves acute to long-tapering at apex (fig. 370g); floating leaves pointed at apex (fig. 370f).
5. Submersed leaves 0.1-0.6 mm wide; lacunae not abundant; floating leaves 3-7-veined, 0.1-2 cm long; embryo
distinctly coiled, with more than 1 complete coil (fig. 370i) ................................. 4. P. bicupulatus
5. Submersed leaves 0.2-1(2.6) mm wide; lacunae abundant (fig. 372e); floating leaves 9-23-veined, 2-4(5.5) cm
long; embryo curved. but without 1 complete coil (fig. 372f) ............................... 5. P. tennesseensis
2. Stipules completely free (figs. 373c, 380a) or with only a few adnate to leaf base.
6. Submersed leaves linear, thread-like to ribbon-like. 0.1-10 mm wide.
7. Fruit keel 0.2-1.2 mm wide (fig. 372c); lacunae prominent in submersed leaves (fig. 372b); floating leaves often
present.
8. Submersed leaves 1-3(7)-veined, usually linear-filiform, not strongly 2-ranked, some with stipules adnate to leaf
base; floating leaves 2-4(5.5) em long. acute at apex; fruit with slightly convex sides. . . . . . . . . . . .. 5. P. tennesseensis
8. Submersed leaves 3-13-veined. ribbon-like, often strongly 2-ranked. with stipules completely free; floating leaves
3-8 em long. bluntly cuspidate or rounded at apex; fruit with flattened sides .................... 6. P. epihydrus
7. Fruit keel less than 0.2 mm wide; lacunae not prominent in submersed leaves (expect in some specimens of P. pusillus
ssp. tenuissimus); floating leaves present or absent.
free }
} portion
st'lpUIe
} sheath
c d
Fig, 368. Stuckenia pectinata: a. habit (F); b. stem. showing adnate stipules (Mason); c, leaf with stipule (F);
d. leaf apex (F); e, flowers (Mason); f. spike (Mason); g, fruit (F),
40
b
inflated _----'~:'\.
stipule
e f
g
a
41
42 ANGIOSPERMS
fruit 1.73.5
44 ANGIOSPERMS
1. P. robbinsii Oakes Robbins' Pondweed Fig, 371 13. P. Xhaynesii Hellquist & Craw Haynes' Pondweed
Ponds, lakes, and slow streams, often deep water, Lab, w. to B.C., Fig. 376
s. to N.J., Md., Ind., Ala., Utah. and Calif. This species rarely Alkaline waters. Vt. w. to Ont.. Mich .. and Minn. This taxon is
forms fruit. of hybrid origin. with P. strictifolius and P. zosteriformis as the
parental species.
2. P. spirillus Tuckerm. Fig. 371
Acid to moderately alkaline waters. Nfld. w. to Man. and S.D., ] 4. P. ogdenii Hellquist & Hilton Ogden's Pondweed
s. to Del., Md., e. Ohio, Iowa, and e. Nebr. Fig. 376
Alkaline waters. Vt.. w. Mass .. N.Y.. and Ont. This taxon is of
3. P. diversifolius Raf. Fig. 370 hybrid origin, with P. hillii and P. zosteriformis as the parental
Moderately alkaline waters. Conn. w. to Pa., n. Ohio, III., Minn., species.
Mont., sw. B.C., and Oreg., s. to Fla., Tex., Calif., and Mex.; WI.
(P. capillaceus Pair.) 15. P. obtusifolius Mert. & Koch Fig. 377
Moderately alkaline waters. Lab. and w. Nfld. W. to Yuk.. s. to
4. P. bicupulatus Fern. Fig. 370 Conn .. N.J.. Pa.. n. Mich.. Wisc.. and Minn.; disjunct in nw.
Acid waters. Me. w. to scattered locations in Ont., Mich., Wisc., Wyo .. nw. Mont.. and Vancouver I., B.C.
and e. Minn., s. to N.J., N.Y., Pa., and n. Ind.; Va. and Tenn. (P.
diversifolius var. trichophyllus Morang; P. capillaceus Poir. is some- 16. P. pusillus L.
times misapplied to this taxon.)
16a. P. pusillus ssp. pusillus Fig. 378
Alkaline waters. Lab. and Nfld. w. to N.WT. and Alask.. s. to
5. P. tennesseensis Fern. Fig. 372
Fla., Calif.. and Mex.; C.Am. (P. panormitanus Biv.; P. panormita-
Streams and rivers. W Pa. and e. Ohio, s. to WVa., w. Va., e. Ky.,
nus var. minor Biv.; P. pusillus var. minor (Biv.) Fern. & Schub.)
and Tenn. This species is very similar to diminutive plants of
P. epihydrus. 16b. P. pusillus ssp. tenuissimus (Mert. & Koch) Haynes &
Hellquist Fig. 378
6. P. epihydrus Raf. Leafy Pondweed Fig. 372 Mostly acid waters. Nfld. W. to Alask .. S. to n. Fla .. La .. and Calif.;
Alkaline and acid waters. S. Lab. and Nfld. w. to Minn. and most abundant in the Northeast. (P. berchtoldii Fieber; P. pusiIlus
Man .. s. to Ga .. Ala .. and Miss.; Ida .. B.C .. and Wash. s. to Colo .. var. tenuissimus Mert. & Koch)
Nev.. and n. Calif. (P. epihydrus var. ramosus (Peck) House; P. epi-
16c. P. pusillus ssp. gemmiparus (Robbins) Haynes &
hydrus var. nuttallii (C. & S.) Fern.)
Hellquist Fig. 379
Acid waters. S. Que .. Me .. N.H .. Mass .. R.I .. and Conn. (P. gemmi-
7. P. confervoides Reichenb. Fig. 373
parus Robbins; P. pusiIlus var. gemmiparus Robbins)
Acid waters. often in bog ponds. Nfld., w. Ont.. Upper Pen.,
Mich .. and Wisc. s. to s. N.J.. Pa .. N.C .. and n. S.C. (P. tucker-
17. P. vaseyi Robbins Vasey's Pondweed Fig. 379
mani Robbins)
Acid to moderately alkaline waters. N.B., s. Que., and Ont. w. to
Wisc. and Minn .. s. to Conn .. Pa .. ne. Ohio. n. Ind .. and n. III.
8. P. zosteriformis Fern. Flat-stemmed Pondweed. Eel-grass
This species is difficult to locate in late summer because plants
Pondweed Fig. 373
with floating leaves typically deteriorate by mid-August. Potamo-
Alkaline waters. W Nfld. and Gaspe Pen., Que. w. to Man ..
getonlateralis Morang is no longer recognized; it was based on a
N.WT.. and Alask.. s. to n. Va .. s. Ohio. Neb.. w. Mont.. Oreg ..
type specimen which consisted of a mixed collection of P. vaseyi
and n. Calif.
and P. pusillus (Hellquist. Philbrick. and Hilton. 1988).
9. P. hillii Morang Hill's Pond weed Fig. 374
18. P. natans L. Floating Brownleaf. Floating-leaved
Clear. cold. calcareous streams and ditches. Vt.. w. Mass .. and
Pondweed Fig. 380
nw. Conn. w. to s. Ont.. n. Mich. and ne. Wisc .. S. to N.Y.. Pa ..
Acid to alkaline waters. Greenl. and Nfld. W. to Alask.. S. to N.J ..
and e. Ohio; w. Va.; somewhat rare. most abundant in w. N.E.
Md., n. Ind .. Kans .. N.M., Ariz., and s. Calif.
and n. Mich. (P. porteri Fern.)
19. P. oakesianus Robbins Oakes' Pondweed Fig. 380
10. P. foliosus Raf. Leafy Pond weed Fig. 374
Acid waters. Nfld. w. to Ont., Mich .. Wisc.. and Minn .. s. to N.J..
Alkaline waters. Nfld .. P.E.I .. and N.S. w. to c. Alask.. s. to Fla ..
Del.. Va .. and ne. S.C.; Mont. and B.C.
Calif.. and Mex.; WI. and C.Am. Our taxon is ssp. foliosus. (P.
foliosus var. macellus Fern.)
20. P. crispus 1. Curly-leaved Pondweed Fig. 381
Alkaline or polluted waters. Sw. Que. and s. N.H. W. to s. Ont..
11. P. friesii Rupr. Fig. 375
Mich .. Minn .. Alta .. and B.C.. s. to Fla., Tex., Ariz .. and Calif.;
Alkaline waters. Lab. and w. Nfld. w. to c. Alask .. s. to Pa .. w.
C.Am. and S.Am.; intro. from Eurasia.
Neb .. n. Utah. and Wash.
21. P. amplifolius Tuckerm. Large-leaved Pond weed. Bass
12. P. strictijolius Ar. Benn. Fig. 375
Weed. Muskie Weed Fig. 382
Alkaline waters. Nw. Nfld .. e. Que .. n. Me .. and vt. w. to Ont..
Acid to alkaline waters. often at great depths. Nfld .. N.S .. and
Man .. N.WT.. and B.C .. s. to nw. Conn .. N.Y.. Ind .. Minn .. Neb..
Gaspe Pen .. Que. w. to s. B.C .. S. to Va .. Ga., Ala .. Ark .. Okla ..
Wyo .. and n. Utah. (P. strictifolius var. rutiloides Fern.; P. longiligu-
Mont .. Ida .. and Calif.
latus Fern.)
c
embryo
coil
keel
Fig. 370. Potamogeton diversi!olius: a. habit (PB); b. leaf apex (F); c. leaf base with stipule (F); d. fruit (PB).
Potamogeton bicupulatus: e. habit (PB); f. upper floating leaves (PB); g. leaf apex (F); h. fruit (F);
i. fruit (PB).
45
a
keel
blade
\I[Iii'f free
stipule
g
} fu,ro
c
e
m f
Fig. 37l. Potamogeton robbinsii: a. upper portion of plant (PB); b. section of stem with leaf base (F).
Potamogeton spirilIus: c. habit (NHAES); d. upper portion of plant (F); e. leaf base with stipule (F);
f. leaf apex (F); g. fruit (F).
46
lacunae
~
keel
d
lacunae
~
c
./
Fig. 372. Potamogeton epihydrus: a. upper portion of plant (PB); b. section of/eaf
showing broad lacunae band
(PB); c. fruit (F).
Potamogeton tennesseensis: d. upper portion of plant; e. section of leaf
showing lacunae band; f. fruit
(PB).
47
peduncle
.t..t--- -blade
+---,11 -+,4,1- ---- stipule
rhizome
48
bristle
Fig. 374. Potamogeton hilIii: a. upper portion of plant (PB); b. section of stem with leaf base and stipule (F);
c. fruit (F).
Potamogeton !oIiosus: d. upper portion of plant; e. habit. showing winter buds; f. section of stem with
leaf base; g. fruit (F).
49
e
nodal
---~:uI
gland
l. d
beak
--H-rfJ,i--- stipule
Fig. 375. Potamogeton strictifolius: a. upper portion of plant (F); b. section of stem with leaf base and stipule
(F); c. winter bud (PE); d. leaf apex (PE); e. fruit (PE).
Potamogeton friesii: f. section of stem with leaf base and fibrous stipule; g. leaf apex; h. midsection of
leaf; i. winter bud; j. fruit (PB).
50
d
Fig. 376. Potamogeton Xhaynesii: a. upper portion of plant; b. leaf apex; c. winter bud; d. fruit (PB).
Potamogeton ogdenii: e. upper portion of plant; f. leaf apex; g. winter bud; h. fruit (PB).
51
'-+.....,~'*- stipule
nodal
1I1Q1F---gland
c b
e
Fig. 377. Potamogeton obtusifolius: a. habit (NHAES); b. leaf apex (NHAES); c. section of stem with leaf base
and stipule (F); d. winter bud (F); e. fruit (PB).
52
h
midrib
Fig. 378. Potamogeton pusillus ssp. pusillus: a. habit (Reed); b. section of stem with leaf and stipule (F); c. flower
(Reed); d. fruit (F).
Potamogeton pusillus ssp. tenuissimus: e, f. upper portion of plant; g. winter bud; h-j. leaf apices, show-
ing lacunae band variations (F).
S3
b
---,fH---- stipule
tfllr-- blade
,.-!~__ nodal
gland
floating
~--Ieaf
a
floating
leaf
f g
e
d
Fig. 379. Potamogeton pusilIus ssp. gemmiparus: a. upper portion of plant (F); b. section of stem with leaf base
and stipule (F); c. winter bud (PB).
Potamogeton vaseyi: d, e. upper portion of plant (PB); f. fruit (PB); g. fruit (F).
54
floating leaf
\\:---- petiole
*---- stipule
55
c
Fig. 381. Potamogeton crispus: a, b. upper portion of plant; c. young inflorescence; d. inflorescence; e. leaf;
f. section of stem with leaf bases and stipules; g. winter bud; h. stem, cross-section; i, j. fruit (Mason).
56
c
Fig. 382. Potamogeton amplifolius: a. upper portion of plant (F); b. fruit (PB).
Potamogeton pulcher: c. habit (G&W); d. flower (G&W); e. fruit (PB).
57
58 ANGIOSPERMS
22. P. pulcher Tuckerm. Spotted Pondweed Fig. 382 Calif.; Bahamas. Potamogeton gramineus hybridizes with many of
Acid waters. Chiefly coastal plain. N.S . s. Me. and c. N.H. s. to the broad-leaved species. especially P. illinoensis. P. richardsonii.
Fla . w. to La .. Tex . and Okla.; Mississippi embayment and n. to and P. perfoliatus. (P. gramineus var. maximus Morong ex Ar.
scattered locations in Ind . Ohio. Ont.. Mich . Wisc.. e. Minn . Benn; P. gramineus var. myriophyllus Robbins)
and se. Mo.
27. P. illinoensis Morong Illinois Pondweed Fig. 386
23. P. oblongus Viviani Fig. 383 Alkaline waters. Que . e. N.B.. Vt.. and Mass. w. to s. Man .. sw.
Acid waters. E. Nfld . St. P. et Miq . and Sable I.. N.S. Les (1983) N.W.T. and B.C . s. to Fla . Tex.. s. Calif.. and Mex.; w.I.. C.Am.
notes that P. oblongus is diploid. whereas the closely related P. and S.Am.
polygonifolius Pourret of Europe is tetraploid.
28. P. praelongus Wulf. Whitestem Pondweed. Muskie Weed
24. P. nodosus Poir. Fig. 383 Fig. 386
Fast-flowing streams of low alkalinity. or ponds and slow- Deep. moderately alkaline waters. Lab. and Nfld. w. to Alask.
moving streams of high alkalinity. Que. w. to B.C .. s. to Fla.. s. to N.J.. Ohio. Ind . Colo . Utah. Calif.. and Mex.
Ala .. Tex.. Ariz . Calif.. and Mex.; WI . C.Am . and S.Am. (P.
americanus C. & S.) 29. P. richardsonii (Ar. Benn.) Rydb. Clasping-leaved
Pondweed. Bass Weed Fig. 387
25. P. alpinus Balb. Fig. 384 Alkaline waters. E. Que. w. to Alask.. s. to Me . Vt.. w. Mass ..
Moderately to strongly alkaline waters. Nfld. w. to Alask.. s. to N.Y.. Ind . Iowa. Colo . Utah. and Calif.; Guat. In very shallow
Me . n. N.H .. Vt.. e. Pa .. N.Y.. Mich .. Minn .. Colo.. Utah. and water. floating leaves may occur.
Calif. (P. alpinus var. subellipticus (Fern.) Ogden; P. alpinus var.
tenuifolius (Raf.) Ogden) 30. P. perfoliatus L. Clasping-leaved Pondweed Figs.387.
388
26. P. gramineus L. Variable Pondweed Fig. 385 Acid. alkaline. and saline waters. Green!' and Nfld. w. to Ont..
Acid to alkaline waters. Green!.. Lab . and Nfld. w. to Alask.. s. s. to N.C . Ala . and La.; C.Am. (P. perfoliatus var. bupleuroides
to N.J. Md . Pa .. n. Ohio. n. Ind .. n. Iowa. Kans . N.M . Ariz . and (Fern.) Farw.)
Perennial or annual aquatic herbs; leaves alternate. stipules adnate to blades; floating leaves absent; flowers 2. enclosed in leaf sheath
at anthesis; fruit on long stalk (podogyne). 4 per flower. usually below water surface.
REFERENCES: Fernald and Wiegand. 1914; Richardson. 1980. 1983; Thieret. 1971.
1. R. maritima L. Widgeon-grass. Ditch-grass Figs. 389. to estuarine sites with chiefly underwater self-pollination. and is
390 diploid. The other is adapted to tidal streams with chiefly surface
Brackish and saline waters. Chiefly coastal; Nfld. s. to Fla.; pollination. and is tetraploid. Although the range of variability
Wash. s. to Calif. and Mex.; scattered locations inland; w.I.. of these two entities overlaps. the two appear to be reproduc-
C.Am.. and S.Am. Studies by Richardson (1980. 1983) have tively isolated and can be differentiated by a combination of
revealed considerable variability in Ruppia. casting doubt on the morphological characters. Robust plants in strongly alkaline
distinctiveness of some varieties recognized by Fernald and Wie- waters of midwestern and western North America are some-
gand (1914). Further. Richardson (1983) demonstrated that in times treated as var. occidentalis (S. Wats.) Graebn. or as a dis-
coastal New Hampshire two entities occur which appear to war- tinct species. R. cirrhosa (Petango) Grande. (R. occidentalis S.
rant recognition at the species leve!. One is an annual adapted Wats.).
Perennial herbs. submersed; stems arising from rhizomes; leaves linear. opposite to subopposite; fruit in axillary clusters of 2-6.
bilaterally symmetrical. usually dentate on one side. short-stalked.
Evidence from molecular studies by D. Les (pers. comm.) strongly suggests that Zannichellia ought to be included as a genus in the
Potamogetonaceae.
1. Z. palustris L. Horned Pondweed Fig. 391 U.S. to Mex.; C.Am. and S.Am. (Z. palustris var. major (Boenn.)
Saline waters along the coast. alkaline and brackish waters in- W. D. J. Koch.)
land. Nfld . e. Que . and Ont. w. to Alask.. s. throughout the
c
\.II1If----- stipule
Fig. 383. Potamogeton oblongus: a. upper portion of submersed plant with floating leaves; b. stranded plant;
c. fruit (PB).
Potamogeton nodosus: d. upper portion of plant with floating leaves; e. fruit (PB).
59
e
Fig. 384. Potamogeton alpinus: a. upper portion of plant, acute-tipped form (F); b. leaf apex, acute form (PB);
c. upper portion of plant, blunt-tipped form (F); d. leaf apex, blunt form (PB); e. fruit (PB).
60
Fig. 385. Potamogeton gramineus: a, b. habit, submersed plant (F); c. habit, stranded plant (F); d. fruit (PB).
61
awl-shaped
e b
Fig. 386. Potamogeton iIlinoensis: a. upper portion of plant (PB); b. upper portion of plant with floating leaves
(PB); c. fruit (F).
Potamogeton pmelongus: d. upper portion of plant (PB); e. boat-shaped apex (F); f. fruit (PB).
62
stipular
fibers
k
6
ed
f
e
I g
IJ h
Fig. 387. Patamogeton richardsanii: a. habit, upper portion of plant; b. fruit (PE).
Patamogeton perfaliatus: c. upper portion of plant; d-j. leaf variations; k. fruit (PE).
63
f
Fig. 388. Potamogeton perfoliatus: a. habit; b. branch with inflorescence; c. flower; d. anthers and sepal;
e. embryo; f. fruit (G&W).
64
Fig. 389. Ruppia maritima: a, b. habit; c. peduncle bearing 2 flowers, each with 2 anthers and 4 pistils; d. 2
flowers after fertilization; c. elongate podogynes with fruits at tips, and recoiled peduncle; f. fruit;
g. stipular sheaths; h. habit (Mason).
65
Fig. 390. Ruppia maritima: fruit, podogyne, and peduncle variations (Fernald and Wiegand, 1914).
66
Fig. 391. Zannichellia palustris: a. habit; b. portion of stem. showing stipular sheaths and axillary flowers;
c. single staminate flower and 2 pistillate flowers surrounded by spathe; d. fruit. longitudinal section;
e. f. fruit variations; g. fruit. outer coat deteriorated away (Mason).
67
68 ANGIOSPERMS
Perennial herbs .. submersed; stems flattened. arising from rhizomes; leaves alternate. up to 2 m long; flowers unisexual. borne in
rows surrounded by a sheathing leaf-like spathe; fruit a utricle.
1. Z. marina L.Eeigrass Fig. 392 Alask. s. to Calif. Our taxon is var. stenophylla Aschers. &
Common in saline water of estuaries and bays and in sea water Graebn.
along the open coast. Green!.. Lab .. and James Bay s. to Fla.;
Annuals. submersed; leaves subopposite or appearing whorled. edges serrate. spinulose. or entire; flowers unisexual. axillary. en-
closed in perianth-like spathe; fruit indehiscent. with a thin pericarp.
New evidence based on seed coat structure in Najas suggests a strong relationship with the Hydrocharitaceae (Shaffer-Fehre.
1991). Molecular evidence obtained by D. Les (pers. comm.) strongly corroborates this viewpoint. On the basis of this information.
Thorne's (1992) classification system deletes the Najadaceae and places Najas within the Hydrocharitaceae.
REFERENCES:Clausen. 1936. 1937; Fernald. 1923; Haynes. 1979; Haynes and HeIIquist. 1996; Meriliiinen. 1968; Stuckey. 1985;
Wentz and Stuckey, 1971.
1. Leaves serrate (fig. 393). midribs spiny on lower surface ofleaf; (fig. 393b); fruit 4-7.5 mm long; plants dioecious ...... 1. N. marina
1. Leaves minutely spinulose (fig. 394b.d) (occasionally seen with unaided eye) or entire, midribs not spiny on lower surface of
leaf; fruit 2-3.5(4.5) mm long; plants monoecious.
2. Leaf margins with 20-60 minute spinules; leaf bases tapered (figs. 394d. 395e).
3. Leaves tapering to long. slender apex; styles slender; seeds shiny ................................... 2. N.jlexilis
3. Leaves appearing more abruptly acute at apex; styles stout; seeds dull.
4. Style 0.5 mm or less long; seeds with lO-20 rows of areolae ................. 3a. N. guadalupensis ssp. guadalupensis
4. Style 0.7-1. 5 mm long; seeds with 30 or more rows of areolae.
5.. Seed smooth. 30-40 rows of areolae. ridge absent .......................... 3b. N. guadalupensis ssp. olivacea
5. ' Seed rough. 50-60 rows of areolae. prominent longitudinal ridge present . . . . . . .. 3c. N. guadalupensis ssp. muenscheri
2. Leaf margins with 6-20 minute spinules; leaf bases truncately lobed (figs. 396b. 397b).
6. Leaves somewhat stiff. often recurved. 0.3-0.5 mm wide; spinules ofleafmargins easily seen with hand lens. basal lobes
flabellate and finely toothed (fig. 396b); seeds with 12-18 rows of areolae. areolae shorter than wide (fig. 396d) ....
. . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4. N. minor
6. Leaves not stiff or recurved. less than 0.3 mm wide; spinules of leaf margins inconspicuous. even with hand lens. basal
lobes cuneate and coarsely jagged (fig. 397b); seeds with ca. 24 rows of areolae. areolae longer than wide (fig. 397d)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. N. gracillima
N.Y. w. to Wisc. and Minn. (N. guadalupensis var. olivacea (Ro- and Mass. w. to Ind. and Mich., s. to Fla., Miss., Ark., La., and
send. & Butt.) Haynes; N. olivacea Rosend. & Butt.) s. Okla. Intro. from Eur., spreading rapidly in the U.S.
3c. N. guadalupensis ssp. muenscheri (Clausen) Haynes &
5. N. gracillima (A. Br.) Magnus Fig. 397
Hellquist
Acid waters of ponds and lakes. N.S. and Me. w. to Ont. and
Tidal waters. E. N.Y. (N. guadalupensis var. muenscheri (Clausen)
Minn., s. to N.C., Ala., Miss., and Mo.; isolated station in Calif.
Haynes; N. muenscheri Clausen)
"""'--style
Fig. 394. Najasflexilis: a. habit (Reed); b. portion of stem (Reed); c. pistillate flower (Reed); d. leaf (F); e. fruit
(F); f. seed (F).
72
c
Fig. 395. Najas guadalupensis ssp. guadalupensis: a, b. upper portion of plant; c. leaf; d. leaf apex; e, f. young
staminate flowers; g. mature anther; h. anther, cross-section; i. pistillate flowers; j. seed (Mason).
73
d
Fig. 396. Najas minor: a. habit (G&W); b. leaf (F); c. pistillate flower (G&W); d. seed (G&W).
74
areolae
d ~\\, ~
leaf base --t I
b
\~-J
Fig. 397. Najas gracillima: a. habit (NHAES); b. leaf (F); c. fruit (NHAES); d. seed (F).
75
Subclass Arecidae
Acoraceae / Sweet-Rag Family
1. Acarus (Sweet-flag. Calamus)
Emersed perennials. arising from thick. creeping. aromatic rhizomes; leaves long-linear; flowers bisexual. 3-merous. greenish; fruit
a gelatinous, few-seeded berry,
1. A, americanus (Raf.) Raf, Fig, 398 been treated by Love and LOve (1957) as A. americanus (Raf.)
Wet meadows. marshes, ponds, and stream banks. Nfld., N.S., Raf. Introductions of A. calamus in the northeast, apparently for
P.E.I., and Que. w. to Minn., Mont., and Ida., s. to Fla. and Tex. medicinal purposes, present identification problems in discern-
This species is often confused in the sterile state with Iris, but is ing the pollen-sterile triploids from native A. american us popula-
easily recognized by its keeled leaves and sweet odor of the lower tions. According to Grayum (1987), studies of pollen morphol-
portion of the plant. Acarus calamus sensu stricto is a sterile trip- ogy in the Araceae support the removal of the genus Acarus and
loid, whereas the North American plants are diploid and have placement of it in its own family, the Acoraceae.
1. Spadix golden-yellow; spathe obscure, forming a sheath at base oflong scape (fig. 398d); leaves broadly oblong-elliptic ... 1. Orontium
1. Spadix green or brown; spathe broad. conspicuous; leaves ovate (fig. 40lb), cordate (fig. 400a), hastate or sagittate (fig. 399b).
2. Leaf blades hastate or sagittate (juveniles unlobed), with prominent vein extending into each basal lobe (fig. 399b) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. PeItandra
2. Leaf blades broadly rounded to cordate, lacking prominent vein extending into basal lobes (if present).
3. Spathe white; spadix on a long peduncle (fig. 400), emerging after leaves; plants lacking a strong odor when bruised
3. Calla
3. Spathe mottled, purplish-green and/or reddish-brown; spadix nearly sessile (fig. 401a), emerging very early. before
leaves; plant with a strong skunk-like odor when bruised ..................................... 4. Symplocarpus
1. Orantium (Golden-club)
Perennial herbs, growing from a deep rhizome; leaves basal.long-petioled, entire, often floating; flowers bisexual. borne in a golden-
yellow spadix; spathe a reduced. obscure sheath at base of scape; fruit a blue-green or brownish utricle.
76
Fig. 398. Acorus americanus: a. habit; b. spadix (NHAES).
Orontium aquaticum: c. habit. terrestrial plant; d. spadix with reduced spathe; e. habit. submersed
plant; f. submersed juvenile stage (NHAES).
77
78 ANGIOSPERMS
2. Peltandra (Arrow-arum)
Emersed perennials, with thick fibrous or subtuberous roots; leaves basal, long-petioled, blades hastate or sagittate; upper flowers of
spadix staminate, lower ones pistillate; fruit an amber or green berry.
1. P. virginica (L.) Schott & End!. Fig. 399 by the conspicuous vein paralleling the leaf margin (fig. 399b)
Bogs, swamps, and margins of ponds, rivers, and streams. C. Me. and by a single prominent midvein and a prominent vein ex-
and s. Que. w. to Mich., Ont., and Wisc., s. to Fla" se. Okla" and tending into each basal lobe. This highly variable species has
Tex. Vegetative plants are often confused with several species of had many dubious forms named, solely on the basis of leaf
Sagitta ria or Pontederia cordata, but can be easily differentiated shape.
Emersed perennials, arising from a long creeping rhizome; leaves cordate, long-petioled; flowers bisexual, borne on a spadix; spathe
ovate, pointed, upper surface white; fruit a red berry.
4. Symplocarpus (Skunk-cabbage)
Emersed perennials, arising from a thick, erect rhizome; leaves ovate to cordate, short-petioled; flowers borne on a spadix, subtended
by a mottled purplish-green and/or reddish-brown spathe; ovaries imbedded in spadix, globose or ovoid mass; entire plant producing
a foul skunk-like odor when bruised.
The duckweeds constitute a family characterized by floating plants which, lacking leaves, consist of globose or flattened fronds, with
or without roots, and very rarely are observed flowering. The family includes Wolffia, the smallest of the flowering plants, consisting
of almost microscopic meal-like bodies without roots. Frequently several species of the Lemnaceae are found intermixed.
REFERENCES: Clark and Thieret, 1968; Daubs, 1965; Hartog and van der Plas, 1970; Hillman, 1961; Hillman and Culley, 1978;
Landolt 1980,1981, 1986.
1. Wolffia (Water-meal)
Minute free-floating plants; rootless; fronds greatly reduced; flowers rarely produced; vegetative reproduction extensive.
This genus contains the smallest known flowering plants. The different species are extremely difficult to identify, especially if
allowed to dry. Every effort should be made to identify Wolffia while material is fresh or preserved in liquid.
IJlitt--- spadix
Fig. 399. Peltandra virginica: a. habit; b. leaf variations; c. spathe and spadix; d. spadix at fruiting stage and
with spathe cut away. below; e. seedlings to juvenile stages (NHAES).
79
c
peduncle ---AI
b
Fig. 400. Calla palustris: a. habit; b. spathe and spadix; c. spadix at fruiting stage (NHAES).
80
c
@
'\1IfI
f
Fig. 401. Symplocarpus fretidus: a. habit; b. leaf; c. spathe and spadix; d. spadix of male flowers; e. spadix of
female flowers; f. fruit (NHAES).
81
82 ANGIOSPERMS
1. Frond globose (fig. 402a). not dotted; some plants floating below surface under crowded conditions 1. W columbiana
1. Frond ellipsoidal (fig. 402b.c). dotted; plants floating on surface even when crowded.
2. Upper surface of frond flattened (fig. 402b) ................................................... 2. W borealis
2. Upper surface of frond raised to a conspicuous papule (fig. 402c) .................................. 3. W brasiliensis
1. W. columbiana Karst. Fig. 402 slow-moving rivers. N.H. and vt. w. to sw. Que., s. Ont., and s.
Floating plants of ditches. ponds. lakes. and slow-moving rivers. Minn., s. to Fla. and e. Tex.; B.C., Wash., and Oreg.; w.I. This
S. Me .. c. N.H .. and sw. Que. w. to s. Ont.. Mich .. Wisc .. and taxon is treated as W. punctata Griseb. in most manuals.
Minn., s. to Fla., Tex .. Calif., and Mex.; C.Am. and n. S.Am. This
and other species of Wolffia are often associated with stagnant 3. W. brasiliensis Wedd. Fig. 402
and/or polluted waters. Floating plants of quiet waters of ditches, ponds, and streams.
Mass. and Conn. w. to sw. Mich., Ind., IlL. Mo., and Kans., s. to
2. W. borealis (Engelm.) Landolt Fig. 402 Fla., Tex., and Mex. This taxon is treated as W. papuliJera C. H.
Floating plants of quiet waters of ditches, ponds. lakes, and Thompson in most manuals.
Tiny plants floating just below water surface; frond reduced. hollow, gradually narrowed from base to apex, 6-g mm long, asymmet-
rical. linear-attenuate or falcate or sigmoid, many times longer than wide; fronds occurring singly or cohering at base and radiating
in a stellate manner; flowers rare, reproduction primarily vegetative.
REFERENCES: Daubs, 1962; Jacobs, 1947; Wohler, Wohler, and Hartman, 1965.
1. Roots 5-20; frond broadly obovate to ovate. usually with an obvious red dot near center of upper surface ........... 1. S. polyrhiza
1. Roots 2-4; frond narrowly obovate to almost reniform. usually lacking a red dot near center of upper surface ........ 2. S. punctata
1. S. polyrhiza (L.) Schleid. Fig. 402 streams. Chiefly coastal plain, N.C. s. to Fla., w. to La. and e.
Standing and slow-moving waters of ditches, ponds. lakes, and Tex.; Mississippi embayment n. to sw. Ky. and s. Ill.; w. Pa.; Calif.
rivers. P.B.I., N.B., and se. Que. w. to s. B.C., s. to Fla., Tex., and and Ariz.; Trop. Am. Intro. from Old World tropics and to be
Mex.; w.I., C.Am., and n. S.Am. looked for throughout the Northeast (S. oligorrhiza (Kurz)
Hegelm.)
2. S. punctaia (Mey.) C. H. Thompson
Standing and slow-moving waters of ditches, ponds, and
4. Lernna (Duckweed)
Small. flattened, free-floating plants; fronds 1-5-nerved, with 1 root per frond; flowers minute. perianth lacking, with 2 staminate
flowers (of a single stamen each) and 1 pistillate flower enclosed in sac-like spathe; fruit a utricle; vegetative reproduction extensive.
1. Fronds long-petiolate. 6-15 mm long. with lateral fronds usually remaining attached to parent frond (fig. 402f.g); plants
usually floating below surface. often in tangled mats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. L. trisulca
1. Fronds sessile or nearly so. less than 6 mm long. forming single plants or small rosettes (fig. 402h-j); plants floating on surface.
2. Fronds elliptic to linear-oblong. less than 0.5-1.5 mm wide with sides somewhat parallel and ends rounded (fig. 402h).
obscurely I-nerved.
a b c
fronds
I---roots
~ k
83
84 ANGIOSPERMS
3. Fronds in clusters of (2)8-10, obovate to elliptic, distinctly asymmetrical at base, 2.5-5 mm long. . . . . . . . . .. 2.1. valdiviana
3. Fronds solitary or in pairs, ovate to broadly elliptic, nearly symmetrical, 1-2.5 mm long ................... 3. L. minuta
2. Fronds broadly oblong, obovate, or subcircular, 1.2-3 mm wide with all sides curved (fig. 402i,j), obscurely 3-nerved.
4. Root sheath with lateral wings; frond with apex pointed, apical and central papilla conspicuous. . . . . . . . . . .. 4. 1. perpusilla
4. Root sheath lacking lateral wings; frond with apex rounded, apical papilla not conspicuous .................. 5.1. minor
1. 1. trisulca L. Fig. 402 s. to Fla., Tex., Calif., and Mex. Landolt (1980,1981) differenti-
Quiet waters of ponds, lakes, and streams. M.l., Que., P.E.I. and ates within this complex L. aequinoctialis Welwitsch, distin-
N.S. w. to Alask., s. to Fla., Ala., Tex., N.M., Calif.. and Mex. guished chiefly by seed characters and the presence of only a
single papule.
2. 1. valdiviana Phil. Fig. 402
Quiet waters. N.H. and N.Y. w. to Ohio, s. Mich., Ill., Neb., Wyo., 5. L. minor L. Fig. 402
and Calif., s. to Fla., Tex., and Mex.; S.Am. Quiet, often highly eutrophic or polluted waters. S. Lab. and
Nfld. w. to B.C., s. to Fla. and s. Calif. Landolt (1980, 1981)
3. 1. minuta Humb. recognizes within this complex the widely distributed 1. turionif-
Quiet waters. Widely scattered from s. Ohio w. to s. Ill., Neb.. era, distinguished by the presence of several papules of nearly
s. Mont., and sw. Wash., s. to Fla., Tex., Calif., and Baja Calif. equal size and freely producing winter buds. (L. obscura (Aust.)
(L. minima Phil.; 1. minuscula Herter) Daubs; 1. minor var. obscura Aust.)
Emersed perennials; leaves grass-like, basal, 2-ranked, blades linear, abruptly or gradually dilated toward base; inflorescence a com-
pact, head-like spike; flowers bisexual, borne in axils of leathery or chaffy, overlapping bracts, often a single flower open at a time;
petals yellow; fruit a capsule.
1. Tips of lateral sepals extending beyond tips of subtending bracts (fig. 403b).
2. Keel of lateral sepals lacerate (fig. 403c) .................................................... 1. X. smalliana
2. Keel oflateral sepals ciliate (fig. 403g), fringed (fig. 404c), or entire (fig. 40Sg).
3. Plants bulbous and hard at base (fig. 403e) .............................................. 2. X. caroliniana
3. Plants soft and flattened at base. not bulbous.
4. Scapes SO-ISO em long. scabrous; keels of lateral sepals fringed ............................... 3. X.fimbriata
4. Scapes 3-45 em long. smooth; keels of lateral sepals entire .................................... 4. X. montana
1. Tips of lateral sepals enclosed by subtending bracts.
5. Keels oflateral sepals ciliate to ciliolate (fig. 40Sc) or scabrous.
6. Plants bulbous and hard at base (fig. 40Sa); lateral sepals tufted at tip (fig. 40Sc) . . . . . . . . . . . . . . . . . . . . . . . .. 5. X. torta
6. Plants soft and flattened at base. not bulbous; lateral sepals not tufted at tip .......................... 6. X. ambigua
5. Keels of lateral sepals lacerate (fig. 407g), erose. or dentate.
7. Plant bases bulbous. lowest outer leaves scale-like (fig. 407a); scapes twisted. often flexuous; upper portion ofleafbase
conspicuously twisted .............................................................. 7. X. platylepis
7. Plant bases not bulbous, lower leaves not scale-like; scapes usually not twisted or flexuous; upper portion of leaf base
not conspicuously twisted.
S. Upper scape 4-ribbed, nearly round in cross-section. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Sb. X. diJJormis var. curtissii
S. Upper scape 2(3)-ribbed, somewhat flattened.
9. Mature fruiting spikes 15 mm long or longer; seed surface mealy. 0.S-1.0 mm long or longer .......... 9. X. laxiJolia
9. Mature fruiting spikes less than 15 mm long (rarely 15 mm); seed surface not mealy. 0.4-0.5 mm long.
10. Plant bases pinkish or purplish. sometimes greenish; leaves spreading ............. Sa. X. diJJormis var. diJJormis
10. Plant bases pale yellow-green to straw-colored; leaves ascending ...... . . . . . . . . . . . . . . . . . . .. 10. X. jupicai
85
b
Fig. 403. Xyris smalliana: a. habit (NHAES); b. flowering spike (NHAES); c. lateral sepal (NHAES); d. seed (Kral,
1966b).
Xyris caroliniana: e. habit (G&W); f. flowering spike (G&W); g. lateral sepal (Kral, 1966b); h. seed
(Kral, 1966b).
86
lateral
sepal
bract
Fig. 4 0 4 .
X ,' " fimb
ria,"' ,. b a
h " (G&Wl'
b . ft o ." ri n g ,p ik e (G
&Wl; c. ,, "
1<9Y66
'" b,mbig""' "" " p ,l ( " " l,
~-ed (K,"I, 1966b).
). c. b a 6 ft ' [, J 9 6 6 6 ), d.
.o " " ri n g ,-pI " c d (K ," I,
ke; g.
87
bract-------!'
~
h
bract - - - 4 j l -
g
!If
~c
Fig. 405. Xyris torta: a. habit (NHAES); b. flowering spike (NHAES); c. lateral sepal (NHAES); d. seed (Kral.
1966b).
Xyris montana: e. habit (NHAES); f. flowering spike (NHAES); g. lateral sepal (NHAES); h. seed
(Kral. 1966b).
88
b c
Fig. 406. Xyris difformis var. difformis: a. habit (NHAES); b. flowering spike (NHAES); c. lateral sepal (NHAES);
d. seed (Kral, 1966b).
Xyris difformis var. curtissii: e. seed (Kral. 1966b).
89
h
Fig. 407. Xyrisplatylepis: a. habit (G&W); b. flowering spike (G&W); c.lateral sepal (Kral. 1966b); d. seed (Kral.
1966b).
Xyris laxifolia var. iridifolia: e. habit (G&W); f. flowering spike (G&W); g. lateral sepal (Kral. 1966b);
h. seed (Kral. 1966b).
90
bract
a c
Fig. 408. Xyris jupicai: a. habit; b. flowering spike; c. lateral sepal; d. seed; e. scape, cross-section (G&W).
91
92 ANGIOSPERMS
5. X. torta Sm. Fig. 405 and Me. w. to N.Y.. Ont.. Mich . n. Ind . and Wisc. s. mainly
Damp shores. sphagnous bogs. acid sandy swamps. and occa- along the coastal plain to Fla . La . and e. Tex. This is the most
sionally on dryish sandy pond shores. C. N.H. and Mass. w. to abundant species throughout the Northeast. (X. caroliniana of
Ont . N.Y. Mich. n. Ind . and Minn. s. to Ga . Tex . and Okla. Gray's Manual. 8th ed. (pro parte). and Fassett)
This is the only bulbous-based species found north of New
8b. X. di!formis var. curtissii (Malme) Kral Fig. 406
Jersey. Wet pine barrens. bogs. and ditches. Se. Can. and the Great
Lakes region; coastal plain. se. Va. s. to Fla . w. to e. Tex. This
6. X. ambigua Beyrich ex Kunth Fig. 404
small plant can be easily confused with X. montana in the north-
Moist or wet sands or sandy peat of bog margins. lakeshores.
ern portion of its range. but is distinguished from X. montana by
and ditches. Coastal plain. se. Va. s. to Fla . w. to e. Tex.; Cuba
lateral sepals that are lacerate toward the tip. and by scales with
andC.Am.
a green central portion. (X. curtissii Malme; X. bayardii Fern.)
7. X. platylepis Chapm. Fig, 407"
9. X. laxi/olia Mart. Fig. 407
Wet pinelands. swamps. peats. and ditches. Coastal plain. e. Va.
Wet sands. clays. and peats of marshes. swamps. and pond mar-
s. to Fla . w. to c. La. This species is most easily confused with
gins. Coastal plain. se. Va. s. to n. Fla. w. to e. Tex. Our taxon is
X. torta because of its scale-like leaf bases. but is easily distin-
var. iridi/olia (Chapm.) Kral. (X. iridi/olia Chapm.)
gUished by the lacerate keels of the sepals.
10. X. jupicai L. C. Rich Fig. 408
8. X. di//ormis Chapm.
Bogs. wet sands. roadside ditches. swamps. and lakeshores.
8a. X. di!formis var. di!formis Fig. 406 Chiefly coastal plain. N.J. s. to Fla .. w. to Tex . Ark . and Tenn.;
Damp sandy shores of ponds and lakes. in ditches and bogs. N.S. C.Am. and S.Am.
1. Scapes glabrous throughout; lacunar tissue in leaves evident without magnification; roots septate (fig. 411a); sepals 2.
petals 2 ............................................................................. 1. Eriocaulon
1. Scapes pubescent. at least toward upper portion (fig. 412a); lacunar tissue in leaves not evident without magnification; roots
not septate; sepals 3. petals absent .......................................................... 2. Lachnocaulon
1. Eriocaulon (Pipewort)
Submersed or emersed perennials. often tufted; roots conspicuously septate; leaves basal. linear. smooth. loosely cellular; inflores-
cence a head; plants monoecious or dioecious; flowers subtended by receptacular bractlets; fruit a loculicidal capsule.
scape
Fig. 409. Eriocaulon compressum: a. habit; b. receptacular bract; c. staminate flower; d. pistillate flower;
e. seed (G&W).
93
d. seed (G&W).
Fig. 410. Eriocaulon decangulare: a. habit; b. stamina te flower; c. pistillate flower;
94
b
J/----- scape
involucral
bracts
Pig. 411. Eriocaulon aquaticum: a. habit; b. flower head; c. pistillate flower; d. staminate flower (NHAES).
Eriocaulon parkeri: e. habit; f. flower head. subtended by involucral bracts; g. pistillate flower; h. stami-
nate flower (NHAES).
9S
Fig. 412. Lachnocaulon anceps: a. habit; b. staminate flower; c. pistillate flower without sepals; d. pistillate flower;
e. bract; f. sepal; g. seed (G&W).
96
Juncaceae I Rush Family 97
3. E. parkeri Robins. Fig. 411 L.I .. N.Y.. N.J. Del.. mts. of N.C.. n. Ohio. n. Ind .. Wisc. and
Saline waters and along muddy shores of estuaries. Rare; Minn. Submersed plants are often noted with scapes of up to 20
coastal. St. Lawrence R. estuary. Que.. N.B .. and N.E. s. to N.C. dm or more in length. (E. pellucidum Michx.; E. septangulare
(E. septangu!are var. parkeri (Robins.) Bovin & Cayouette) With.)
Usually perennial herbs. arising from rhizomes. grass-like; stems round in cross-section. pithy or hollow. rarely branching; leaves
round in cross-section. channeled. or flat. lower blades reduced to sheaths; flowers small. perianth parts similar (tepals). sepals 3.
petals 3. scale-like; stamens 3 or 6; fruit a 3-valved capsule. with numerous seeds.
There are numerous species of funcus. and they are often distinguished with some difficulty. Fruiting material is usually critical
for identification. Rushes are sometimes mistaken for sedges (see pp. 121-251) or grasses (see pp. 252-295). but are easily distin-
guished by flowers with 6 tepals and many-seeded capsules. The most easily recognized species is funcus effusus (Soft Rush). which
may occur in large clumps of several hundred stems and is often abundant in wet meadows. The true aquatics are f. militaris. J.
repens. J. subtilis. and dwarfed forms of J. marginatus and J. pe]ocarpus. The others are typically found in wet sites. the plants sometimes
occurring in shallow water.
REFERENCES: Brooks and Kuhn. 1986; Clements. 1990; Cope and Stace. 1978; Eleuterius. 1975; Hiimet-Ahti. 1980; Hermann. 1975;
Reinking. 1981; Snogerup. 1963. 1980; Stuckey. 1980b. 1981.
1. Inflorescence apparently borne laterally. lowest involucralleaf erect. appearing as a continuation of stem (fig. 413a.e).
2. Leaf sheaths often terminated by cylindrical blade about 1 mm in diameter (fig. 415d) ..................... 1. J. coriaceus
2. Leaf sheaths bladeless.
3. Individual flowers sub tended by 1 bracteole at base of pedicel; flowers/fruits in clusters of 2-6 in branched inflores-
cences; plants of tidal marshes ....................................................... 2.]. roemerianus
3. Individual flowers subtended by 3 bracteoles at base of pedicel (fig. 413f); flowers/fruits 1 per inflorescence axis. or in a
single. compound cluster; plants of inland freshwater sites. or if coastal. then non-tidal.
4. Stems borne in large clumps (fig. 413d); stamens 3 ........................................... 3. J. eJJusus
4. Stems arising singly in rows. or in small clumps along creeping rhizomes (fig. 414a); stamens 6.
5. Perianth shorter than capsule (fig. 414d). brown; seeds 0.7-1 mm long; rhizome thick.
6. Stem with longitudinal ribs; basal sheaths reddish-purple; perianth 2-2.7 mm long. brown. much shorter
than capsule ........................................................... 4. J. gymnocarpus
6. Stem without longitudinal ribs; basal sheaths straw-colored. grayish. brown. or dark brown; perianth 3.5-6
mm long. with dark reddish-brown markings. only slightly shorter than capsule ................. 5. J. arcticus
5. Perianth slightly longer than capsule. green; seeds 0.5 mm long; rhizome more or less slender . . . . . . . .. 6. J. filiJormis
1. Inflorescence terminating stem. subtended by 1 or 2 ascending or spreading involucralleaves (fig. 416aJ).
7. Leaves not septate.
8. Leaf blades round in cross-section. at least toward apex. but sometimes channeled basally.
9. Seeds long-tailed at each end (fig. 416h).
10. Heads with 1-4 flowers; capsule pale brown. mucronate; seeds 2.5-4 mm long ................... 7. J. stygius
10. Heads with 15-50 flowers; capsule green. blunt (fig. 416g); seeds 1-1.5 mm long ................... 8. J. vaseyi
98 ANGIOSPERMS
26. Mature fruit and perianth 2-4 mm long; seeds 0.7-0.9 mm long. with short tails. about a third as
long as body length; heads narrower. hemispherical. 5-20-flowered ............... 24. J. subcaudatus
25. Capsule much longer than perianth. dark brown; heads 2-7-flowered; inflorescence narrow. 3-6 times
longer than wide. branches ascending ..................................... 25. J. brevicaudatus
22. Seeds lacking tails (fig. 424d). or tails very short and inconspicuous.
27. Stems wealdy ascending to creeping or floating. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26. J. bulbosus
27. Stems erect.
28. Heads globose (fig. 424a). with lower flowers reflexed.
29. Capsules lanceolate. tapering to long slender tip (fig. 424c).
30. Blades of lower leaves laterally compressed; leaf sheaths approximately same width as blade
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . " 27. J. validus
30. Blades of lower leaves round to nearly round in cross-section; leaf sheaths distinctly wider than
blade.
31. Uppermost leaf blade (not the bract) as long as or longer than its sheath; sepals hard and firm.
32. Petals as long as or longer than sepals; sepals 2.5-4 mm long; lobes at summit of leaf
sheath 0.5-2.5 mm long; stems 0.7-1.5 mm in diameter near middle.
33. Lobes at summit of sheath 1-2.5 mm long; heads 2-12(30). each 15-40-flowered.
pale green to dull brown; stamens 3 ............................ 28. J. scirpoides
33. Lobes at summit of sheath 0.5-1 mm long; heads (1)2-15. each 8-20-flowered. red-
dish brown; stamens 6 ....................................... 29. J. nodosus
32. Petals slightly shorter than sepals; sepals 4-5.5 mm long; lobes at summit of leaf sheath
2.5-3.5 mm long; stems 2-4 mm in diameter near middle .................. 30.]. torreyi
31. Uppermost leaf blade (not the bract) shorter than its sheath; sepals soft and flexible ...... .
31. J. megacephalus
29. Capsules obovoid. abruptly contracting to a very short tip (fig. 425h) ................ 32. J. brachycarpus
28. Heads hemispherical or obconic (fig. 431b). with lower flowers not reflexed.
34. Stamens 3.
35. Perianth as long as or slightly longer than capsule.
36. Inflorescence of (40)50-500 heads; capsule rounded or minutely mucronate; sepals 2.2-2.9
mmlong.
37. Leaves 3.5 mm in diameter. with conspicuous ring-like bands at cross-partitions; roots
lacking tubers; petals 1.9-2.3 mm long; capsule 2.3-2.5 mm long ............ 33.]. nodatus
37. Leaves 1-3 mm in diameter. with obscure ring-like bands at cross-partitions; some roots
bearing tuberous thickenings; petals 2.4-2.8 mm long; capsule 2.4-2.9 mm long .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34. J. elliottii
36. Inflorescence usually of fewer than 50 heads; capsule acute and apiculate; sepals 3.3-4 mm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35. J. acuminatus
35. Perianth much shorter than capsule.
38. Sepals shorter than petals; petals more or less blunt; seeds 0.8-1.2 mm long. tails sometimes
inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22. J. brachycephalus
38. Sepals and petals of same length; petals acute to very sharp-pointed; seeds less than 0.7 mm
long. lacking tails.
39. Capsule lanceolate. 2.3-3 mm long; perianth lanceolate-acute. two-thirds the length of
capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36.]. debilis
39. Capsule slender.lanceolate-linear. 4-5 mm long; perianth linear-subulate. half the length
of capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37. J. difJusissimus
34. Stamens 6.
40. Flowers single or in heads of 2 or 3(5). often replaced by bulblets.
41. Stems lax and creeping. floating. or submersed.
42. Plants with firm base. not forming mats. if completely submersed. then consisting of tufts
of basal leaves. with stems often absent (usually occurring as an erect form) .... 9. J. pelocarpus
42. Plants with soft base. often forming mats. with elongate floating leaves and/or creeping
branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. J. subtilis
100 ANGIOSPERMS
1. J. coriaceus Mackenz. Fig. 415 Nfld. w. to Ont. and Minn., s. to N.E., Del., Pa., n. Ind., and Wisc.
Swamps, marshes, and wet woods. S. N.]. w. to Ky., Ark., and Submersed plants (forma submersus Fassett) are very similar to
Okla., s. to Fla. and e. Tex. (J. setaceus sensu Auctt. non Rostk.) the typical floating state of J. subtilis.
2. J. roemerianus Scheele Needle Rush Fig. 413 10. J. dichotomus Ell. Fig. 416
Saline or brackish marshes. Coastal. s. N.J. and Md. s. to Fla., Damp to wet sands and peats. Coastal plain, Mass. s. to Fla.,
w. to Tex. w. to Tex. and Mex.; inland n. to Okla.
perianth - - - t - H t t
involucral
leaf
a
c
Fig. 413. funcus roemerianus: a. inflorescence (PB); b. perianth and capsule (Beal); c. seed (Beal).
funcus effusus: d. habit (F); e. inflorescence (F); f. perianth and capsule (Beal); g. seed (Beal).
]01
capsule - - - H , -
perianth
involucral
leaf
l
T
capsule - - - + - - -
102
.Q i.~
, ;;!--
r.,. .,. . . . '. . . .
,~~t
. g
blade
Fig. 415. Juncus coriaceus: a. habit; b. perianth and capsule; c. seed; d. leaf sheath
(G&W).
Juncusfiliformis: e. habit; f. perianth and capsule; g. seed (HeOT).
103
tail
II!
nI l
Fig. 416. funcus dichotomus: a. habit; b. perianth and capsule; c. seed; d. leaf sheath (G&W).
funcus vaseyi: e. habit (HeOT); f. inflorescence (PB); g. perianth and capsule (HeOT); h. seed (HeOT);
i. leaf sheath (HeOT).
104
Fig. 417. 'uncus pelocarpus: a. habit. terrestrial form (F); b. habit. submersed plant (F); c. branch with bulblets
(F); d. capsule and perianth (PB); e. seed (PB).
'uncus repens: f. habit. terrestrial form (PB); g. habit. submersed form (PB); h. perianth and capsule
(G&W); i. seed (G&W); j. leaf sheath (G&W).
lOS
capsule
0. ,"
.
.'
','
Fig. 41 8.
Jo nM J,",i.""
funcus bufonius: a.
li" e.hah,
bit: b. c. pe ria nt h
b it; f. pc '",n th ",an d capsule; d. seed (H T
d cu P' " Ie; g. "e d;
'co,
b.
eO ).
,h o, tb . two vieW'
(H CO'j')
10 6
,....--- perianth
Fig. 419. Juncus marginatus var. biflorus: a. inflorescence; b. perianth and capsule; c. seed (PB).
Juncus bulbosus: d. habit (Gleason); e. habit (PB); f. perianth and capsule (Gleason).
107
a
Fig. 420. Juncus marginatus var. marginatus: a. habit; b. perianth and capsule; c. seed (NYS Museum).
108
Juncaceae / Rush Family 109
17. ,. gerardii Loisel. Black-grass Fig. 421 Me., sw. Que., Vt., and w. Mass. w. to Sask. and Wash., s. to Ala.,
Salt marshes and disturbed saline habitats. Coastal, Nfld. and Tex., Calif., and n. Mex. This species is similar to J. nodosus, but
Que. s. to Va.; locally inland w. to N.Y., n. Ky., Ont., s. Man., se. tends to be a taller, coarser plant, and its heads have a some-
N.D., and Mo.; Colo. and Utah; sw. B.C. and nw. Wash. (J. gerardii what more bristly appearance.
var. pedicellatus Fern.)
31. ,. megacephalus M. A. Curtis Fig. 427
18. J. compressus Jacq. Freshwater to brackish marshes, peats, sands, marly ponds, and
Disturbed saline sites, highway and railroad ditches, and some- ditches. Coastal plain, se. Va. s. to Fla., w. to Tex.
times invading salt marshes. Very local, w. Nfld., C.B.I., St. Law-
rence Seaway, Que., e. Me., c. N.Y., Ont., e. Wisc., nw. Minn., 32. ,. brachycarpus Engelm. Fig. 425
Man., Wyo., w. Mont., and nw. Utah; intro. from Eur. Wet sands and peaty soils. Ne. Ohio and s. Ont. w. to s. Mich.,
Ill., Minn., Mo., and Okla., s. to Ga. and Tex.; locally in e. Mass.
19. J. tenuis Willd. Fig. 422 and 1.1., N.Y.
Damp, wet, or dry open soil of roadsides, thickets, and swamps.
Throughout N.Am.; subcosmopolitan. The presence of well- 33. ,. nodatus Cov. Fig. 428
developed auricles is critical to separate J. tenuis from J. dudleyi. Shallow water, shores, sloughs, peats, and ditches. Ind. w. to Ill. ,
(J. tenuis var. anthelatus Wieg.; J. tenuis var. williamsii Fern.) Mo., and Kans., s. to Ala., Miss., La., and Tex.
20. ,. dudleyi Wieg. Fig. 422 34. J. elliottii Chapm. Fig. 428
Damp to dry soil, swamps, and lakeshores, often in calcareous Wet sands, swales, peats, and pond shores. Coastal plain, Del. s.
soil. Nfld. w. to B.C., s. to N.E., N.J., Pa., Va., Tenn., Mo., Okla., to Fla., w. to se. Tex.
Ariz., Calif., and n. Mex. (J. tenuis var. dudleyi (Wieg.) F. J. Herm.)
35. ,. acuminatus Michx. Fig. 429
21. J. caesariensis Cov. Fig. 423 Marshes, and shores of ponds, lakes, and streams. N.S. w. to
Sphagnous swamps and mossy woods. Very rare; N.S.; s. N.J. s. Ont. and Minn., s. to Fla., Tex., and n. Mex.; B.C. s. to Calif.
to Md. and se. Va.
36. J. debilis Gray Fig. 430
22. J. brachycephalus (Engelm.) Buchenau Fig. 424 Marshes, shores of acid streams, and wet clearings. Mass., R.I.,
Calcareous shores, marshes, and meadows. Nfld. and N.S. w. to and Conn. w. to N.Y., Ky., Tenn., and Mo., s. to n. Fla. and e. Tex.
Ont., s. to w. N.E., n. N.J" Pa., Ind., and Ill.
37. ,. diffusissimus Buck!. Fig. 430
23. J. canadensis J. Gay Fig. 421 Wet clay and sand, marshy shores, ditches, and clearings, often
Marshes, swamps, meadows, peats, and saline and brackish in shallow water. Coastal plain, se. Va. s. to n. Fla., w. to Tex.;
marshes. Nfld. w. to s. Ont. and Minn.; Ia. and Neb.; coastal inland n. to s. Ind. and Tenn.,; w. to Kans. and Okla. This species
plain s. to Fla., w. to La.; inland n. to Tenn. and Mo. Similar to is similar to J. debilis but with a more diffuse inflorescence.
,. brachycephalus and J. brevicaudatus, but usually a larger plant.
38. J. subtilis Mey. Fig. 421
24. J. subcaudatus (Engelm.) Cov. & Blake Fig. 425 Shores of ponds and streams, often floating in currents. Green!.,
Bogs, mossy woods, and pond margins. Nfld., N.S., Mass., and Lab., and Nfld. w. to s. Que. and s. Ont., s. to n. Me.
s. Conn. w. to Pa., W.Va., and Mo. s. to Ga. and Tenn.
39. J. abortivus Chapm. Fig. 428
25. J. brevicaudatus (Engelm.) Fern. Fig. 425 Wet shores of lakes, ponds, and streams, often in acid waters of
Muddy, sandy, and wet shores. Lab. and Nfld. w. to n. Alta., s. to flowing streams. Coastal plain, se. Va. s. to Fla.
N.B., e. N.Y., Pa., mts. of N.C. and Tenn., s. Ont., Mich., n. Ill.,
and Minn. 40. J. militaris Bigel. Fig. 427
Shallow acid waters, sandy gravelly, and peaty shores. Nfld. and
26. J. bulbosus L. Fig. 419 N.S. s. to N.E., N.Y., e. Md., and Del.; local in Ont. and Mich.
Sandy and peaty shores of ponds and streams, often floating.
Nfld., St. P. et Miq., and Sable I., N.S.; B.C.; intro. from Eur. 41. J. alpinus Viii. Fig. 431
Wet sandy and gravelly shores, often calcareous. Greenl. and
27. J. validus Cov. Fig. 424 Nfld. w. to Alta. and B.C., s. to c. Me., w. Vt., N.Y., n. Ohio, n.
Swales, damp prairies, marshy shores, damp fields, and ditches. Ind., Ill., Mo., Neb., and Wash. Hiimet-Ahti (1980) believes that
Se. Va. s. to Fla., w. to Tex.; inland n. to w. Tenn., Mo., and Okla. the correct name is J. alpinoarticulatus Chaix, but notes that it is
not clear to her that Chaix meant his taxon to apply to this spe-
28. J. scirpoides Lam. Fig. 426 cies. A natural hybrid between this species and ,. torreyi has
Damp soil, pools, wet pinelands, peats, and pond shores. Coastal been described from a local area along Lake Erie in Ohio as ,.
plain, s. and c. N.Y. s. to Fla., w. to Tex.; n. inland to Ky., n. Ind., Xstuckeyi Reinking (Reinking, 1981). (J. alpinoarticulatus Chaix)
Ill., sw. Mich., s. Mo., and Okla. (J. scirpoides var. meridionalis
Buchenau) 42. J. articulatus L. Fig. 431
Wet sandy and boggy shores, and pond and stream shores. Nfld.
29. ,. nodosus L. Fig. 426 w. to B.C., s. to N.E., N.J., N.Y.. W.Va .. Ohio, n. Ind .. Minn., Utah,
Moist soil, swamps, and gravels. Nfld. w. to Alask., s. to N.E., Nev., and Oreg. This species is very similar to J. alpin us, and
n. N.J., w. Va., Ohio, Ill., Iowa, Neb .. Tex .. N.M .. Nev., and Calif. more than a single character is needed to distinguish them. (J.
articulatus var. obtusatus Engelm.)
30. J. torreyi Cov. Fig. 423
Moist to wet sandy areas, river and stream banks, and ditches.
tail
Fig. 421. Juncus subtilis: a. habit; b. fascicle of leaves; c. perianth and capsule (Gleason).
Juncus gerardii: d. habit (F); e. portion of inflorescence (HCOT); f. seed (HCOT).
Juncus canadensis: g. inflorescence; h. perianth and capsule; i. seed (Beal).
no
a
c I
auricle
Fig. 422. Juncus tenuis: a. inflorescence (HCOT); b. perianth and capsule (G&W); c. seed (G&W); d. leaf sheath
(G&W).
Juncus dudleyi: e. habit; f. perianth and capsule; g. seed; h. leaf sheath, two views (HCOT).
111
h f
d
rhizome
Fig. 423. funcus torreyi: a. habit. with tuberous rhizome (HeaT); b. inflorescence (F); c. perianth and capsule
(HeaT); d. seed (HeaT); e. leaf sheath (HeaT).
funcus caesariensis: f. inflorescence; g. perianth and capsule; h. seed (PB).
112
c
Fig. 424. funcus validus: a. habit (G&W); b. inflorescence (PB); c. perianth and capsule (G&W); d. seed (G&W).
funcus brachycephalus: e. inflorescence; f. perianth and capsule (PB).
113
capsule
OJ
perianth
Fig. 425. funcus brevicaudatus: a. inflorescence (F); b. perianth and capsule (Beat); c. inflorescence. modificd as
insect gall (F).
funcus subcaudatus: d. inflorescence; e. perianth and capsule; f. seed (Beal).
funcus brachycarpus: g. inflorescence; h. perianth and capsule; i. seed (Beal).
114
stigma
11 5
"
"
Fig,427. funcus megacephalus: a, habit (G&W); b. perianth and capsule (Beal); c. seed (G&W); d. leaf sheath
and leaf section. showing septae (G&W).
funcus militaris: e. habit (F).
116
g
Fig. 428. Juncus elliottii: a. habit; b. perianth and capsule; c. seed; d. leaf sheath (G&W).
Juncus nodatus: e. inflorescence; f. perianth and capsule; g. seed (PB).
Juncus abortivus: h. inflorescence; i. perianth and capsule; j. seed (Beal).
117
b
Fig. 429. funcus acuminatus: a. lower portion of plant: b. upper portion of plant: c. perianth and capsule;
d. seed; e. leaf sheath (HeOT).
118
Fig. 430. Juncus debilis: a. habit; b. perianth and capsule; c. seed (G&W).
Juncus diffusissimus: d. habit; c. perianth and capsule; f. seed (G&W).
119
~r~
~' ..~
J
I
i I
auricle
~ 1'
h
1.1
I \"
a
Fig, 431. Juncus alpin us: a, habit; b, portion of inflorescence; c. perianth and capsule; d, seed; e, leaf sheath
(HCOT).
Juncus articulatus: f. habit; g. perianth and capsule; h. seed; i. leaf sheath (HeOT).
120
Cyperaceae / Sedge Family 121
The sedges constitute a very large family. superficially resembling the grasses. but they are easily distinguished by several vegetative
and reproductive features (fig. 432): stems usually 3-angled in cross-section. solid (except Dulichium and some Carex); leaves 3-
ranked. sheathing stem. with sheaths closed; flowers in axils of overlapping scales. each having a single pistil. with 2 or 3 stigmas
and 1-3 stamens (unisexual in Carex and ScIeria); perianth absent or reduced to bristles and/or scales; fruit a 2-sided (biconvex) or
3-sided (3-angled) achene. The fruits of sedges are important food for wildfowl.
In this family divergent opinions exist as to the status of genera. A few genera (i.e . Dichromena. Psilocarya. and Hemicarpha)
traditionally recognized by most authors are not recognized here.
REFERENCES: Mackenzie. 1931-1935; Naczi et al.. 1986; Tucker. 1987; Svenson. 1957.
1. Carex (Sedge)
Perennial herbs. arising from creeping rhizomes or cespitose; stems usually 3-angled; leaves grass-like. 3-ranked; flowers unisexual.
staminate and pistillate flowers in separate spikes (fig. 433i). different parts of same spike. or scattered. mixed in same spike; fruit a
3-angied or biconvex achene enclosed in a perigynium.
r--T---::::>" stigmas
style
anther}
pistil stamen
filament
base of
blade
b
stem
stigmas ------'
remains of scale - - -
sheath filaments
(sometimes present)
sheath
attached bristle
to stem
here
a
c
Fig. 432. Structure ofCyperaceae (sedges): a. section of stem with leaf base, generalized; b. flower, generalized;
c. achene, generalized; d. spike/spikelet, generalized (F).
122
13. Lower leaf sheaths glabrous; perigynia somewhat inflated, obovoid with abrupt beak.
13. Lower leaf sheaths scabrous to hispidulous; perigynia flattened, elliptic, tapering to minute beak.
124 ANGIOSPERMS
1 7. Stems not arising from last year's leaves, bearing bladeless sheaths at base; leaf bases distinctly fringed.
21. Ligules rounded, about as long as wide or (fig. 43ge); pistillate spikes 2-10(15) em long .... " 15. C. emoryi
21. Ligules acute, longer than wide (fig. 4390); pistillate spikes 2-6(11) em long.
22. Stems in dense tussocks; sheaths smooth on side opposite blade; blades not glaucous ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 16a. C. stricta var. stricta
22. Stems stoloniferous, sometimes forming large colonies, but not tussocks; sheaths minutely scab-
rous on side opposite blade; blades glaucous. . . . . . . . . . . . . . . . . . . . . . .. 16b. C. stricta var. strictior
5. Lateral spikes sessile, short; terminal spike usually partly pistillate.
23. Plants with stems usually solitary or few, arising from stolons or olTshoots from stolon.
24. Perigynia wingless, lacking sharp margins. not flattened in cross-section.
25. Spikes. especially lower ones. somewhat separated; terminal spike with flowers pistillate on upper portion.
staminate on lower ............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37. C. mackenziei
25. Spikes crowded. often appearing as a single head; terminal spike with flowers staminate on upper portion.
pistillate on lower . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. C. chordorrhiza
24. Perigynia winged or with sharply edged margins (fig. 440h). flattened in cross-section ............ 18. C. sartwellii
23. Plants loosely to densely tufted.
26. Spikes with staminate flowers on upper portion (androgynous).
27. Stems thick. sharply angled. soft, easily compressed.
28. Perigynia 3-5 mm long; beak shorter than or as long as body. . . . . . . . . . . . . . . . . . .. 19. C. alopecoidea
28. Perigynia 4.5-9 mm long; beak longer than body.
29. Perigynia 6-9 mm long. base dilated. disk-shaped (fig. 441b); thin portion of leaf sheath
strongly purple-dotted. ventral band not transversely wrinkled ................. 20. C. crus-corvi
29. Perigynia 4-5(6) mm long. base not dilated; thin portion ofleaf sheath not purple-dotted. ven-
tral band strongly transversely wrinkled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21. C. stipata
27. Stems very slender. not sharply angled, firm. not easily compressed.
30. Inflorescences simple. 1-5 perigynia per spikelet; perigynia essentially beakless .......... 22. C. disperma
30. Inflorescences with spikelets on lateral branches. several to many perigynia per spikelet; perigynia
beaked.
31. Perigynia more or less biconvex; pistillate scales acute to slightly cuspidate or mucronate; leaf
sheaths not transversely wrinkled.
32. Leaves (2.5)5-8 mm wide; inflorescences 7-18 em long; perigynia abruptly beaked. taper-
ing basally to short stipe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. C. decomposita
32. Leaves 1-3 mm wide; inflorescences 1-6.5(10) cm long; perigynia gradually tapering to
beak. rounded to truncate at base.
33. Leaf sheath pale at summit; mature perigynia olive-green to blackish. inner face
convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24. C. diandra
33. Leaf sheath bronze-colored at summit; mature perigynia light brown to brown. inner
face flat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 25. C. prairea
31. Perigynia plano-convex; pistillate scales long-awned; leaf sheaths transversely wrinkled
(fig. 443f) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26. C. vulpinoidea
26. Spikes with pistillate flowers on upper portiun (gynaecandrous).
34. Perigynia with thin-winged margins; achene not filling perigynium tu margin; luwer portion of perigyn-
ium not spongy-thickened; stems hollow.
35. Pistillate scales obtuse, acute, or short-acuminate. lacking awned tip.
36. Perigynia more than 2 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27. C. suberecta
36. Perigynia not more than 2 mm wide.
37. Perigynia about (3)3.5-5 times as long as wide. appressed-ascending.
38. Perigynia 1.3-2 mm wide, 4.1-5.5 mm long . . . . . . . . . . . . . . . . . . . . . . . . 28. C. scoparia
38. Perigynia 0.6-1.1 mm wide. 2.7-4(4.5) mm long ................... 29. C. crawfordii
37. Perigynia 1.5-2.5(3.5) times as long as wide; ascending to spreading or somewhat re-
curved.
39. Perigynia ascending to slightly spreading; spikes in a dense, stilT inflorescence ... 30. C. bebbii
39. Perigynia widely spreading to somewhat recurved; spikes mostly in a more open inflo-
rescence with lowest 2 spikes slightly separated . . . . . . . . . . . . . . . . . . . .. 31. C. cristatella
46. Leaves glaucous, 22.7(3.7) mm wide; spikes ellipsoidal to subglobose, with
10many loosely spreading perigynia . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. C. canescens
46. Leaves green, 12 mm wide; spikes elongate, with (3)510 ascending to
appressed perigynia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38. C. brunnescens
126 ANGIOSPERMS
72. Leaves flat, not wiry; staminate spikes 2 or more; pistillate scales lanceolate, acute
to acuminate.
73. Perigynia firm; beak minutely roughened ....................... 63. C. bullata
73. Perigynia membranous; beak smooth ........................ 64. C. vesicaria
65. Perigynia 11-20 mm long, with 15-20 nerves; achene 2.5-6 mm long.
74. Achene as wide or wider than long, with conspicuously knobbed angles (fig. 462c) ...... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65. C. lupulijormis
74. Achene about two-thirds as wide as long, with obscurely knobbed angles ......... , 66. C. lupulina
64. Styles deciduous (at least upper portion with stigmas), articulated with achenes; perigynia entire or
minutely toothed at tip.
75. Perigynium beakless or with short, obscure beak 0.5 mm or less long.
76. Lower bracts of pistillate spikes sheathless or with short sheath.
77. Terminal spikes pistillate on upper portion; perigynia many-nerved .......... 67. C. buxbaumii
77. Terminal spikes staminate throughout; perigynia few-nerved.
78. Scales as wide or wider than and barely longer than perigynia; pistillate spikes 0.5-
2.5 em long.
79. Stems with 3 sharp angles; leaves involute or flat, not thick or stiff.
80. Leaves involute, 1-2 mm wide; scales ovate to subcircular, as wide as peri-
gynia, persistent .................................... 68. C. limosa
80. Leaves flat, 1-3 mm wide; scales lanceolate, narrower than perigynia,
readily deciduous .................................. 69. C. paupercula
79. Stems with 3 rounded angles; leaves flat, thick, and stiff. . . . . . . . . . .. 70. C. rariflora
78. Scales narrower than and much longer than perigynia; pistillate spikes 2-5.5 cm
long .................................................. 71. C. barrattii
76. Lower bracts of pistillate spikes with well-developed sheath.
81. Leaves whitish, glaucous, becoming involute with age; perigynia whitish, glaucous .. 72. C. livida
81. Leaves green to slightly glaucous, flat except at apex of older leaves; perigynia green, not
glaucous .................................................. 73. C. tetanica
75. Perigynium with distinct beak 0.5 mm or more long.
82. Perigynia ascending; pistillate spikes 1 or 2; plants stoloniferous, forming large colonies ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 47. C. striata
82. Perigynia reflexed or spreading; pistillate spikes 2-6; plants forming tufts.
83. Perigynia 2-3.5 mm long, beak a fourth to half as long as body. . .. 74a. C. viridula ssp. viridula
83. Perigynia 3-6 mm long, beak half as long as or longer than body.
84. Stems obtusely angled at tip; perigynia obovoid, abruptly beaked ............ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 74b. C. viridula ssp. brachyrrhyncha
84. Stems acutely angled at summit; perigynia lanceolate-ovoid to subulate, gradually
tapering to beak.
85. Perigynia 3-4.5(5) mm long, beak smooth .................... 75. C. cryptolepis
85. Perigynia 4-6 mm long, beak often serrulate toward tip (fig. 468b) ....... 76. C. flava
ortion
k f
Fig. 433. Carex pauciflora: a. habit; b. inflorescence; c. perigynium; d. achene within perigynium (HCOT).
Carex leptalea: e. habit; f. inflorescence; g. perigynium; h. achene (G&W).
Carex torta: i. inflorescence; j. perigynium (F).
Carex salina: k. inflorescence; l. perigynium; m. achene with constriction; n. pistillate scale (Mac-
kenzie).
128
c
h g
Fig. 434. Carex mitchelliana: a. inflorescence; b. perigynium; c. achene; d. pistillate scale (Mackenzie).
Carex paleacea: e. inflorescence; f. perigynium; g. achene with constriction; h. pistillate scale .(Mac-
kenzie).
129
130 ANGIOSPERMS
7. e. lyngbyei Hornem. Fig. 435 above the water in shallow pond margins. P.E.1. and N.B. w. to
Salt marshes, beach gravels, and coastal bluffs. Atlantic coast, ant.. Ohio, Ill., and Minn., s. to N.E., N.C., and sw. Ky.
Green!' and Lab. s. to Anticosti I. and e. Que.; Pacific coast,
16b. C. stricta var. strictior (Dew.) Carey Fig. 439
Alask. s. to Calif.
Calcareous swamps, meadows, and low woods. Que. w. to ant.
and Minn., s. to N.S., N.B., N.C., Tenn., and Mo.; more common
8. C. crinita Lam.
westward. (C. strictior Dew.)
8a. e. crinita var. crinita Fig. 435
Swales and damp thickets. Nfld. w. to n. Man., s. to N.S., N.E., 17. C. chordorrhiza Ehrh. ex L. f. Fig. 440
Md., n. Ga., Tenn., Mo., e. Okla., and e. Tex. Sphagnum bogs, peaty ground, and quagmires. S. Baffin I. w. to
Alask., s. to Ntld., e. Que., Me., sw. Vt., Mass., C. N.Y., n. II!., n.
8b. e. crinita var. brevicrinis Fern.
Iowa, and Sask.
Wooded swamps and bottomlands. Coastal plain, s. N.E. s. to
Fla., w. to e. Tex.; inland n. to Ky. and Mo.
18. C. sartweIlii Dew. Fig. 440
Calcareous bogs, marshes, swales. and lakeshores. S. Que. W. to
9. C. gynandra Schwein. Fig. 436
James Bay, N.WT., and n. B.C .. s. to WVa., Ind., Ill., Mo., Neb.,
Swales and damp thickets. Nfld. w. to Wisc. and ne. Minn., S. to
and Colo.
N.B., Md., N.C., Tenn., and n. Ga. (e. crinita Lam. var. gynandra
(Schwein.) Schwein. & Torr.)
19. C. alopecoidea Tuckerm. Fig. 440
Marshes, meadows. and swales. S. Que. w. to Minn., s. to N.B.,
10. C. aurea Nutt.
n. N.J., Ind., and Iowa.
Wet meadows, damp shores, bogs, swamps, and low woods.
Nfld. W. to Alask., S. to N.E., n. Pa., n. Ohio, n. Ind., n. Ill., Minn.,
20. C. crus-corvi ShuttIw. ex Ktze. Fig. 441
Neb., N.M., and Calif.
Marshes, sloughs, swamps, and riverbanks. Coastal plain, Va. s.
to Fla., W. to e. Tex.; inland n. to Ohio. s. ant., s. Mich .. S. Minn.,
11. e. garberi Fern. Fig. 436
e. Okla. and e. Neb.
Calcareous sands, and marly or gravelly shores. Que. W. to nw.
ant. and ne. Minn., S. to N.Y., Ohio, Mich., and nw. Ind.; most
21. C. stipata Muh!. ex Willd. Fig. 441
abundant along shores of Great Lakes and vicinity.
Wet meadows, marshy shores, swampy woods, floodplains, low
woods, and damp shaded ground. S. Lab. and Nfld. w. to S.
12. C. aquatilis Wahlenb.
Alask., s. to Fla., Tex., N.M., and Calif. A stouter growth form of
12a. C. aquatilis var. aquatilis Fig. 437 the coastal plain and Gulf states has been called var. maxima
Shallow pools, wet meadows, marshes, shores of ponds, lakes, Chapm. (e. uberior (e. Mohr) Mackenz.)
and streams. Green!' and Lab. W. to Alask., S. to Nfld., C.B.I., nw.
Mich., Minn., N.M., Ariz., and Calif. 22. C. disperma Dew.
Sphagnum bogs, cedar swamps, and mossy woods. Lab. and
12b. e. aquatilis var. altior (Rydb.) Fern. Fig. 437
Nfld. W. to N.WT. and Alask.. s. to N.E .. n. N.J,. Pa., n. Ind.,
Calcareous waters, marshes, bogs, and ditches. Nfld. and e. Que.
Wisc., Minn., S.D., N.M., and Calif.
W. to B.e., S. to n. N.J., Ind., Wisc., Mo., Neb., and Oreg. (C. aqua-
tilis var. substricta Kuek.; C. substricta (Kuek.) Mackenz.)
23. C. decomposita Muh!. Fig. 441
Wooded swamps and pond margins, often on rotting logs. N. Y.
13. C. lenticularis Michx. Fig. 438
W. to s. Mich., s. to Fla. and e. Tex.
Wet gravelly and sandy shores, and meadows. Lab. and Nfld. W.
to N.WT., s. to Mass., ne. N.Y., ant., n. Mich., ne. Minn., and
24. C. diandra Schrank Fig. 442
Man. Our iaxon is var. lenticularis.
Bogs and peaty swamps, especially calcareous sites. Lab. and
Nfld. w. to Yuk. and Alask., S. to N.]., Pa .. Md., Ill.. Neb., Colo.,
14. C. nigra (L.) Reichard Fig. 438
and Calif.
Wet meadows, swales, salt marshes, and wet gravels and rocks,
especially disturbed siies. Chiefly near the coast; Green!', se.
25. C. prairea Dew. Fig. 442
Lab., and Ntld. W. along St. Lawrence R. to se. Que. and n. Vt.,
Calcareous bogs, meadows, and wet thickets. Que. W. to Alta.,
s. to Mass., R.I., and Conn.; n. OnL, n. Mich. Standley (198 7a)
S. to W. N.E., n. N.J" IIl., and Neb.
notes that C. nigra is morphologically variable in North America
and that infraspecific taxa should not be recognized. Further,
26. C. vulpinoidea Michx. Fig. 443
North American populations have not diverged significantly
Wet. low ground, marshes, shores of rivers and streams. and
from those in Europe to warrant the recognition of them as a
near margins of wet wooded areas. Ntld. W. to S. B.C .. s. through-
separate taxon. (c. nigra var. strictijormis (Bailey) Fern.)
out U.S.
15. C. emoryi Dew. Fig. 439
27. C. subaecta (Olney) Britt. Fig. 442
Swamps, shores, and river margins. N.Y. and N.J. w. to ant.,
Calcareous soils of meadows, marshy shores, ditches, and prai-
Ind., Wisc., Minn., and Man., s. to Fla., Tex., Colo., and N.M.
ries. S. ant. w. to Minn .. s. to w. Va., Ind .. Ill., and Mo.
16. C. stricta Lam.
28. C. scoparia Schkuhr ex. Willd. Fig. 444
16a. e. stricta var. stricta Fig. 43 Y Open swamps, wet meadows, marshy shores, thickets. and
Acid swamps, meadows, and low woods. Large tussocks of this moist Lo dry open sites. Nfld. w. to B.C .. s. to S.C .. Tenn., Ark.,
species are distinctive in meadows and, at times. stand well N.M., Ariz., and Oreg.
d
er in
d. scale of pistillate flow
b. per igy niu m; c. ach ene with constriction; ke (Mason).
ei: a. habit; portion of spi
Fig. 43 5. Carex lyngby n of spike; e. scale of pistillate flower in lower ction; i. pistillate
Upper portio enc e; g. per igy niu m; h. ach ene with constri
var . crinita: f. inf loresc
crin
ex (M
Carle
sca ackitaenz ie).
131
h
c awn
g
b
Fig. 436. Carex gynandra: a. inflorescence; b. perigynium; c. achene with constriction; d. pistillate scale (Mac-
kenzie).
Carex garberi: e. habit; f. perigynium; g. achene; h. pistillate scale (Mackenzie).
132
a
( /
scale
'I' )1
,,;./
~'
Fig. 437. Carex aquatilis var. aquatilis: a. inflorescence; b. pistillate spike; c. perigynium (HCOT).
Carex aquatilis var. altior: d. inflorescence; e. perigynium (F).
133
d
b a
134
a
/
n g h
Fig. 439. Carex emoryi: a. inflorescence; b. perigynium; c. achene; d. pistillate scale; e. ligule (Mackenzie).
Carex stricta var. stricta: f. inflorescence (F); g. perigynium (Mackenzie); h. achene (Mackenzie);
i. pistillate scale (Mackenzie); j. section of stem with sheath (F).
Carex stricta var. strictior: k. inflorescence; l. perigynium; m. achene; n. pistillate scale; o. ligule (Mac-
kenzie).
135
c
f
h
6
ligule
1[
.::
...
p
l/
Jf n
m
Fig. 440. Carex chordorrhiza: a. upper portion of plant; h. inflorescence; c. perigynium; d. achene; e. pistillate
scale (Mackenzie).
Carex sartwellii: f. inflorescence; g. single spike; h. perigynium; i. achene; j. pistillate scale (Macken-
zie).
Carex alopecoidea: k. inflorescence; l. single spike; m. perigynium; n. achene; o. pistillate scale; p.
ligule (Mackenzie).
136
b
c
d
;)
.
)
~'.::.
\_0 j
Fig.44l. Carex crus-corvi: a. inflorescence (Mackenzie); b. perigynium (G&W); c. achene (Mackenzie); d. pistil-
late scale (G&W).
Carex stipata: e. inflorescence; f. ligule (F).
Carex decomposita: g. inflorescence (Mackenzie); h. single spike (Mackenzie); i. perigynium (G&W);
j. achene (Mackenzie); k. pistillate scale (G&W); I. ligule (Mackenzie).
137
ligule
e d
staminate
flowers
m n
q r
p
Fig. 442. Carex diandra: a. inflorescence (F); h. single spike (Mackenzie); c. perigynium (Mackenzie); d. achene
(Mackenzie); e. pistillate scale (Mackenzie); f. ligule (Mackenzie).
Carex prairea: g. h. inflorescences; i. perigynium; j. achene; k. pistillate scale; I. ligule (Mackenzie).
Carex suberecta: m. inflorescence; n. single spike; o. perigynium; p. achene; q. pistillate scale; r. lig-
ule (Mackenzie).
138
Fig. 443. Carex vulpinoidea: a. habit (HCOT); b. inflorescence (G&W); c. perigynium (G&W); d. scale variations
(G&W); e. achene (G&W); f. leaf sheath. two views (HCOT).
139
a
y
c
e d
k m
Fig. 444. Carex scoparia: a. inflorescence; b. single spike; c. perigynium; d. achene; e. pistillate scale; f. ligule
(Mackenzie) .
Carex crawfordii: g. habit (HCaT); h. inflorescence (HCaT); i, j. perigynium, two views (HeaT);
k. achene (Mackenzie); 1. pistillate scale (Mackenzie); m. ligule (Mackenzie).
140
Cyperaceae / Sedge Family 141
29. C. crawfordii Fern. Fig. 444 41b. C. atlantica ssp. capillacea (Bailey) Reznicek Fig. 449
Wet meadows. swamps. shores. and wet to dry open ground. Swamps. bogs, wet. acid thickets. floodplains. and low woods.
Nfld. w. to B.C .. s. to n. N.J.. mts. of Tenn .. Mich.. Minn., Sask., Chiefly coastal plain, N.S. s. to N.J., S.C .. and Fla., w. to La. and
Ida . and Wash. e. Tex.; inland w. to s. Que . N.Y.. Pa .. Ont.. Ohio. Mich.; sw.
Tenn. This taxon has long been known as C. howei or C. interior
30. C. bebbii (Bailey) Fern. Fig. 445 var. capillacea. Sometimes ssp. capillacea intergrades with ssp. at-
Wet meadows, swales, and shores, chiefly calcareous soils. Nfld. lantica in non-glaciated portions of their ranges (Reznicek and
w. to B.C . s. to N.E . n. N.J.. n. Ohio, n. Ind., n. Iowa. Neb . Colo . Ball. 1980). (c. interior var. capillacea Bailey; C. howei Mackenz.)
Ida., and Wash.
42. C. wiegandii Mackenz. Fig. 448
31. C. cristatella Britt. Fig. 445 Sphagnum bogs. boggy thickets, lakeshores, and wet peaty soils.
Open swamps. marshes, wet meadows, shores, thickets, and wet Rare; Nfld. w. to Ont.. s. to Me. and n. Mass.; n. Pa.
woods. Sw. Que. w. to N.D., s. to N.B., Md., Ky., Mo., and Neb.
43. C. interior Bailey Fig. 450
32. C. alata Torr. Fig. 446 Shores of lakes. ponds. and streams, ditches. wet meadows. fens.
Marshes, swamps, bogs. shores. meadows, and wet woods. wet prairies, and swamps; chiefly calcareous sites. Nfld. w. to sw.
Mass. w. to N.Y. Pa., s. Ont., and Mich . s. to Fla .. s. Mo .. and N.W.T. and s. Alask., s. to N.J. Ohio. Mo . Kan . Colo. n. Ariz.,
Tex. Nev.. and n. Calif.; Mex. (C. interior var. keweenawensis F. J.
Herm.; C. interior ssp. charlestonensis Clokey; C. stellulata var.
33. C. straminea Wil\d. Fig. 446 scirpoides (Wil\d.) Dar!.)
Swamps and marshes. Mass. w. to Mich., s. to Md. (c. richii
(Fern.) Mackenz.) 44. C. echinata Murr. Fig. 450
Wet acid soils, sphagnum bogs, boggy meadows. and sandy lake
34. C. hormathodes Fern. Fig. 446 and river shores. Green!' and Lab. w. to Ont. and Minn., s. to
Saline and freshwater marshes. Coastal. w. Nfld., St. P. et Miq., N.E., N.].. mts. to N.C . Pa .. n. Ind., and n. Iowa; Sask. w. to B.C..
and e. Que. s. to Va.; disjunct station. n. Ind. s. in mts. to Wyo .. Colo., Utah, and Calif. (c. angustior Mackenz.;
C. angustior var. gracilenta Clausen & Wahlenb.; C. muricata var.
35. C. trisperma Dew. cephalantha (Bailey) Wieg. & Eames; C. cephalantha (Bailey)
Coniferous bogs and swamps, mossy woods, and boggy hollows. Bickn.; C. leersii Wil\d.; C. josselynii (Fern.) Mackenz. ex Pease;
Lab. and Nfld. w. to N.W.T., s. to N.B.. n. N.J.. Md., mts. to N.C. C. laricina Mackenz. ex Bright; C. stellulata Wil\d.)
and e. Tenn., Ohio. n. Ind., II!.. Minn., and Alta.
45. C. lanuginosa Michx. Fig. 451
36. C. tenuiflora Wahlenb. Shores, meadows, swales, and riverbanks; often in shallow wa-
Sphagnum bogs, rich woodland fens, mossy woods. and peaty ter. Que. w. to B.C., s. to N.E., Va . Tenn . Ark., Tex . N.M., and
shores. Lab. and Nfld. w. to N.W.T. and Alask.. s. to c. Me. s. Calif.
w. Mass., N.Y., Mich .. Wisc. Minn .. Man., Sask., and Alta.
46. C. lasiocarpa Ehrh. Fig. 451
3 7. C. mackenziei Krecz. Fig. 447 Bogs. streams, swales, and shallow water. Nfld. w. to B.C .. s. to
Saline shores and marshes. Coastal; Green!' and Lab. w. to Hud- n. N.J., Pa., n. Ohio, n. Ill. , ne. Iowa, Man., Sask., Ida., and
son Bay and Alask., s. to N.S. and Me. Wash. The North American plants have been segregated as var.
americana Fern.
38. C. canescens L. Fig. 447
Swamps, bogs, ponds, and stream margins. Green!. and Lab. w. 47. C. striata Michx. Fig. 453
to Alask., s. to N.E., N.J., Va., Ohio, Ind., Minn., Wyo . Ida., Ariz. Pine barren swamps and pond margins. Coastal plain, se. Mass.
and Calif. Sometimes var. subloliacea (Laestad.) Hartm. and var. s. to L.r.. N.Y., N.J., S.C., and n. Fla. Our taxon, var. brevis Bailey.
disjuncta Fern. are recognized. is distinguished from the southern taxon by having glabrous
perigynia. Reznicek and Catling (1986) have determined C. stri-
39. C. brunnescens (Pers.) Poir. ata to be the correct name for this plant, widely referred to as C.
Low woods. margins of bogs and swamps, and wet turfy hollows walteriana Bailey.
and slopes. Lab. and Nfld. w. to N.W.T. and Alask., s. to N.B.,
N.J.. Pa., mts. to N.C. and e. Tenn., Ohio, Mich., Wisc., Minn., 48. C. trichocarpa Muh!. ex Schkuhr Fig. 451
Colo., and Oreg. Calcareous meadows, swales, marshes, wet deciduous woods,
and riverbanks. Sw. Que. and vt. w. to Ont. and Minn., s. to
40. C. exilis Dew. Fig. 448 Conn., DeL. Ohio, IlL. and n. Iowa; w. N.C.
Bogs. fens, and open wet sites. Lab. and Nfld. w. to Ont. and
n. Minn., s. to N.Y., N.J., De!., Md., n. Mich.; disjunct to N.C. 49. C. atherodes Spreng. Fig. 452
and Miss. Calcareous meadows. shores, marshes, and shallow water. S.
Me. w. to Ont.. Man., N.W.T., and Alask., s. to w. N.Y.. W.Va ..
41. C. atlantica Bailey Ohio, Wisc .. Mo., Neb., e. Mont., Utah, and Oreg.
41a. C. atlantica ssp. atlantica Fig. 449
Bogs and wet, acid thickets, and edges of forests. N.S. and N.B. 50. C. lacustris Wil\d. Fig. 453
w. to N.Y. and Mich., s. to n. Fla . se. Mo . and e. Tex.; most Swamps, ditches. marshes, swales, and river and stream mar-
abundant near the coast. (C. stellata var. conferta Chapm.; c. gins. Nfld., P.E.!., and Que. w. to Man. and Sask.. s. to N.E., Va.,
echinata var. conferta (Chapm.) Bailey; C. incomperta Bickn.; Ohio. Ill., Mo., e. Kan . and Ida. (C. riparia var. /acustris (Wil\d.)
C. atlantica var. incomperta (Bickn.) F. J. Herm.) Kuek.)
c
e
b h
Fig. 445. Carex bebbii: a. habit (HCOT); b. inflorescence (HeOT); c. perigynium (Mackenzie); d. achene (Mac-
kenzie); e. pistillate scale (Mackenzie); f. ligule (Mackenzie).
Carex cristatella: g. h. inflorescence; i. j. perigynium. two views; k. achene; I. pistillate scale (Mac-
kenzie).
142
k
e
ligule
Fig. 446. Carex alata: a. inflorescence (F); b. perigynium (G&W); c. pistillate scale (G&W).
Carex straminea: d. inflorescence; e. perigynium (F).
Carex hormathodes: f. g. inflorescences; h. single spike; i. j. perigynium. two views; k. achene; I. pistil-
late scale; m. ligule (Mackenzie).
143
h
Fig. 447. Carex canescens: a. habit (HeOT); b. perigynium (Mackenzie); c. achene (Mackenzie); d. pistillate scale
(Mackenzie) .
Carex mackenziei: e. inflorescence; f. single spike; g. perigynium; h. achene; i. pistillate scale (Mac-
kenzie).
144
e
Fig. 448. Carex exilis: a. habit; b. single spike; c, d. perigynium, two views; e. achene; f. pistillate scale; g. ligule
(Mackenzie).
Carex wiegandii: h. inflorescence; i. single spike; j, k. perigynium, two views; l. achene; m. pistillate
scale (Mackenzie).
145
c d e
'i\,
\:"
l:i 1i
h
i\\l!
L n
Fig. 449. Carex atlantica ssp. atlantica: a. habit (G&W); b. inflorescence (G&W); c, d. perigynium, two views
(Mackenzie); e. achene (Mackenzie); f. pistillate scale (Mackenzie); g. ligule (Mackenzie).
Carex atlantica ssp. capillacea: h. inflorescence; i. single spike; j, k. perigynium, two views; l. achene;
m. pistillate scale; n. ligule (Mackenzie).
146
"
pistillate
flowers
staminate
flowers
h
k
e
c
d
Fig. 450. Carex interior: a. habit (HeOT); b. single spike (Mackenzie); c. perigynium (Mackenzie); d. achene
(Mackenzie); e. pistillate scale (Mackenzie).
Carex echinata: f. habit; g. single spike; h. i. perigynium. two views; j. achene; k. pistillate scale (Mac-
kenzie).
147
stigma
I{
1
1 :.:.-.( g
\ ~, I ..
-.)
il'V:I::.'1
, d
\'
c
b
h
e
a
148
ry e
Fig. 452. Carex atherodes: a. inflorescence: b. lower portion of plant with thick rhizomes: c. perigynium:
d. achene: e. achene. cross-section: f. pistillate scale: g. staminate scale: h. ligule (Mason).
149
e
Fig. 453. Carex striata: a. habit (G&W); b. inflorescence (F); c. perigynium (G&W); d. achene (G&W); e. stami-
nate scale (G&W); f. pistillate scale (G&W); g.ligule (G&W).
Carex lacustris: h. inflorescence; i. perigynium (F).
150
Cyperaceae / Sedge Family 151
51. C. hyalinolepis Steud. Fig. 454 63. C. bullata Schkuhr Fig. 460
Calcareous or brackish swamps. shores. ditches. and wet woods. Acid swales. meadows. and bogs. Coastal plain. N.S. s. to Ga.
N.J. and se. Fa. w. to s. Ont .. se. Mich .. s. Ind .. s. II!.. and e. Neb .. (e. bullata var. greenii (Boeck!.) Fern.)
s. to Fla. and Tex. (e. lacustris var. laxiflora Dew.; C. riparia var.
impressa (Wright) Mackenz.; e. impressa Mackenz.) 64. e. vesica ria L. Fig. 461
Swamps. marshes. and swales. Nfld. w. to B.C .. s. to N.E .. Del..
52. C. michauxiana Boeck!. Fig. 455 Pa .. n. Ky.. s. Ind .. s. IlL. Minn .. Colo .. and Calif. Numerous vari-
Bogs. wet sands. and meadows. Nfld. w. to Ont.. s. to N.S .. n. eties have been described.
N.B.. w. Mass . n. N.Y.. n. Mich .. and ne. Minn.: Sask.
65. e. lupuliformis Sartwell ex. Dew. Fig. 462
53. C. folliculata L. Fig. 455 Calcareous swamps. meadows. swales. prairies. and wet woods.
Swamps. swales. bogs. pond and lake margins. stream banks. Sw. Que. and w. vt. w. to Ont. and Minn .. s. to Conn . Va . Ky..
and wet woods. Nfld. w. to Wise. s. to N.B.. mts. and piedmont and e. Tex.
to N.C .. W.Va .. and Tenn. Our taxon is chiefly the northern var.
folliculata. which passes into the southern var. australis Bailey. 66. C. lupulina Willd. Fig. 464
a coastal plain taxon ranging from s. N.J. to Fla .. w. to La .. char- Swamps. marshes. and wet woods. N.S. and s. Que. w. to s. Ont..
acterized by having shorter pistillate scales. usually long- Mich .. Wisc .. Minn .. and Neb .. s. to Fla. and Tex.
acuminate. and seldom awned.
67. C. buxbaumii Wahlenb. Fig. 463
54. C. pseudocyperus L. Fig. 454 Swamps. wet shores. and bogs. Nfld. w. to Alask .. s. to N.B..
Marshes. swamps. bogs. and shores; often in shallow water. e. Va .. N.C .. w. Ky.. Tenn .. Ark.. Kan .. Colo .. and Calif.
Nfld. w. to Sask .. s. to Me .. n. Conn .. n. Pa .. Mich .. and N.D.
68. C. limosa L. Fig. 464
55. C. comosa Boott Fig. 456 Marly bogs. sphagnum bogs. and marshy pond margins. Lab.
Swamps. bogs. marshes. and shallow water. N.S. and Me. w. to and Nfld. w. to Alask.. s. to N.B.. DeL. Pa .. n. Ohio. n. TI!.. ne.
sw. Que .. Mich .. Minn .. S.D.. Ida .. Wash .. and Calif.. s. to Fla. Iowa. Sask.. Mont.. Utah. and Calif.
and e. Tex.
69. C. paupercula Michx. Fig. 465
56. C. schweinitzii Dew. ex Schwein. Fig. 457 Sphagnum bogs. fens. peaty hollows. coniferous swamps. wet
Swamps. meadows. cold springs and streams. and low. wet meadows. and alpine peats. Lab. and Nfld. w. to N.W.T. and
woods. Sw. vt. and w. Mass. w. to s. Ont. and n. Mich .. s. to w. Alask.. s. to Mass . w. Conn .. ne. Pa .. N.Y.. n. Ohio. n. Ind .. Wisc ..
Conn .. n. N.J.. Pa .. and mts. to N.C. and Tenn. Minn .. Sask .. Colo.. Utah. n. Ida .. and Wash.
57. C. lurida Wahlenb. Fig. 458 70. C. rariflora (Wahlenb.) Sm. Fig. 465
Mostly acid soils; swamps. marshes. wet woods. lakeshores. and Cold bogs. peat barrens. and pond margins. Chiefly arctic;
stream margins. N.S. and s. Que. w. to Minn .. s. to Fla .. Tex .. Green!' and Baffin I. w. to Alask .. s. to Nfld .. e. Que .. n. Me ..
and e. Mex. James Bay. and n. Man.
58. C. hystericina Willd. Fig. 457 71. C. barrattii Schwein. & Torr. Fig. 465
Calcareous soils; swamps. swales. wet meadows. and shores. W. Peaty swamps. pinelands. and wet woods. Coastal plain. Conn.
Nfld. and Que. w. to Alta. and Wash .. s. to N.B .. N.B.. Md .. Tenn .. s. to N.C.; mts. of Tenn. and Ala.
Ark.. Tex .. Ariz .. and Calif.
72. C. livida (Wahlenb.) Willd. Fig. 466
59. C. rastrata Stokes Calcareous meadows and bogs. Lab. and Nfld. w. to Alask.. s. to
Swamps. shallow water. and wet shores. Green!' and Lab. w. to n. Me .. ne. Mass .. w. Conn .. s. N.J.. c. N.Y.. Mich .. Minn .. Sask..
Alask .. s. to n. Nfld .. n. C.B.I .. n. N.B .. n. Mich .. n. Minn . Sask.. Ida .. and n. Calif. Plants in the southern portion of the range
nw. Mont.. and B.C. According to Reznicek (pers. comm.) much have been treated as C. livida var. radicaulis Paine. (C. grayana
of what has been called C. rastrata by American authors is refer- Dew.; C. livida var. grayana (Dew.) Fern.)
able to e. utriculata (= e. rastrata var. utricu/ata)
73. C. tetanica Schkuhr Fig. 469
60. C. utriculata Boott Fig. 459 Calcareous bogs. meadows. and low woods. E. Mass. w. to Ont.
Swamps. shallow water. and wet shores. Nfld. and Lab. w. to and Sask.. s. to Va .. Ohio. II!.. and Neb.
Alask .. s. to e. Va .. Pa .. n. W.Va .. Ind .. Iowa. Neb.. N.M .. Ariz ..
Calif.. and n. Baja Calif. (e. rostrata var. utriculata (Boott) Bailey; 74. C. viridula Michx.
e. rhynchophysa Fisch .. Mey. & Lall.) 74a. C. viridula ssp. viridula Fig. 467
Calcareous gravels. shores. and springy areas. Nfld. w. to Alask .
61. C. tuckermanii Dew. Fig. 459
s. to N.B.. n. N.J.. Pa .. n. Ind .. Ill .. Wise.. Minn .. S.D.. Mont.. N.M ..
Calcareous swamps. meadows. marshes. and low woods. Que.
Utah. Nev.. and Calif. (C. flava var. viridu/a (Michx.) Bailey; C.
w. to Ont. and Minn .. s. to N.B .. n. N.J.. Ohio. Ind .. and ne. Iowa.
oederi var. viridula (Michx.) Kuek.)
62. C. oligosperma Michx. Fig. 459 74b. e. viridula ssp. brachyrrhyncha (Celakovsky) B. Schmid
Acid swamps. sphagnum bogs. and occasionally in shallow wa- Fig. 469
ter. Lab. w. to N.W.T.. s. to Mass .. Pa .. Ind .. Minn .. and Sask. Calcareous bogs. fens. swamps. and gravels. Chiefly coastal.
staminate
spike
PIg. 45 4.
"",."no",I"
~
"' "' "
a. b" ,i t; b. tn
ll i> '= -; ""
Carex pseudocype
rus: e. infloresc
ence; f. perigy
"" ," ," "" d.
nium (F ).
,,,till"" ",,), (G&W),
152
g
Fig. 455. Carex michauxiana: a. habit; b. perigynium; c. achene; d. pistillate scale; e. ligule; f. section of leaf
(Mackenzie) .
CarexfolIiculata: g. inflorescence; h. perigynium; i. pistillate scale (G&W).
153
c
.
d. pistillate scale (G&W)
: a. inf lor esc enc e; b. perigynium; c. ach ene ;
Fig. 45 6. Carex comosa
15 4
staminate
spike
e
g
pistillate ---~IJ2re1:'/
spike
~---style
Fig. 457. Carex schweinitzii: a. habit; b. perigynium; c. achene; d. pistillate scale; e. ligule (Mackenzie).
Carex hystericina: f. inflorescence; g. perigynium (F).
155
Fig. 458. Carex lurida: a. habit: b. perigynium: c. achene: d. staminate scale: e. pistillate scale: f. ligule (G&W).
156
II I,
'~I I
I e
d f( /
~r
style
Fig. 459. Carex utriculata: a. inflorescence (F); b. perigynium (Mackenzie); c. achene (Mackenzie); d. pistillate
scale (Mackenzie); e. section of leaf blade (F).
Carex tuckermanii: f. inflorescence; g. tip of perigynium (F).
Carex oligosperma: h. inflorescence; i. perigynium (F).
157
!~"fr
1<
j
~.
i;:,
; .
, II
I e
Fig. 460. Carex bullata: a. habit; b. perigynium; c. achene; d. pistillate scale; e. ligule (Mackenzie).
158
....--anther
g c
stigma -----..
style - - - - - \
Fig.461. Carex vesicaria: a. inflorescence; b.lower portion of plant; c. perigynium; d. achene; e. achene, cross-
section; f. staminate flower and subtending scale; g. pistillate scale; h. ligule (Mason).
159
r-~i ':
",/
:1\ 'I1')
I
, '(
, "
,I
I'
"'!-:
~I~' ,
I
~:II
,e ,I
r "
I I,J e
Fig. 462. Carex lupuliJormis: a. habit; b. perigynium; c. achene; d. pistillate scale; e. ligule (Mackenzie).
160
g
Fig. 463. Carex buxbaumii: a. habit; b. terminal spike; c. perigynium; d. achene; e. achene, cross-section;
f. staminate scale with filaments; g. pistillate scale; h. ligule (Mason).
161
g
ciiiii::;:..
Fig. 464. Carex Iupulina: a. habit; b. perigynium; c. achene; d. pistillate scale (G&W).
Carex limosa: e. habit (HCOT); f. section of root (HCOT); g. perigynium (Mackenzie); h. achene (Mac-
kenzie); i. pistillate scale (Mackenzie).
162
k
Fig. 465. Carex rariflora: a. habit; b. perigynium; c. achene; d. pistillate scale (Mackenzie).
Carex barrattii: e. inflorescence; f. perigynium; g. achene; h. pistillate scale (Mackenzie).
Carex paupercula: i. inflorescence; j. perigynium; k. achene; l. pistillate scale (Mackenzie).
163
anther --",\.\
Fig. 466. Carex livida: a. habit; b, c. perigynium variations; d. achene; e, f. achenes, cross-sections, showing
variations; g. staminate flower with subtending scale; h, i. pistillate scales; j, k. ligules (Mason).
164
f
V d
Fig. 467. Carex viridula ssp. viridula: a. habit; b. perigynium; c. achene; d. achene. cross-section; e. pistillate
scale; f. ligule (Mason).
165
\
rt\
Fig. 468. Carexflava: a. habit; b. perigynium; c. achene; d. pistillate scale; e. section ofleafblade (Mackenzie).
166
, :.... : ..
..< .....
':';' .:~
'\'i)',
I a
b h
,e
'.
~~
;
:..
.<.1 <
.
~
Fig. 469. Carex tetanica: a. inflorescence; b. perigynium; c. achene; d. pistillate scale (Mackenzie).
Carex cryptolepis: e. inflorescence; f. perigynium (F).
Carex viridula ssp. brachyrrhyncha: g. inflorescence; h. perigyniuni; i. achene; j. pistillate scale (Mac-
kenzie).
167
168 ANGIOSPERMS
Nfld., Anticosti I., M.I., Gaspe Pen., coastal and c. Que., n. N.B., 76. C. flava L. Fig. 468
C.B.I., and Brier I., N.S. (c. Iepidocarpa Tausch) Wet shores, marshes, swales, and meadows; chiefly calcareous
sites. Nfld. w. to Man., s. to N.E., n. N.J., Pa., Ohio, Ind., Wisc ..
75. C. cryptolepis Mackenz. Fig. 469 and ne. Minn.; Alta., B.C., and s. Alask. s. to w. Mont. and Ida.
Meadows, sandy shores, ditches, and swales. Nfld. w. to w. Ont.,
s. to N.B., n. N.J., n. Ohio, n. Ind., Wisc., and Minn. (C.flava var.
fertilis auctts., non Peck)
2. Scleria (Nut-rush)
Perennial or annual herbs, grass-like; stems sharply 3-angled, leafy; flowers unisexual, pistillate spikelets I-flowered, intermixed
with clusters of few-flowered staminate spikelets; fruit a white stony achene without a tubercle.
1. Achene smooth, shiny white; disk (hypogynium) subtending achene 3-angled, white crusty-papillose (fig. 470b,c); leaves
(3)5-9 mm wide ..................................................................... 1. S. triglomerata
1. Achene roughened, reticulate or irregularly warty, with low transverse ridges (figs. 470e. 473b), dull white; disk (hypogynium)
absent. or if present, 3-lobed, not papillose; leaves 0.5-5 mm wide.
2. Achene reticulate. subtended by 3-lobed. bract-like disk (fig. 470f) .................................. 2. S. reticularis
2. Achene irregularly warty. with low transverse ridges. disk absent ................................... 3. S. verticillata
l. S. triglomerata Michx. Fig. 470 Mass. s. to Fla., w. to Tex. and Mex.; nw. Ind .. sw. Mich.. and c.
Wet to dry sites, borders of marshes, wet peats, thickets, savan- Wisc.; w.1., C.Am., and S.Am.
nas, open woods, and prairies. Mass. and Conn. w. to Ont..
Wisc., se. Minn., Iowa, and Neb., s. to Fla. and Tex. 3. S. verticilIata Muhl. ex Willd. Fig. 473
Wet sands, sandy peats and marls, swamps, meadows, and
2. S. reticularis Michx. Fig. 470 marshes. Mass. w. to s. Ont., Mich., Wisc., and Minn., s. chiefly
Wet open sands, peaty sands, pond shores, shallow water of cy- along the coast to Fla., w. to se. Tex.; Trop. Am.
press ponds, meadows, wet pinelands, and savannas. S.N.H. and
3. Rhynchospora (Beak-rush)
Perennial herbs (rarely annual), usually tufted, some producing elongate rhizomes; leaves mostly shorter than stems; stems 3-angled
or nearly round in cross-section; flowers in clustered or panicled spikelets, (1)2-4(6) or few to many, lower 2 or 3 spikelet scales
empty, upper flower often staminate; perianth consisting of bristles. sometimes absent; fruit a biconvex or flattened achene with a
persistent tubercle.
1. Involucral bracts spreading or drooping, basal portions whitish, green toward tips (fig. 471a,b) . . . . . . . . . . . . . . . . .. 1. R. colo rata
1. Involucral bracts ascending, green throughout.
2. Perianth bristles absent; spikelets few- to many-flowered.
3. Tubercles longer than wide. styles persisting (fig. 472f); achene finely roughened or wrinkled ............. 2. R. scirpoides
3. Tubercles much wider than long, styles deciduous (figs. 472d); achene strongly roughened or wrinkled ........ 3. R. nitens
2. Perianth bristles present (fig. 473e,g); spikelets (1)2-4(6)-f1owered.
4. Tubercles 10 mm or more long (fig. 473e,g).
5. Bristles longer than body of achene.
hypogynium
Fig. 47 0.
s,"", ~tkUl""'
Sc1eria triglomerata:
a.
d. b,ha bi
b\ " e.t b. achenef.; ""
"hen" c. achene. bottom
vie w (G&W).
,," c, "" "" '" v,"w
16 9
,h ow ln g 3Io""d b,,
<c tI;'" dl'" (G&W).
b
Fig.471. Rhynchaspam calamta: a. habit (C&C); b. inflorescence (G&W); c. spikelet (C&C); d. achene (G&W).
170
Fig. 472. Rhynchospora nitens: a. habit; b. sheath; c. spikelet; d. achene (G&W).
Rhynchospora scirpoides: e. habit (F); f. achene (G&W).
171
\
6. Spikelets in tight clusters of 10-50 (fig. 473c.f); tubercle 15-23 mm long; plants cespitose.
7. Achenes 2.6-3.1 mm wide; tubercle 1-1.8 mm wide at base; leaves firm, 3.5-12 mm wide
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4a. R. macrostachya var. macrostachya
7. Achenes 3-3.8 mm wide; tubercle 1.8-2.4 mm wide at base; leaves subflaccid, 9-15 mm wide ........ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4b. R. macrostachya var. colpophila
6. Spikelets in open clusters of 1-6 (fig. 473d); tubercle 10-18 mm long; plants stoloniferous ........... 5. R. inundata
5. Bristles shorter than body of achene.
8. Achenes 5-6 mm long, 2.8-3.3 mm wide; base of tubercle half the width of achene (fig. 473g) .......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6a. R. corniculata var. corniculata
8. Achenes 4.4-5.3 mm long, 2.4:-2.6 mm wide; base of tubercle nearly as wide as achene ............... .
6b. R. corniculata var. interior
4. Tubercles 5 mm or less long.
9. Achenes wrinkled (figs. 475c,g), transversely ridged, rugose, or honeycomb-reticulate.
10. Bristles long-fringed (fig. 475c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. R. oligantha
10. Bristles merely barbed (fig. 474c).
11. Stems lax and reclining; leaves 1 mm or less wide, involute ............................. 8. R. rariflora
11. Stems erect or strongly ascending; leaves 1-8 mm wide, flat.
12. Tubercle with a distinct raised rim at base (fig. 4740 ............................... 9. R. harveyi
12. Tubercle lacking a distinct raised rim at base.
13. Bristles shorter than body of achene.
14. Achene faces flat, lacking a bulge toward tip.
15. Scales awned; achene 1.3-2 mm long, with 10 or more transverse ridges; bristles 6
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10. R. torreyana
15. Scales blunt to mucronate; achene 1-1.3 mm long, with 8 or fewer transverse ridges;
bristles (0)1-3(6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. R. perplexa
14. Achene faces biconvex, with a bulge toward tip ......................... 12. R. globularis
13. Bristles longer than body of achene.
16. Leaves 1.5-4 mm wide; achene 2-2.5 mm long, faces flat .................. 13. R. inexpansa
16. Leaves 4-8 mm wide; achene 1-1.8 mm long, faces biconvex.
17. Achenes 1-1.3 mm long; tubercles smooth; branches ofinflorescence spreading, almost
at right angles to main rachis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14. R. miliacea
17. Achenes 1.4-1.8 mm long; tubercle margins setaceous (fig. 477h); branches of inflo-
rescence ascending . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. R. caduca
9. Achenes smooth.
18. Bristles retrorsely barbed.
19. Bristles 8-12, villous at base (fig. 477b); scales of spikelet white to pale brown .................. 16. R. alba
19. Bristles 6, generally glabrous at base (fig. 477d); scales of spikelet dark brown to dark reddish-brown.
20. Clusters of spikelets subglobose, the lowest spikelet in each cluster spreading or reflexed (fig. 477c,e).
2l. Achenes 2-2.4 mm long, 1.4-1.6 mm wide .............................. 17. R. cephalantha
2l. Achenes 1.4-1.6 mm long, 0.9-l.1 mm wide ........................... 18. R. microcephala
20. Clusters of spikelets ellipsoidal, all spikelets ascending (figs. 478c).
22. Achenes with pale margins, obovoid; leaves flat.
23. Spikelets I-fruited ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19. R. chalarocephala
23. Spikelets 2-fruited, rarely I-fruited, with a sterile floret present.
24. Leaves 3-7 mm wide; spikelets 5-6 mm long; achenes 1.7-2 mm long with conspicuous
central bulge, summit nearly truncate (fig. 478b) .................... 20. R. glomerata
24. Leaves 0.5-4 mm wide; spikelets 3-5 mm long; achenes l.3-1.8 mm long, without
central bulge, summit rounded (fig. 478d) ......................... 21. R. capitellata
22. Achenes lacking pale margins, ellipsoidal to narrowly obovoid; leaves involute-filiform ...... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 22. R. capillacea
18. Bristles antrorsely barbed or smooth, occasionally absent.
25. Base of plants swollen and bulbous; spikelets I-flowered; bristles absent or 1-3, rudimentary . . . . .. 23. R. pallida
25. Base of plants not swollen and bulbous; spikelets 2-4( 6 )-flowered; bristles 5 or 6.
26. Achenes 0.6-l.1 mm wide, distinctly prolonged to a stipe-like base (fig. 480b,d).
174 ANGIOSPERMS
27. Plants stoloniferous; achenes 1.3-1.7 mm long; tubercle usually green; spikelets 2- or 3-flowered
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24. R. fusca
27. Plants cespitose; achenes 1-1.3 mm long; tubercle tan; spikelets 3- or 4(6)-flowered ..... 25. R.filifolia
26. Achenes 1.1-1.7 mm wide. not stipitate.
28. Tubercles elongate. 1-2.6 mm long. as long as or longer than achene body; bristles as long as or
longer than tip of tubercle ........................................... 26. R. gracilenta
28. Tubercles deltoid. 0.4-0.7 mm long. shorter than achene body; bristles typically shorter than or
longer than achene body. but shorter than tubercle.
29. Stems and leaves tending to be flexuous and reclining; scales bifid. shorter than achenes;
spikelets 2.5-3.2 mm long ......................................... 27. R. debilis
29. Stems and leaves erect. stilT; scales not 2-c1eft. longer than achenes; spikelets 3-5 mm long.
30. Scales awned; achenes 1.5-2 mm long; bristles absent or much shorter than achene
body ....................................... 28a. R. fascicularis var. fascicularis
30. Scales mucronate; achenes 1.3-1.5 mm long; bristles longer than achene body .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28b. R. fascicularis var. distans
1. R. colorata (L.) H. PfeilTer White-top Sedge Fig. 471 8. R. rariflora (Michx.) Ell. Fig. 474
Wet peats. peaty sands. savannas. borders of wet depressions. Damps sands and peats of shores. Coastal plain. s. N.J. s. to Fla ..
and ditches. Coastal plain. e. Va. s. to Fla .. w. to e. Tex. and e. w. to Tex.; inland n. to Tenn.; WI. and C.Am.
Mex.; WI. and C.Am. The more robust R. latifolia (Baldwin)
Thomas. although noted in some manuals from Virginia. is re- 9. R. harveyi Boott Fig. 474
ported by Thomas (1984) only as far north as North Carolina. Moist to well-drained peats. pinelands. and depressions. Coastal
(Dichromena colo rata (1.) Hitchc.) plain. Va. s. to Fla .. w. to Tex.; inland n. to Mo. and Okla.
2. R. scirpoides (Torr.) Gray Fig. 472 10. R. torreyana Gray Fig. 475
Wet. sandy. peaty. and mucky shores. Chiefly coastal plain. se. Damp to dryish sands and peats of pond margins and pinelands.
Mass. s. to Fla. panhandle; disjunct to Mich .. n. Ind . and c. Coastal plain. se. Mass. s. to Ga.
Wisc. (Psilocarya scirpoides Torr.; P. scirpoides var. grimesii
Fern. & Grisc.) 11. R. perplexa Britt. ex Small Fig. 476
Wet peat. shores. and shallow water. Coastal plain. se. Va. s.
3. R. nitens (Vahl) Gray Fig. 472 to Fla .. w. to Tex.; inland n. to c. Tenn.; WI. (R. perplexa var.
Wet sands and peats. Coastal plain. Mass. s. to Fla .. w. to Tex.; virginiana Fern.)
disjunct to nw. Ind. (Psilocarya nitens (Vahl) Wood)
12. R. globularis (Chapm.) Small Fig. 476
4. R. macrostachya Torr. ex Gray Wet peaty and sandy soils of pine lands. depressions. and shores.
N.J. s. to Fla .. w. to interior N.C . s. Ky. Tenn .. Mo .. Okla .. and
4a. R. macrostachya var. macrostachya Fig. 473
Tex.; n. Ohio and s. Mich. w. to ne. Ill.; n. Calif.; WI. and C.Am.
Wet sand and peat. along the shores of ponds. lakes. streams.
(R. globularis var. recognita Gale; R. cymosa ElL)
and marshes. Coastal plain. s. Me. s. to Fla . w. to Tex.; inland
in c. N.Y.. c. Ky.. sw. Mich .. s. Ill.. Mo .. and Kans.
13. R. inexpansa (Michx.) Vahl Fig. 475
4b. R. macrostachya var. colpophila Fern. & Gale Wet to dry sandy and peaty soils of swamps. ditches. and pond
Freshwater tidal marshes. Md. s. to se. Va. margins. Coastal plain. se. Va. s. to n. Fla .. w. to Ark. and e.
Tex.; WI.
5. R. inundata (Oakes) Fern. Fig. 473
Peaty and mucky shores of ponds. swamps. and ditches. Coastal 14. R. miliacea (Lam.) Gray Fig. 478
plain. e. Mass. s. to Fla.. w. to Ala. (La.?); rare and local in the Stream margins. pine woods. thickets. swamps. and ditches.
northern portion of its range. Coastal plain; se. Va. s. to Fla . w. to La.
tubercul e
175
bristle
176
a
e
g
Fig. 476. Rhynchospora globularis: a. inflorescence (F); b. inflorescence (G&W); c. scale (G&W); d. achene
(G&W).
Rhynchosporaperplexa: e. inflorescence; f. scale; g. achene (G&W).
177
Fig. 477. Rhynchospora alba: a. habit (HeOT); b. achene (G&W).
Rhynchospora cephalantha: c. inflorescence (F); d. achene (G&W).
Rhynchospora microcephala: e. inflorescence; f. achene (G&W).
Rhynchospora caduca: g. inflorescence; h. achene (G&W).
178
bristle
18. R. microcephala Britt. Fig. 477 to ant.. s. Mich .. n. Wise.. ne. Minn . and Sask.. s. to L.r.. N.Y.
Sandy and peaty shores of swamps. savannas. and wet pine- and Del.
lands. Coastal plain. N.J. s. to Fla.. w. to Miss.; Cuba. (R. cepha-
Iantha var. microcephala (Britt.) Kuek.) 25. R.filifolia Gray Fig. 480
Pond margins. swamps. and pine savannas. Coastal plain. N.J.
19. R. chalarocephala Fern. & Gale Fig. 479 s. to Fla .. w. to se. Tex.; inland n. to Ark. and Tenn.
Pond margins. swamps. savannas. and wet pinelands. Coastal
plain. N.J. s. to Fla . w. to La. 26. R. gracilenta Gray Fig. 480
Wet pine barrens. peats. swales. and sandy depressions. Chiefly
20. R. glomerata (L.) Vahl Fig. 478 coastal plain. N.J. s. to n. Fla. w. to e. Tex.; inland n. to Tenn.
Wet peaty or sandy soil of bogs. savannas. and meadows. and Ark.
Coastal plain. N.J. s. to Fla . w. to e. Tex.; inland n. to Ky.. Tenn.
Ark.. Kans .. and Okla. 27. R. debilis Gale Fig. 480
Damp sandy and peaty soil of bogs and pine savannas. Coastal
21. R. capHellata (Michx.) Vahl Fig. 478 plain. se. Va. s. to n. Fla . w. to Ala. and La.
Bogs. sandy shores. low ground. and wet pine savannas. N.B. w.
to sw. Que.. s. ant.. and Wisc.. s. to n. Fla. and e. Tex. 28. R. fascicularis (Michx.) Vahl
28a. R. fascicularis var. fascicularis Fig. 480
22. R. capillacea Torr. Fig. 479
Moist to wet pinelands. savannas. pond margins. and ditches.
Calcareous bogs. sands. and ledges. W. Nfld. w. to Sask.. s. to
Coastal plain. se. Va. s. to Fla .. w. to e. Tex.
Me.. Conn . n. N.J. se. Pa. sw. Va . e. Tenn . Ind .. Mo . and S.D.
28b. R. fascicularis var. distans (Michx.) Chapm.
23. R. pallida M. A. Curtis Fig. 479 Moist to wet pinelands. wet peats. and ditches. Coastal plain. se.
Wet peats and wet pine barrens. Coastal plain. L.r.. N.Y. s. to Va. s. to Fla .. w. to Miss.; Ber. This variety may not be worthy of
N.C. taxonomic recognition.
4. Cladium (Twig-rush)
Perennial herbs. arising from stout rhizomes or stolons; stems leafy; inflorescence much-branched. each branch bearing a cluster of
spikelets; spikelets 1- or 2-flowered. perianth absent; fruit an ovoid to globose achene without a tubercle.
1. Leaves narrow. 1-3 mm wide. channeled. margins and midrib nearly smooth ............................ 1. C. mariscoides
1. Leaves broad. 5-10 mm wide. flat. Stiff. margins and midrib saw-toothed ................................ 2. C. jamaicense
1. C. mariscoides (Muhl.) Torr. Twig-rush Fig. 481 2. C. jamaicense Crantz Saw-grass Fig. 481
Swamps. sandy. boggy. and marshy shores. and swales. often in Freshwater swamps. marshes. and shores. and saline marshes.
shallow water. fresh or brackish. Nfld. and N.S. w. to s. ant.. Coastal. Va. s. to Fla . w. to Tex. and Mex.; w.r. (Mariscus ja-
Mich . and Minn . s. to Va . Ohio. and Iowa; mts. to N.C . S.C. maicensis (Crantz) Britt.)
Ga .. Ala .. and Miss. (Mariscus mariscoides (Muhl.) Ktze.)
5. Kyllinga (Sedge)
Perennial herbs. usually arising from rhizomes. or tufted annuals; stems 3-angled; leaf blades with scaberulous margins and keel.
or sometimes reduced to sheaths; involucral leaves 2-4. subtending dense inflorescence; spikelets 2-scaled. I-flowered; fruit an
achene. without a tubercle.
1. K. pumila Michx. Fig. 482 2. K. brevifolioides (Thieret & Delahoussaye) Tucker Fig. 482
Swamps. marshes. ponds. wet clearings. and ditches. L.r.. N.Y. Wet sites. marshy shores. and swamp margins. Conn. s. to N.C.
and e. Pa. w. to e. Kans .. s. to Fla .. Tex. and Mex.; w.r. and and Tenn. This taxon is similar to K. brevifalia Rottb.. which is
S.Am. (Cyperus tenuifolius (Steud.) Dandy; Kyllinga tenuifolia more widespread in the southeastern United States and tropical
Steud.) areas. Both species form large mats from numerous rhizomes.
(Cyperus brevifaliaides Thieret & Delahoussaye)
f
Fig. 481. Cladium jamaicense: a. inflorescence; b. base of plant; c. section of stem with leaf sheath; d. section of
leaf; e. achene (G&W).
Cladium mariscoides: f. inflorescence; g. achene (F).
183
Fig. 482. Kyllinga pumila: a. habit; b. scales; c. achene (G&W).
Kyllinga brevifolioides: d. scales; e. achene (Delahoussaye and Thieret, 1967).
184
Cyperaceae / Sedge Family 185
6. Eriophorum (Cotton-grass)
Perennial herbs. grass-like. cespitose. or stems arising from creeping rhizomes; flowers bisexual. sub tended by membranous scales;
perianth of numerous bristles. greatly elongating with maturity; achenes lacking tubercle. subtended by persistent cottony bristles.
186
nt h
r- -_ :: :: - pe ria
bristles
c. ac he ne (TF).
m pylaiea num: a. habit; b. scale;
Fig. 48 4. Eriophoru ; f. ac he ne (TF).
iophorum va ginatum; d. habit; e. scale
Er
18 7
e
e d
peduncl e
~."'.(".'
.'
,
,.
..-.'.\
~
~ ' ~/ I.
t-'.."-... .- ..:.)
..'.' " " " '..'.'.'
\ -.'
l~,c i
. f
b
a
190
Cyperaceae / Sedge Family 191
7. E. gracile W. D. J. Koch Fig. 486 10. E. viridicarinatum (Engelm.) Fern. Fig. 487
Bogs, wet peat, and inundated shores. Lab. and Nfld. w. to Bogs, wet peats, peaty meadows, and conifer swamps. Lab. and
Alask., 's. to N.E., Del., Pa., Mich., n. Ill., Iowa, Colo., Ida., and Nfld. w. to Alask., s. to N.B., N.Y., Ohio, n. Ind., n. Ill., n. Minn.,
c. Calif. N.D., nw. Mont., Colo., Ida., and s. B.C.
7. Eleocharis (Spike-rush)
Perennial or annual herbs, tufted or arising from rhizomes or stolons; leaves reduced to bladeless sheaths; spikelet solitary, with few
to many bisexual flowers; perianth bristles usually 6, occasionally absent; achenes biconvex or 3-angled, with a persistent tubercle.
The shape and size of the tubercle are of great taxonomic importance in the genus, and identification is almost impossible without
mature achenes.
1. E. equisetoides (El!.) Torr. Jointed Spike-rush Fig. 488 Coastal plain. se. Mass. s. to N.J.. Va .. and Fla.; disjunct in w.
Shallow water of marshes. lakes. ponds. and ditches. Coastal Mich.
plain. e. Mass. s. to n. Fla . w. to e. Tex.; inland n. to Mo. and
the Great Lakes region. 11. E. compressa Sulliv. Fig. 492
Shallow water. calcareous gravels. sands. and peats. W. Que. w.
2. E. quadrangulata (Michx.) R. & S. Square-stem Spike-rush to Sask.. s. to ne. Va . Ga . n. Ala . Mo . ne.Tex. and Colo. (E.
Fig. 489 elliptica var. compressa (Sulliv.) Drapalik & Mohlenbrock)
Shallow water of marshes. lakes. and ponds; often tidal along
coast. E. Mass. w. to Ont.. Mich.. and Wisc.. s. to Fla.. Okla . 12. E. nitida Fern. Fig. 493
Tex . and Mex. (E. quadrangulata var. crassior Fern.) Damp peaty and sandy sites. Ntld. w. to Wisc. and ne. Minn .
s. to N.S . Me . and n. N.H.; B.C. and the Aleutian I.. Alask.
3. E. robbinsii Oakes Triangle Spike-rush Fig. 489
Shallow water. often submersed. forming large mats in ponds. 13. E. elliptica Kunth Fig. 493
lakes. and marshes. N.S. w. to Ont. and n. Wisc.. s. to c. N.Y. and Damp marly ground. shores. swales. and ditches. Ntld. w. to
n. Ind.; s. along the coastal plain to n. Fla. and sw. Ala. In the Minn. s. to N.B.. n. N.J. Ohio. Iowa; infrequent w. to Man .. s.
vegetative state this species is easily confused with two other Alta .. nw. Mont.. and sw. B.C. (E. compressa var. atrata Svens.)
aquatics that produce fine hair-like submersed leaves. Scirpus
subterminalis and Juncus militaris. 14. E. tenuis (Willd.) Schultes
14a. E. tenuis var. tenuis Slender Spike-rush Fig. 493
4. E. smallii Britt. Fig. 490
Wet. damp. and dry sands. gravels and peats. N.S. sw. to Ohio.
Shallow water and along shores of streams. rivers. lakes. and
s. to S.C.
ponds. Nfld. w. to Ont. and Wisc. s. to Md .. w. Va. s. Ind . and
Mo. This taxon is a member of the extremely complex group 14b. E. tenuis var. pseudoptera (Weath.) Svens. Fig. 493
subseries Palustres. Svenson (1957) includes New England Meadows. L.I .. N.Y.. and N.J. s. to Va. and mts. of N.C. and e.
plants fitting Fernald's concept of E. palustris (L.) R. & S. within Tenn. (E. capitata var. pseudoptera Weath.; E. elliptica var. pseu-
E. smallii. which he considers distinct from the European E. pal- doptera (Weath.) Harms)
ustris. Some American authors take a broader taxonomic view
of this complex and treat E. smallii. E. macrostachya. E. uniglumis. 15. E. verrucosa (Svens.) Harms Fig. 493
and even E. erythropoda as synonyms under E. palustris. (E. Moist to wet sandy and peaty soils. Pa. se. Va . and Ky. w. to s.
smallii var. major (Sonder) Seymour) Ind .. s. IlL. Iowa. and Neb. s. to Ark.. e. Okla .. and e. Tex. (E.
capitata var. verrucosa Svens.; E. tenuis var. verrucosa (Svens.)
5. E. macrostachya Britt. Fig. 490 Svens.)
Marshes. ditches. and shores. Ill. and Minn. w. to B.C. s. to La .
Mex . and Calif. E. xyridi!ormis Fern. & Brack .. a southwestern 16. E. radicans (Poir.) Kunth Fig. 494
taxon sometimes mistaken for E. macrostachya. does not appear Wet. sandy and mucky shores. Local; se. Va., Fla .. and Mich.;
to be in our range. (E. mammillata sensu auctt. non Lindb. f.) Okla. and e. Tex. w. to Ariz . Calif. and Mex.; WI. and S.Am.
6. E. erythropoda Steud. Fig. 491 17. E. acicularis (L.) R. & S. Needle-rush Fig. 494
Shallow water. wet shores. and marshes. C.B.I. and Que. w. to Wet sands and mucky shores. often submersed. Green!' w. to
Hudson Bay and Man . s. to N.B.. Va .. Tenn .. Neb .. and Okla. (E. Alask .. s. to Fla.. Okla . n. Mex . and s. Calif. This species fre-
calva Torr.; E. palustris var. calva (Torr.) Gray) quently remains vegetative. forming large mats. (E. acicularis
var. submersa (Hj. Nilss.) Svens.)
7. E. uniglumis (Link) Schultes
Wet shores and marshes. Nfld. and Lab. w. to Alask.. s. to N.S.. 18. E. pauciflora (Lightf.) Link Fig. 495
e. Que. N.D.. Wyo .. and Oreg. Damp. marly shores. ledges. and swamps. Green!' and Ntld. w.
to n. Ont.. Sask.. and B.C .. s. to Mass . nw. N~J. N.Y.. nw. Pa ..
8. E. halophila (Fern. & Brack.) Fern. & Brack. Fig. 492 Ind . Iowa. Colo. N.M.. and Calif. Our plants are separated as
Salt marshes and saline or brackish shores. Ntld. and e. Que. var. !ernaldii Svens. from the Eurasian var. pauciflora. (E. quin-
w. to James Bay. s. to N.Y. and Va. (E. uniglumis var. halophila queflora (Hartm.) sw.)
Fern. & Brack.)
19. E. parvula (R. & S.) Link Fig. 495
9. E.!allax Weath. Fig. 491 Wet saline or brackish shores. Ntld. s. along the coast to Fla . w.
Freshwater pond margins and brackish marshes. Coastal plain. to Tex. and Mex.; locally inland w. to w. N.Y.. Mich .. w. Ont..
e. Mass. s. to Fla .. w. to Tex.; Cuba. (E. ambigens Fern.) Minn . Mo . and Sask .. s. along the Pacific coast from B.C. to
Calif.; W.I. and S.Am.
10. E. tricostata Torr. Fig. 492
Sandy and peaty shores of freshwater to brackish marshes.
c
tubercule
Fig. 488. Eleocharis equisetoides: a. habit; b. spikelet; c. achene; d. leaf sheath (G&W).
194
spikelet
Fig. 489. Eleocharis quadrangulata: a. habit (Mason); b. spikelet (F); c. achene (Mason)
; d. stem. cross-section
(Mason); e. tubers (F).
Eleocharis robbil1sii: f. habit (F); g. achene (Beal).
195
perianth
bristle
sterile ....c:"---_-\".../.-4~
scales
196
p e r ia n th _
-- -#
bristle
solitary
sterile scale
r\)"."
\it f
Fig. 492. Eleocharis tricostata: a. habit (G&W); b. spikelet (G&W); c. achene (G&W); d. leaf sheath (F).
Eleocharis compressa: e. habit (Svenson, 19 32b); f. spikelet (F).
Eleocharis halophila: g. achene; h. leaf sheath (Beal).
198
spikelet
~ (
~
\
\i
scale--
I
rr I I
e r
.1
.\
\
;
I . II)
I i I'
\\ I f
Fig. 493. Eleocharis tenuis var. tenuis: a. habit (HeOT); b. spikelet (F); c. achene (F).
Eleocharis tenuis var. pseudoptera: d. spikelet (F).
Eleocharis verrucosa: e. achene (F).
Eleocharis nitida: f. habit (F).
Eleocharis elliptica: g. habit; h. spikelet; i. achene (F).
199
a
Fig. 494. Eleocharis acicularis: a. habit (G&W); b. habit. submersed plant (F); c. spikelet (G&W); d. achene
(G&W); e. leaf sheath (G&W).
Eleocharis radicans: f. achene (G&W).
200
~~~~tubers
b
d
20. E. intermedia Schultes Fig. 496 27. E. wolfii (Gray) Gray Fig. 498
Wet calcareous soils, stream margins, marshy ground, and mud Wet shores and prairies. L.r., N.Y.; s. Ind. nw. to Minn. and s.
flats. E. Que. and N.B. w. to Minn., s. to N.B., n. N.J., Md., w. Va., Sask., s.locally to Tenn., Ala., La., Mo., Kans., and Colo.
Tenn., Ill., and Iowa.
28. E. microcarpa Torr. Fig. 499
21. E.flavescens (Poir.) Urban Fig. 496 Damp sands, swamps, and shores, often in shallow water.
Wet peats and sands. Coastal plain, N.J. s. to Fla., w. to e. Tex. Coastal plain; e. Conn. s. to Fla., w. to La.; Tenn. and nw. Ind.
and Mex.; w.I. and S.Am. (E. microcarpa var.filiculmis Torr.; E. torreyana (Boeckl.) Boeckl.;
E. brittonii Svens. ex Small; E. lundellii Svens.)
22. E. olivacea Torr. Fig. 497
Wet sands, peats, and mud flats, occasionally in flowing water. 29. E. albida Torr. Fig. 499
Coastal. N.S. s. to Fla.; local inland, Me.; w. to s. Ont., Mich., Brackish marshes and saline shores. Coastal; Md. s. to Fla.,
and Minn., s. to w. Pa. and Ohio. w. to Tex. and e. Mex.; Ber.
23. E. caribaea (Rottb.) Blake Fig. 497 30. E. tuberculosa (Michx.) R. & S. Fig. 500
Wet, fresh and brackish sands and gravels. Great Lakes region Wet, sandy, and peaty shores, swamps, and ditches. Chiefly
of s. Ont., Mich., and nw. Ind.; S.C. s. to Fla., w. to Tex.; s. Calif.; coastal plain, N.S. s. to Fla., w. to Ark. and se. Tex.; Tenn.
Trop. Am. (E. caribaea var. dispar Blake; E. geniculata (L.) R. & S.)
31. E. melanocarpa Torr. Fig. 500
24. E. ovata (Roth.) R. & S. Fig. 498 Sandy, peaty, and mucky shores. Coastal plain, se. Mass. s. to se.
Muddy and wet open areas. Local; Nfld. w. to Minn., s. to ne. Va. and n. Fla., w. to Tex.; disjunct in s. Mich. and n. Ind.
Mass., Conn., c. N.Y., and n. Ind.; Wash. and Oreg. (E. obtusa
var. ovata (Roth.) Drapalik & Mohlenbrock) 32. E. rostellata Torr. Walking Sedge Fig. 501
Saline, brackish, and calcareous marshes, bogs, and swamps.
25. E. obtusa (Willd.) Schultes Fig. 498 Coastal. N.S. and Me. s. to Fla.; inland, N.Y. and Ont. w. to Wisc.
Muddy. sandy, and peaty shores, often in shallow water. P.E.I. and Minn., s. to IlL, Kans., Okla., Tex., and Mex.; Wyo., Utah,
and N.S. w. to Minn., s. to nw. Fla., Miss., e. Tex., Colo., and N.M., and Calif.; B.C.; Ber., w.I., and S.Am.
N.M.; Ida. and B.C. s. to n. Calif. (E. obtusa var. ellipsoidalis Fern.;
E. obtusa var. jejuna Fern.; E. obtusa var. peasei Svens.; E. dian- 33. E. tortilis (Link) Schultes Fig. 502
dra Wright) Springy swamps, wet peat, wet woods, and thickets. Coastal
plain, L.r., N.Y., s. to Fla., w. to e. Tex.
26. E. engelmannii Steud. Fig. 498
Wet sand, peat, and mud. S. Me. w. to Sask., Alta., and Wash., 34. E. vivipara Link Fig. 503
s. to e. Va., Ga., Ark., Tex., Ariz., and s. Calif. (E. engelmannii var. Peaty shores, marshes, pools, and ditches. Coastal plain, se. Va.
robusta Fern.; E. obtusa var. engelmannii (Steud.) Gilly) s. to Fla. (E. curtissii Small).
Perennial herbs, arising from rhizomes; stems round in cross-section, hollow, leafy; lower leaves reduced to sheaths; upper leaves
with flat blades, conspicuously 3-ranked; inflorescences axillary; spikelets racemose, several-flowered; scales strictly 2-ranked; flowers
bisexual; bristles 6-9; achenes biconvex, slender style persistent, tubercle absent.
Perennial herbs with rhizomes, stolons, or enlarged, hardened bases, or tufted annuals; stems usually 3-angled, leafy at base; involu-
cralleaves 1 or more, subtending terminal, umbellate inflorescence; spikelets 3-many per spike or head; scales 2-ranked; achene
biconvex or 3-angled, without a tubercle.
It is essential to collect the entire plant for accurate identification.
Fig. 498. Eleocharis obi usa: a. habit (F); b. spikelet (G&W); c. achene (F); d. leaf sheath (G&W).
Hleocharis ovata: e. achene (F).
Eleocharis engelmannii: f. achene (F).
Hleocharis wolfii: g. achene (F).
205
d
Fig. 499. Eleocharis microcarpa: a. habit; b. spikelet; c. scale; d. leaf sheath; e. achene
(G&W).
Eleocharis albida: f. habit; g. spikelet; h. achene; i. leaf sheath (G&W).
206
tubercle
",we ""'"'0
"'"",""w"'"" "",d <
o"""""';,.,.h,p,b"'" b. ,p " d o t ; c. " h
,ca' 'h<~h (G&v
f ig . 5 00
.
d i t ; ,." h , n,; , n '; d
.
g.
2 0 7 '' '" ,h c ath (G& n
).
Fig. 501. Eleocharis rostellata: a. habit; b. spikelet; c. achene; d. summit ofleaf sheath, two views (G&W).
208
d
Fig. 502. EIeocharis tortilis: a. habit; b. spikelet; c. achene; d. leaf sheath (G&W).
209
Fig. 503. Eleocharis vivipara: a. habit; b. proliferated inflorescence; c. achene; d. summit of leaf sheath (G&W).
210
b
Fig. 504. Dulichium arundinaceum: a. habit (F); b. inflorescence (C&C); c. achene (C&C).
211
212 ANGIOSPERMS
1. C. j1avicomus Michx. tened mound. (C. polystachyos var. texensis (Torr.) Fern.; C. mi-
Pond shores. swales. and ditches. L.I.. N.Y.. Md . and e. Va. w. to crodontus Torr.; C. paniculatus Rottb.; C. careyi Britt. ex. Small; C.
e. Ky.. Ark . N.M . and Ariz . s. to Ga . Tex . and Mex.; Trop. Am. odoratus sensu Small. not L.)
(C. albomarginatus (Nees) Steud.; Pycreus albomarginatus Nees)
5. C. diandrus Torr. Low Flatsedge Fig. 506
2. C. j1avescens L. Fig. 505 Sandy to muddy shores. occasionally wet peat. N.B. and Que. w.
Wet sands and peats along pond and lake shores. and ditches. to Wisc. and N.D.. s. to S.C . n. Ala . and Mo.
Mass. w. to Ont.. Mich .. Mo . and Kans .. s. to Fla. and Tex.;
Trop. Am. 6. C. bipartitus Torr. Fig. 506
Wet. sandy. gravelly. or muddy soils of marshes. shores of lakes
3. C. filicinus Vahl and ponds. and ditches. N.B. and c. Me. w. to s. Ont.. Minn .
Saline or brackish marshes. rarely freshwater shores. Coastal Neb. and Oreg . s. to Ga . Miss . Tex . and Calif.; S.Am. (c. rivu-
plain. Me. s. to Fla.. w. to La. (C. polystachyos var. filicinus laris Kunth)
(Vahl) O'Neill)
7. C. squarrosus L. Fig. 507
4. C. polystachyos Rottb. Fig. 505 Damp sands. silts. riverbanks. shores. and wet depressions in
Damp sands. peats. marshy shores. and seasonally wet areas. rock outcrops. N.B. w. to s. Ont.. Minn .. s. Man .. s. Sask. and s.
Coastal plain and piedmont. Me. and Mass. s. to Va . W.Va .. S.C . B.C .. s. to Fla .. Tex.. Calif. and Mex.; W.I. and S.Am. (C. aristatus
and Fla .. w. to Tex. and Mex.; inland n. to Mo. and Okla.; Trop. Rottb.; C. inj1exus Muhl.)
Am. In the North this species typically grows in a short. flat-
c
213
e
8. C. acuminatus Torr. & Hook. Fig. 508 Man., and Alask., s. to Fla., Tex., Calif., and Mex.; C.Am. and
Wet limestone soils. N.H.; Va. w. to Ohio, s. Mich., Minn., N.D., S.Am.
and B.C., s. to Ga., Tenn., La., Okla., Tex., Ariz., Nev., Calif., and
ne. Mex. 14. C. dentatus Torr. Fig. 510
Sandy and gravelly shores of ponds and lakes. Chiefly coastal
9. C. haspan L. Fig. 508 plain, N.S. and N.B. w. to s. Que. and se. Ont., s. to N.E., Del.,
Tidal, fresh to saline waters, marshy shores, stream borders, Md., N.C., and S.C.; nw. Ind.
and temporary pools. Coastal plain, se. Va. s. to Fla., w. to Tex.;
inland n. to Tenn.; Trop. Am. 15. C. pseudovegetus Steud. Fig. 511
Wet marshy shores, ditches, and swales. Se. Mass. (adv.); s. N.J.
10. C. ida L. Fig. 509 w. to Ind., Ill., Mo., and se. Kans., s. to Fla. and Tex. The name
Clearings, ditches, moist fields, often in shallow water. Coastal C. virens Michx. occasionally has been misapplied by some au-
plain and piedmont, Va. and W.Va. s. to Fla., w. to Tex. and thors to C. pseudovegetus. The spikelets appear similar but the
Mex.; inland n. to Ky., Tenn., and Okla.; W.r. and C.Am.; intro. more southern C. virens is a large, more robust plant with scab-
from Eurasia. rous stems.
11. C. strigosus L. Fig. 509 16. C. erythrorhizos Muhl. Red-rooted Cyperus Fig. 511
Marshes, meadows, damp clearings, and shores. N.E. w. to sw. Alluvial or damp soil along marshes, swales, and ditches. Se.
Que., Mich., Minn., and S.D., s. to Fla., Tex., and Ariz.; Wash. s. Me., se. N.H., and Mass. w. to s. Ont., Wisc., N.D., and B.C., s. to
to Calif. (C. strigosus var. robustior Britt.) Fla., Tex., N.M., Wash., and Calif.
l. F. pumila (Torr.) Spreng. Fig. 512 mont, L.r., N.Y. and s. N.J. s. to n. Fla., w. to Tenn., s. Ark., se.
Moist to wet sandy or sandy-peaty shores. Chiefly coastal plain. Okla . and e. Tex. (F. hispida Ell., F. squarrosa var. hispida (Ell.)
se. Mass. s. to Fla., w. to se. Tex.; disjunct in s. Ont.. s. Mich . Chapm.)
n. Ind., and c. Wisc. (F. squarrosa var. pumila Torr.)
4. F. simplex Vahl Fig. 514
2. F. breviseta (Cov.) Cov. Fig. 513 Swamps, swales, wet peats, seeps, and ditches. W. Ill., Mo., and
Wet sands. peats, and ditches. Coastal plain, se. Va. s. to Fla .. e. Neb. sw. to Tex., N.M., and Mex.; W.I. and C.Am.
w. to se. Tex. (F. squarrosa var. breviseta Cov.)
221
Fig. 513. Fuirena breviseta: a. habit; b. summit ofleafsheath; c. scale; d. perianth and achene (Kral. 1978).
Fuirena squarrosa: e. habit; f. summit of leaf sheath; g. scale; h. perianth and achene; i. perianth scale
variations (Kral. 1978).
222
c
involucral
bract
I
b
scale
Fig. 514. Fuirena simplex: a. upper portion of plant (F); b. summit of leaf sheath
(Kral. 1978); c. scale (Kral,
1978); d. perianth of scales and bristles (Kral. 1978).
Scirpus cespitosus: e. habit; f. inflorescence; g. achene (HeOT).
223
224 ANGIOSPERMS
Perennial or annual herbs. arising from rhizomes or tufted; stems 3-angled to nearly round in cross-section; leaves with blades or
reduced to sheaths; inflorescences variable. terminal or lateral; involucral bracts numerous. single. appearing as continuation of
stem. or absent; perianth of (0)1-6(8) bristles; stamens 2 or 3. style 2- or 3-cleft; achenes often apiculate. but lacking a tubercle.
The treatment of the leafy species follows that of Schuyler (1967). Certain species differ only in small technical characters (e.g .
Scirpus atrovirens vs. S. hattorianus). and hybridization within the genus contributes to differences of opinion as to the distinction of
taxa in some species complexes. In these cases. careful comparison of specimens is required.
In reevaluating the genus Scirpus for the Flora of North America Smith (1995. in press) has recognized the segregate genera
Schoenoplectus. Bolboschoenus. Isolepis. and Trichophorum. While we have retained the traditional concept of Scirpus here. we have
also provided synonymy for our taxa in those segregate genera.
REFERENCES: Beetle.
1947; Browning. Gordon-Gray. and Smith. 1995; Koyama. 1962. 1963; Miller and Beal, 1972; Raymond. 1958;
Schuyler. 1962a. 1962b. 1964a. 1964b. 1966. 1967. 1968. 1974a. 1974b; Smith. 1969; Tucker. 1992; Tucker and Miller. 1990.
1. Spikelets subtended by a single. very short. obscure involucral bract nearly as long as or shorter than spikelets (fig. 514e,f)
(occasionally longer than spikelet; in S. hudsonianus appearing absent).
2. Spikelets solitary.
3. Perianth bristles 10-30 mm long. cottony; stems 3-angled; plants in small tufts along creeping rhizomes .... 1. S. hudsonianus
3. Perianth bristles less than 4 mm'long. not cottony; stems nearly round in cross-section; plants in dense tufts. not
rhizomatous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. S. cespitosus
2. Spikelets several, crowded into a flattened (2-ranked) compound spike .............. , . . . . . . . . . . . . . . . . . . .. 3. S. rufus
1. Spikelets subtended by 1 or more involucral bracts conspicuously longer than spikelets (fig. 517a).
4. Principal involucral bracts 1. appearing as continuation of stem (inflorescence lateral) (fig. 517a).
5. Spikelets usually sessile (fig. 517a).
6. Fertile stems weakly ascending. usually supported by water. often accompanied by long. hair-like leaves; spikelet
solitary .................................................................... 4. S. subterminalis
6. Fertile stems erect or arching; spikelets (1}2 or more. in glomerules.
7. Plants with elongate rhizomes; stems sharply 3-angled. at least at tip; anthers 1-3.5 mm long; bristles present.
8. Rhizomes flexible; scales lacking apical notch. margins smooth; styles 3-cleft; achenes 3-4 mm long ... 5. S. torreyi
8. Rhizomes firm; scales with obvious apical notch. margins (at least some) erose; styles 2-cleft (rarely 3-cleft);
achenes 1.9-3 mm long.
9. Involucral bract 1-3.5(5) cm long; scale tips less than 0.5 mm long .................... 6. S. americanus
9. Involucral bract 3.5-18 cm long; scale tips 0.5-1 mm long.
10. Spikelets usually more than 5 per inflorescence; achenes less than 2.5 mm long. 1.6 mm wide; bristles
slender. often longer than achenes .......................................... 7. S. deltarum
10. Spikelets fewer than 5 per inflorescence; achenes (2)2.4-3 mm long. greater than 1.6 mm wide;
bristles stout. rarely longer than achenes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 8. S. pungens
7. Plants with rhizomes absent or inconspicuous; stems round in cross-section or obscurely angled; anthers less
than 1 mm long; bristles usually reduced or absent.
11. Spikelets 1-3. 2.5-6 mm long. 2-3 mm wide; scales boat-shaped and prominently keeled (fig. 518g) .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .,. . . . . . . . . . . . . . . . . . . . . 9. S. koilolepis
11. Spikelets 1-12, 4-15 mm long, 2.5-5 mm wide; scales concave and slightly keeled.
12. Scales not striate, tips awned or cuspidate to acuminate; achenes conspicuously wrinkled transversally
(fig. 519f,g).
13. Styles 2-cleft; achenes plano-convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 10. S. hallii
13. Styles 3-cleft; achenes 3-angled ...................................... 11. S. saximontanus
12. Scales striate. tips blunt or slightly mucronate; achenes smooth or slightly pitted.
14. Achenes smooth, plano-convex; beak abruptly differentiated from achene body; bristles lacking, or
if present, slender .................................................... 12. S. smithii
14. Achenes slightly pitted, biconvex;' beak gradually tapering from achene body; bristles stout .... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13. S. hotarui
5. Spikelets usually pedunculate (figs. 520a, 521a), or occasionally short-stalked and appearing subsessile.
15. Involucral bracts 10-25 em long; stems 3-angled; emersed leaves with long triangular, channeled blades; sub-
mersed leaves with long ribbon-like blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14. S. etuberculatus
Cyperaceae / Sedge Family 225
15. Involucral bracts 1-7 cm long; stems round in cross-section; emersed and submersed leaves bladeless or with
inconspicuous blades.
16. Achenes obtusely 3-angled; style 3-cleft; bristles 2-4(5); spikelets all pedunculate ............ 15. S. heterochaetus
16. Achenes plano-convex; style usually 2-cleft; bristles usually 6; spikelets, at least some of them, sessile, glomeru-
late (rarely all pedunculate).
17. Stems soft and easily compressed, pale green when fresh; inflorescence open and lax, spikelets 12-125,
usually solitary or in clusters of 2 or 3; achenes usually partly exposed beyond scales; scales bright
orange-brown, with obscure darker spots, awns nearly straight, 0.2-1 mm long ....... 16. S. tabernaemontani
17. Stems hard, not easily compressed, dark olive-green when fresh; inflorescence usually stiff, spikelets 3-40,
usually solitary or in clusters of 2-7; achenes hidden by scales; scales usually dull grayish-brown, with
conspicuous bright brown spots, sometimes dark reddish-brown, awns usually contorted, about 0.5-1.5
mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 7. S. acutus
4. Principal involucral bracts 2 or more, leaf-like, not appearing as continuation of stem (inflorescence terminal) (figs. 523a,
524a,b).
18. Stems sharply 3-angled; rhizomes tuberous; spikelets 1-5 em long, 5-11 mm wide.
19. Inflorescence comparatively open; bristles usually persisting at base of achene; achenes 4-5 mm long, rounded
at summit, gradually tapering to beak.
20. Achenes acutely 3-angled; plants mostly of freshwater and fresh tidal zones .................. 18. S. fluviatilis
20. Achencs plano-convex to obtusely 3-angled; plants mostly of mildly saline tidal shores and marshes 19. S. cylindricus
19. Inflorescence comparatively compact or congested; bristles usually dropping off base of achene; achenes 3-3.5
mm long, flattened at summit, with abruptly differentiated beak.
21. Firm portion of lower leaf sheaths truncate or slightly concave at summit; scales reddish-brown (brown n. of
Mass.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20. S. robustus
21. Firm portion of lower leaf sheaths V-shaped at summit; scales pale grayish-brown to brown . . . . . . 21. S. maritimus
18. Stems obtusely 3-angled; rhizomes not tuberous; spikeIets 0.2-2.5 em long, 1-4(5) mm wide.
22. Bristles smooth, strongly contorted or curled, much longer than achene.
23. Scale midribs conspicuous, green; mature bristles usually hidden by scales; achene 1.1-1.5 mm long.
24. Mature stems lax with inflorescences drooping to, or near to, ground; lateral inflorescences 2 or 3, rays
with axillary bulblets; bristles usually 2-4 mm long ................................ 22. S. lineatus
24. Mature stems nearly erect; lateral inflorescences 0 or 1(2), rays lacking axillary bulblets; bristles usually
longer than 5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. S. pendulus
23. Scale midribs obscure, not green; mature bristles extending beyond scales, appearing wooly; achene 0.6-1
mm long.
25. Involucral bracts sticky at base; scales 2.2-3.1 mm long; achenes reddish or reddish-brown; plants with
thick elongate rhizomes, not forming tussocks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 24. S. longii
25. Involucral bracts not sticky at base; scales 1.1-2.1 mm long; achenes whitish; plants with short
branching rhizomes, forming tussocks.
26. Scales usually blackish; achenes maturing in late June and early July ............... 25. S. atroeinetus
26. Scales pale brown, brown, reddish-brown, or occasionally blackish; achenes maturing mid-July
to September.
27. Spikelets usually solitary, with distinct pedicels; scales pale brown; achenes maturing from
mid- to late July. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26. S. pedieellatus
27. Spikelets in glomerules with poorly developed pedicels; scales brown, reddish-brown, or some-
times blackish; achenes maturing from early August to September .............. 27. S. eyperinus
22. Bristles with teeth (obscurely so in S. divarieatus), usually straight, occasionally with 1 or 2 bends, barely, if at all,
longer than achene, or bristles sometimes absent.
28. Achenes biconvex, styles usually 2-cleft; leaf sheaths red-tinged at base ................... 28. S. microcarpus
28. Achenes 3-angled, styles usually 3-cleft: leaf sheaths greenish on lower portion of stem (but with lower
sheaths reddish in S. expansus).
29. Spikelets solitary with distinct pedicels; achenes sharply 3-angled, sides concave .......... 29. S. divarieatus
29. Spikelets usually in glomerules with poorly developed pedicels; achenes not sharply 3-angled, sides con-
vex, flat, or slightly concave.
30. Scales reddish-brown. as wide as long (excluding tips); stems 10-20-leaved ......... 30. S. polyphyllus
30. Scales brown or black, usually longer than wide (excluding tips); stems 2-10-leaved.
226 ANGIOSPERMS
31. Bristles with thick-walled, sharp-pointed teeth, densely arranged almost to bristle base (fig.
529b,f).
32. Rays of inflorescence ascending to spreading; leaf sheaths on lower portion of stem
strongly red-tinged at base; plants with long reddish rhizomes; achenes with bristles
readily dropping off; spikelets 1-2.5 mm wide ......................... 31. S. expansus
32. Rays of inflorescence arching; leaf sheaths on lower portion of stem green at base; plants
cespitose, with short brownish rhizomes; achenes with bristles persistent; spikelets 2-3
mm wide .............................................. 32. S. ancistrochaetus
31. Bristles with thin-walled, round-tipped teeth, restricted to upper two-thirds of bristle, or bris-
tles absent.
33. Bristles 0-3, shorter than achenes ................................ 33. S. georgian us
33. Bristles 5 or 6, shorter to slightly longer than achenes.
34. Scale tip awn-like (fig. 529j), about 0.4-0.7 mm long .................. 34. S. pallidus
34. Scale tip slightly mucronate, less than 0.4 mm long.
35. Mature stems lax, with inflorescence drooping to or near to ground; glomerule
usually with fewer than 15 spikeiets; plants of se. Va. in our range ... 35. S. flaccidifolius
35. Mature stems erect; glomerules often with more than 15 spikeiets; plants
throughout our range.
36. Lower leaf blades and sheaths usually nodulose-septate; scales usually
brown; bristles frequently longer than achene .............. 36. S. atrovirens
36. Lower leaf blades and sheaths smooth to slightly nodulose-septate; scales
usually black; longer bristles usually shorter than or as long as achene
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 37. S. hattorianus
1. S. hudsonianus (Michx.) Fern. Fig. 515 gular flowering stems. (Schoenoplectus subterminalis (Torr.)
Bogs, boggy meadows, marly swamps, wet gravels, and springy Sojak)
places, especially in calcareous areas. Se. Lab. and Nfld. w. to
Alask., s. to N.E.. n. N.Y., n. Mich., Minn., and w. Mont. Some 5. S. torreyi Olney Torrey's Three-square Fig. 515
authors have placed this taxon in the segregate genus Trichopho- Sandy and peaty shores of fresh to brackish waters. N.B. w. to
rum (as T. alpinum), and others have placed it in the genus Erio- Man., s. to 1.I., N.Y.. Del., Va .. n. Ohio, Ill., and Mo. (Schoenoplec-
phorum (as E. alpinum). Tucker and Miller (1990) have recently tus torreyi (Olney) Palla)
studied achene micromorphology in Eriophorum, including taxa
sometimes placed in Scirpus, and suggest that this taxon is bet- 6. S. americanus Pers. Fig. 516
ter treated in the genus Scirpus, and that it is strongly aligned Saline and brackish marshes. Coastal, N.S. s. to Fla., w. to Tex.
with S. cespitosus. (Eriophorum alpinum L.; Trichophorum alpinum and Mex.; inland in Mich., Mo., and Ida.; Pacific coast, Wash. s.
(1.) Pers.) to Calif.; S.Am. This species is treated as S. olneyi Gray in most
manuals, but Schuyler (1974a) has shown that name to have
2. S. cespitosus L. Deer-hair Sedge Fig. 514 been misapplied to this taxon. He further notes that hybrids
Cold bogs, fens, and peats of lowlands and mountain tops. Arc- with S. pungens may be abundant in transitional habitats, par-
tic regions s. to Nfld., N.S., Me., mts. of N.E.. disjunct to higher ticularly brackish tidal marshes and shores, and have been
mts. of N.C., Ga., and Tenn., w. to Mich., n. Ill., Minn., Man., called S. Xcontortus (Eames) T. Koyama (Schoenoplectus Xcon-
Mont., higher elevations to Utah and Oreg. Our plants are var. tortus (Eames) S. G. Smith). (Schoenoplectus americanus (Pers.)
callosus Bigel. (S. cespitosus var. delicatulus Fern.; Trichophorum Volk.)
cespitosum (L.) Hartm.)
7. S. delta rum Schuyler
3. S. rufus (Huds.) Schrad. Fig. 515 Alluvial tidal freshwater shores, rarely in saline areas. Fla., Ala.,
Saline and brackish marshes, rarely in freshwater peaty and La.; disjunct in Mo. and Kans. (Schoenoplectus deltarum
marshes. Coastal, Nfld., P.E.I., and e. Que. s. to sw. N.S. and sw. (Schuyler) Sojak)
N.B.; scattered localities, Hudson Bay, Man., N.W.T., and Alask.
Our taxon is var. neogaeus Fern. (Blysmus rufus (Huds.) Link; 8. S. pungens Vahl Three-square Bulrush Fig. 517
Schoen us rufus (Huds.) Schrad.) Freshwater, brackish, and saline shores and marshes. Nfld. w.
to s. Ont., Man., and s. B.C., s. to Fla., Tex., Calif., and Mex.;
4. S. subterminalis Torr. Water Bulrush Fig. 518 SAm. Schuyler (1974a) has shown that the name S. americanus
Ponds, bogs, lakes, and streams. Nfld. w. to Ont. and Minn., s. Pers. has been misapplied to this taxon. (Schoenoplectus pungens
to N.J., Pa., Mich., and n. Ill.; s. Alask. s. to nw. Wyo., nw. Mont., (Vahl) Palla)
n. Ida., and Calif.; disjunct stations in S.C., Ga., and Mo. This
species is often confused with Eleocharis robbinsii, because both 9. S. kOilolepis (Steud.) Gleason Fig. 518
species are often encountered only in the vegetative state, pro- Sands, depressions, seeps, shores, and damp woodlands. N.C.
ducing fine hair-like submersed leaves; E. robbinsii has more an- and Tenn. w. to sw. Ky., s. Mo., and Okla., s. to Fla., Ala., and
pe ria nt h
bristle involucral
br ac t
h
involucral
br ac t
.
'
bristle
Fig. 516. Scirpus american us: a. habit; b. stem, cross-section; c. inflorescence; d. achene (G&W).
228
Fig. 517. Scirpuspungens: a. habit; b. scale; c. achene; d. stem, cross-section (G&W).
229
b
230
a
I
i
I
b c
Tex.; Calif. (S. carinatus (Hook & Arn. ex Torr.) A. Gray; Isolepis 17. S. acutus Muhl. ex Bigel. Hardstem Bulrush Fig. 522
koilolepis Steud.) Freshwater to brackish marshes, ponds, and lakes. Nfld. w. to
Alask., s. to N.E., W.Va., N.C., Tenn., Mo., Tex., Ariz., Calif., and
10. S. hallii Gray Fig. 519 Mex. This species may hybridize with S. tabernaemontani and S.
Peaty and sandy shores. Rare, scattered localities; e. Mass., s. heterochaetus. Our taxon is var. acutus; however, fig. 522 is
Mich., Ill., Wisc., s. Mo., Kans., Neb., Ky., and Ga. (Scirpus supi- drawn from a plant of the western var. occidentalis S. Wats.; var.
nus var. hallii (Gray) Gray; Schoenoplectus hallii (Gray) S. G. acutus typically has scales with highly contorted awns in con-
Smith) trast with the shorter, straighter awns of fig. 522d,e. (Schoe-
noplectus acutus (Muhl. ex Bigel.) LOve & Love)
II. S. saximontanus Fern. Fig. 519
Shores of ponds, depressions, river bottomlands, and ditches. 18. S. j1uviatilis (Torr.) Gray River Bulrush Fig. 523
S.D. s. to Mo., Neb., Kans., Tex., Colo., and Mex.; Ohio. (Scirpus Fresh tidal shores along the coast, calcareous or alkaline shores
supinus var. saximontanus (Fern.) T. Koyama; Schoenoplectus saxi- and marshes inland. N.B. w. to Sask., s. to ne. Va., w. Pa., Ind.,
montanus (Fern.) J. Raynal) sw. Ky., and Mo.; N.M.; nw. Ida., Wash., and Oreg. (Bolboschoe-
nus j1uviatilis (Torr.) Sojak)
12. S. smithii Gray Fig. 519
Sandy, peaty, and muddy shores of lakes, ponds, and rivers; oc- 19. S. cylindricus (Torr.) Britt.
casionally tidal. N.B., Que., and Ont. w. to Minn., s. to N.E., Del., Brackish tidal transition zones in river systems. Coastal, Me. s.
ne. Va., nw. Pa., Ohio, n. Ind., and Ill. (S. smithii var. setosus to Ga.; inland in c. N.Y. This species often grows with S. robust us
Fern.) in regions where the habitat is transitional from brackish to sa-
line. Scirpus cylindricus grows with and is similar to S. robustus
1 3. S. hotarui Ohwi Fig. 519 and S. maritimus. and is therefore not easily distinguished. The
Marshes, bogs, wet peats, acid swamps, and sandy shores. Me. location in the tidal marsh is helpful in the identification of the
w. to s. Mich. and Minn., s. to Ga., Ala., and Miss. This species three species, but S. cylindric us is typically found farther up-
is treated as S. purshianus Fern. in most manuals, but the correct stream than S. robust us (Schuyler, 1967). (S. novae-angliae Britt.,
name is S. hotarui (Raymond, 1958; Tucker, 1992). (Scirpus de- S. robustus var. novae-angliae (Britt.) Beet!; Bolboschoenus novae-
bilis Pursh non Lam.; Scirpus smithii var. williamsii (Fern.) Bee- angliae (Britt.) S. G. Smith)
tle; Schoenoplectus purshianus (Fern.) M. Strong)
20. S. robustus Pursh Salt Marsh Bulrush Fig. 524
14. S. etuberculatus (Steud.) Ktze. Swamp Bulrush Fig. 520 Saline portions of tidal rivers and marshes. Coastal, M.l. and n.
Pond margins, freshwater and brackish marshes. Coastal plain; N.S. s. to Fla., w. to Tex. Schuyler (1974b) notes that plants
R.I.; Del. s. to Fla., w. to e. Tex.; inland n. to s. Mo. (Schoenoplec- found from Massachusetts southward usually have ovate spike-
tus etuberculatus (Steud.) Sojak) lets with reddish-brown scales, whereas those northward often
have narrowly ovate spikelets with brownish scales. Much inter-
IS. S. heterochaetus Chase Slender Bulrush Fig. 519 gradation occurs between the geographical regions. According
Calcareous shores of ponds, lakes, and marshes. Mass., w. Vt., to Schuyler (1974b) S. maritil11us var. fernaldii f. agonus Fern.
and sw. Que. w. to N.Y., Ont., Wisc., N.D., s. Man., and ne. Wyo., was misapplied by Fernald to this taxon. (S. maritimus var. l11ac-
s. to Pa., sw. Ky., Ill., Mo., and Okla; nw. Ida., Wash., Oreg., and rostachyus Michx.; S. maritil11us var. agonus (Fern.) Beetle; Bol-
ne. Calif. A similar Eurasian species, S. lacustris L. (Schoenoplec- boscllOenus robustus (Pursh) Sojak)
tus lacustris (1.) Palla), also with 3-cleft styles and 3-angled
achenes, but with many spikelets clustered and with 6 bristles, 21. S. maritimus L. Bayonet-grass, Prairie Bulrush Fig. 525
has been reported from Camden, New Jersey, and may be spar- Saline portions of tidal rivers and estuaries along the coast;
ingly naturalized near the East Coast. (Schoenoplectus heterochae- marshes, salt flats, and muddy shores inland. Coastal, Nfld. and
tus (Chase) Sojak) st. Lawrence R. system, Que., s. to N.J.; James Bay; Minn. w. to
s. B.C., s. to Tex., Calif., and Mex. Schuyler (1974b) notes that
16. S. tabernaemontani K. C. Gmel. Great Bulrush, Soft-stem variation within S. maritil11us (S. paludosus) is correlated with the
Bulrush Fig. 521 upstream-downstream distribution along rivers or with topo-
Fresh to brackish shores of ponds. lakes, streams, and marshes. graphical distribution in marshes. Upstream plants or plants in
Nfld. w. to Alask., s. to Fla., Tex., N.M., Calif., and Mex.; S.Am. the upper portions of marshes tend to be taller with open inflo-
This species is referred to as S. validus Vahl in most manuals. A rescences and browner scales than plants in more saline sites
similar species, S. californicus (C. A. Mey.) Steud. (Schoenoplectus downstream or in lower portions of marshes. Plants with more
californicus (C. A. Mey.) Palla), with obtusely 3-angled upper open inflorescences occasionally have achenes with prominent
parts of stems and plumose bristles (fig. 521), is reported near dorsal bulges. Schuyler (pers. comm.) now treats S. maritimus
the edge of our range in Oklahoma. This species hybridizes com- as a worldwide taxon, including S. paludosus. (S. paludosus Nels.;
monly with S. acutus and rarely with S. heterochaetus. Smith S. paludosus var. atlanticus Fern., S. l11aritimus var. paludosus
(1995) considers S. Xsteinmetzii Fern. (Schoenoplectus Xstein- (Nels.) Kuek., S. maritil11us var. fernaldii (Bickn.) Fern.; Bolbo-
l11etzii (Fern.) S. G. Smith) to be a hybrid between S. tabernael11on- schoenus l11aritimus (1.) Palla)
tani and S. heterochaetus, a plant previously known only from a
single stream in Maine, but with new localities recognized from 22. S. lineatus Michx. Fig. 525
Massachusetts, New York, Quebec, Ontario, Wisconsin, Iowa, Bottomlands, marshes, ditches and wet woods. Coastal, se. Va.
and Minnesota; although it is often sterile, sometimes achenes s. to Fla., w. to La.; se. Mo. (S. fontinalis Harper, S. fontinalis var.
with fully formed seeds are produced. (Schoenoplectus tabernae- virginiana Fern.)
l110ntani (K. C. Gmel.) Palla; Scirpus validus Vahl; Schoenoplectus
validus (Vahl) Love & LOve)
h
II:
',il
Ii', c
,
a I
" III
I I,
f
I: I
',I
b e
Fig. 521. Scirpus tabernaemantani: a. habit; b. upper stem. cross-section; c. scale; d. achene (G&W).
Scirpus cali/arnicus: e. upper stem. cross-section; f. leaf sheath; g. scale; h. achene (G&W).
234
g
involucral
bract
stamen ----li!!,.~
Fig. 522. Scirpus acutus: a. habit; b, c. inflorescence variations; d. spikelet; e. scale; f. flower; g. achene;
h. achene, cross-section (Mason).
235
~
V Q )k O I J
Fig. 524. Scirpus robustus: a. habit; b. inflorescence; c. scale; d. achene; e. achene, cross-section (Mason).
237
g
23. S. pendulus Muhl. Fig. 525 30. S. polyphyllus Vahl Fig. 528
Meadows. marshes. stream banks. wet peat. and ditches. N.B. Low woods. swamps. stream banks. and marshes. Se. N.H. and
and Me. w. to Iowa and e. Colo . s. to Fla .. Miss . Okla . Tex .. and Mass. sw. to s. Ill., s. to Ga. and Ala.
n. Mex.; Oreg. and Calif. (S. lineatus sensu acutt. non Michx.)
31. S. expansus Fern. Fig. 529
24. S. longii Fern. Fig. 527 Marshes. meadows. and low thickets. Me. w. to Ohio and Mich .
Meadows. swamps. and marshes. Coastal. N.S. and se. N.H. s. to s. to Ga. (S. sylvaticus L. pro parte)
s. N.}.; inland to w. Me. Rare. locally abundant in s. N.J. Schuyler
(1962b) has noted that a large portion of herbarium specimens 32. S. ancistrochaetus Schuyler Fig. 529
have charred leaves and has suggested that fire may be an im- Pond margins. stream banks. and bogs. Very rare and local. se.
portant factor in stimulating flowering. Vt.. c. Mass . N.Y.. Pa . and Va.
25. S. atrocinctus Fern. Fig. 526 33. S. georgianus Harper Fig. 529
Meadows. swamps. marshes. and ditches. often in disturbed Moist meadows. marshes. and ditches. P.E.I. w. to Wisc. and
areas. Nfld. w. to n. Alta. and B.C.. s. to W.Va . Mich . ne. Iowa. Neb. s. to Ga . Tenn . and e. Tex. This species is rare in the
S.D.. and Wash. This species often grows with and hybridizes northern part of its range. (S. atrovirens var. georgianus
with S. cyperinus (Schuyler. 1967). (S. cyperinus var. brachypodus (Harper) Fern.)
(Fern.) Gilly)
34. S. pallidus (Britt.) Fern. Fig. 529
26. S. pedicellatus Fern. Fig. 527 Low ground. marshes. stream margins. and ditches. Minn. and
Marshes. wet lowlands. and stream margins. E. Que. w. to ne. Man. w. to Wash .. s. to Wisc . Mo .. e. Tex. N.M .. and Ariz.;
Minn .. s. to n. N.J. n. Ind . s. Ill.. and n. Mo. This species closely e. Pa. (S. atrovirens var. pallidus Britt.; S. atrovirens ssp. pallidus
resembles S. atrocinctus. but tends to be more robust and has D. LOve & Bern.)
pale brown instead of blackish scales. (S. cyperinus var. pedicella-
tus (Fern.) Schuyler) 35. S.flaccidifolius (Fern.) Schuyler Fig. 531
Bottomlands. Se. Va. and ne. N.C. (S. atrovirens var. flaccidifolius
27. S. cyperinus (L.) Kunth Fig. 527 Fern.)
Wet meadows. swamps. marshes. and ditches. Nfld. w. to Minn.
and Man .. occasionally to s. B.C . s. to Fla. and e. Tex. This spe- 36. S. atrovirens Willd. Fig. 530
cies is extremely variable. the northern plants tending to have Meadows. marshes. stream margins. bogs. and ditches. Nfld.
black involucres and bracts. and spikelets with short. ovate. and s. Que. w. to Minn. and Sask. s. to Ga . Tenn . Mo . and e.
brownish scales (Schuyler. 1967). (S. cyperinus var. pelius Fern . Tex.; Ariz. In late summer. plants frequently produce leafy tufts
S. cyperinus var. eriophorum (Michx.) Ktze . S. cyperinus var. ru- in the inflorescence (fig. 530c). This species is closely related to
bricosus (Fern.) Gilly. S. eriophorum Michx .. S. rubricosus Fern.) S. ancistrochaetus. S. flaccidifolius. S. georgian us. S. hattorianus.
and S. pallidus. and occasionally hybridizes with them (see
28. S. microcarpus Presl Fig. 528 Schuyler. 1967).
Marshes. low thickets. meadows. and ditches. S. Lab. and Nfld.
w. to Alask.. s. to N.E.. W.Va .. Mich .. n. Ill.. Iowa. N.M .. Ariz .. 37. S. hattorianus Makino Fig. 526
and s. Calif. This species may hybridize with S. expansus. (S. mi- Meadows. marshes. and ditches. Nfld. to w. Ont.. s. to N.C .
crocarpus var. rubrotinctus (Fern.) M. E. Jones; S. microcarpus var. Ohio. Ind.. and Wisc. This species has frequently been treated
longispicatus Peck; S. rubrotinctus Fern.; S. expansus f. globulosus as S. atrovirens var. georgianus by American authors.
Fern.)
12. Lipocarpha
Annuals. densely tufted; stems 3-angled. leafy at base; leaves narrow. involute; inflorescence a cluster of several sessile. densely
flowered spikelets. subtended by 2 long and several short involucral bracts; flowers bisexual. subtended by 1 or 2 small delicate
bractlets (easily overlooked). bristles absent; fruit an achene. lacking a tubercle.
1. Leaves reduced to sheaths at base of stem. with very short blades (fig. 532a); achenes 0.5-0.8 mm long. subtended by 1 small
bractlet as well as spikelet scale ............................................................ 1. L. micrantha
1. Leaves with well-developed blades (fig. 533a); achenes 0.8-1 mm long. subtended by 2 small bractlets as well as spikelet scale
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. 1. maculata
b
c
d c
Fig. 528. Scirpus microcarpus: a. habit; b. portion of inflorescence; c. scale; d. achene (Mason).
Scirpus divaricatus: e. inflorescence (Beal).
Scirpus polyphyllus: f. achene (F).
242
a
, c
Fig. 530. Scirpus atrovirens: a. basal portion of plant (C&C); b. upper portion of plant (C&C); c. inflorescence
with leafy tufts (Braun); d. portion of inflorescence (Braun); e. spikelet (Braun); f. achene (C&C);
g. scale (C&C).
244
Fig. 531. ScirpusjIaccidi!oIius: a. inflorescence; b. achene; c. scale (PB).
245
e
Fig. 532. Lipocarpha micrantha: a. habit (G&W); b. habit (F); c. inflorescence (G&W); d. scale (G&W);
e. achene (G&W).
246
Fig. 533. Lipocarpha maculata: a. habit; b. inflorescence; c. scale and bractlets surrounding flower; d. achene
(G&W).
247
248 ANGIOSPERMS
1. 1. micrantha (Vahl) Tucker Fig. 532 2. 1. maculata (Michx.) Torr. Fig. 533
Sandy shores of ponds and streams. Que. and s. Me. w. to ant., Wet sands along pond and stream margins, wet peaty areas,
Minn., s. Ida., B.C., and Wash., s. to Fla., Tex., Ariz., Calif., and springy places, borders of swamps. Coastal plain, Va. s. to Fla.,
Mex.; C.Am. and S.Am. Previously treated in Hemicarpha, e. to Ala.
Tucker (pers. comm.) feels that group should be recognized as
congeneric with Lipocarpha. (Scirpus micranthus Vahl; Hemicar-
pha micrantha (Vahl) Britt.)
13. Fimbristylis
Perennial or annual herbs, grass-like, forming tufts, or culms arising singly from a creeping rhizome; leaves basal; spikelets solitary
and terminal on scapes or arranged in umbelliform inflorescences; flowers bisexual, perianth absent; stamens 1-3; styles 2- or 3-
cleft, style base flat or swollen, but not persistent at fruit maturity; achenes biconvex or 3-angled, shiny, lacking a tubercle.
Fimbristylis differs from Scirpus in that the base of the style is swollen; it differs from Rhynchospora by the style dropping off and
not forming a tubercle.
l. Styles 3-cleft (fig. 534d); achenes 3-angled (fig. 534c) or obovoid and obscurely 3-angled (fig. 535b), but not biconvex.
2. Spikelets narrowly linear to lanceolate (fig. 534a); achene surface smooth to warty ..................... l. E autumnalis
2. Spikelets broadly ovoid to globose (fig. 535a); achene surface reticulate (fig. 535b) ........................ 2. E miliacea
l. Styles 2-cleft; achenes biconvex.
3. Spikelets appearing sessile, in dense, capitate clusters (fig. 534g); achenes 0.4-0.5 mm long ................... 3. E vahlii
3. Spikelets on rays (fig. 534e), rarely congested; achene 1-l.5 or 4-6 mm long.
4. Plants densely tufted, forming large tussocks, up to 1.5(2) m tall, bases deeply set in substrate .............. 4. E castanea
4. Plants rhizomatous or tufted, but not forming large tussocks, bases not deeply set in substrate.
5. Plants perennial; bases hard and bulbous, with rhizomes either slender and somewhat elongate, or short, thick, and
knotty or bulbous.
6. Rhizomes thick, with hard, bulbous, persistent bases, giving knotty appearance; leaf blade lacking ligule .....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5. E puberula
6. Rhizomes slender; leaf blade with ligule of short hairs .................................... 6. E caroliniana
5. Plants annual; bases soft, lacking rhizomes.
7. Leaves linear, 1-2(4) mm wide, ligule of short hairs present; achenes 1-1.2 mm long ................. 7. E annua
7. Leaves linear-filiform, less than 1 mm wide, ligule absent; achenes 4-6 mm long ................. 8. F. perpusilla
1. F. autumnalis (L.) R. & S. Fig. 534 5. F. puberula (Michx.) Vahl Fig. 535
Moist to wet sands, peats, clays, and shores, often in disturbed Wet savannas and pinelands, wet peats, meadows, and prairies.
sites. C. Me. and sw. Que. w. to ant., Mich., Minn., and se. S.D., s. Coastal plain, 1.1., N.Y., s. N.J., and se. Pa. s. to Fla., w. to e. Tex.;
to Fla., Tex., and Mex.; Trop. Am. (F. autumnalis var. mucronulata inland n. in the Midwest to Okla., Neb., Mo., Ill., Ind., Mich.,
(Michx.) Fern.; F. /rankii Steud.) and sw. Onto Our taxon is var. puberula. (F. drummondii (Torr. &
Hook) Boeck.; F. puberula var. drummondii (Torr. & Hook) Ward)
2. F. miliacea (L.) Vahl Fig. 535
Wet sandy peats, mud flats, shores, savannas, and wet ditches 6. F. caroliniana (Lam.) Fern. Fig. 536
and fields. Coastal plain, N.C. s. to Fla., w. to Tex. and Mex.; Saline, alkaline, or weakly acid sands or sandy peats of beaches,
inland n. to sw. Ky. and Ark.; w.I. and C.Am. dune swales, lakeshores, ditches, and savannas. Coastal plain,
N.J. s. to Fla., w. to e. Tex. and Mex.; Cuba.
3. F. vahlii (Lam.) Link Fig. 534
Damp sands, silts, and clays on margins of ponds, lakes, and 7. F. annua (All.) R. & S. Fig. 536
streams, and disturbed bottomlands. S.C. w. to Ohio, sw. Ky., Ill., Moist open sites, savannas, grasslands, and disturbed sites.
s. Mo., se. Kans., and Okla., s. to Fla. and Tex.; Calif., Ariz., and Coastal plain, se. Pa. s. to Fla., w. to Tex., Ariz., and Mex.; ex-
Mex.; C.Am. tending n. along the Mississippi embayment to e. Okla., s. Mo.,
and s. Ill.; w.I. and C.Am.; intro. from Eurasia. (F. baldwiniana
4. F. castanea (Michx.) Vahl Fig. 534 (Schult.) Torr.)
Moist sands, muck, or marl, saline coastal marshes, and dune
swales. Strictly coastal, L.I., N.Y., s. to Fla., w. to e. Tex. and 8. F. perpusilla Harper Fig. 536
Mex.; w.I. Alluvial soils. Delmarva Pen.; se. N.C., e. S.C., and sw. Ga.
stigmas
It--- style
c
d
Fig. 534. Fimbristylis autumnalis: a. habit (C&C); b. ligule (Kral, 1971); c. achene (C&C); d. stigmas and styles
(Kral. 1971).
Fimbristylis castanea: e. habit, portion of plant removed from tussock; f. achene (C&C).
Fimbristylis vahlii: g. habit; h. achene (G&W).
249
Fig. 535. Fimbristylis miliacea: a. habit; b. achene (G&W).
Fimbristylis puberula: c. habit; d. achene (G&W).
250
b
Fig. 536. Fimbristylis caroliniana: a. habit; b. scale; c. achene; d. section ofleaf (Kral. 1971).
Fimbristylis annua: e. habit; f. achene (Kral. 1971).
Fimbristylis perpusilIa: g. habit; h. achene (G&W).
251
252 ANGIOSPERMS
The grasses belong to the fourth-largest family, yet relatively few of its members occupy aquatic or wetland habitats, unlike the
sedges (see pp. 121-251) and the rushes (see pp. 97-120), which they superficially resemble. The grasses can be readily distinguished
by the following features (fig. 537): stems round in cross-section and hollow, except at nodes; leaves 2-ranked, sheathing stem,
sheath open with edges often overlapping, and having a ligule at its junction with blade; flowers borne in spikelets, each spikelet
with 2 glumes at base and I-several florets, each floret consisting of a lemma and a palea enclosing the flower with its lodicules
(modified petals), a pistil, and 3 or 6 stamens; fruit a grain.
Some grasses, especially Glyceria (Manna Grass) and Zizania (Wild-rice), are sometimes found submersed and in a vegetative state
with floating leaves similar to those of Vallisneria. These can be recognized as grasses by the presence of a ligule at the junction of
the blade and sheath. The presence of a rhizome in Glyceria will help to distinguish it from Zizania, which being annual lacks a
rhizome and is easily pulled.
REFERENCES: Campbell, 1985; Dore and McNeill, 1980; Gould, 1975; Gould and Shaw, 1983; Hitchcock, 1950; Pohl, 1980; Soder-
strom et aI., 1987; Voss, 1972.
1. Plants woody, with much-branched stems, often only vegetative, but when flowering (at intervals of 4-6 years), spikelets 4-7
cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Arundinaria
1. Plants herbaceous (if appearing woody, then unbranched), flowering regularly, spikelets less than 4 cm long.
2. Florets unisexual.
3. Plants usually less than 0.6 m tall, dioecious; florets several per spikelet.
4. Inflorescence of both pistillate and staminate plants a dense, spike-like panicle (fig. 542a); plants of salt marshes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. Distichlis
4. Inflorescence of pistillate plants subglobose, inflorescence of staminate plants an open panicle; plants of non-saline
sites (E. reptans) ................................................................. 10. Eragrostis
3. Plants usually more than 1 m tall, monoecious; pistillate florets 1 per spikelet, staminate florets sometimes 2.
5. Pistillate spikelets borne on upper branches, staminate spikelets borne on lower branches of inflorescence (fig. 539b)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Zizania
5. Pistillate and staminate spikelets borne on same branches of inflorescence.
6. Pistillate spikelets on upper portion of inflorescence branches, staminate spikeIets on lower portion (fig. 540a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4. Zizaniopsis
6. Pistillate spikelets on lower portion of inflorescence branches, staminate spikelets on upper portion (fig. 541a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Tripsacum
2. Florets bisexual, or with 1 bisexual floret along with empty lemmas, or with 1 or 2 staminate florets.
7. Spikelets with 2-several bisexual florets.
8. SpikeIets sessile (fig. 542c,e), 2 at a node, 2-ranked on opposite sides of rachis, forming simple spikes; glumes and
lemmas tipped with long awn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Elymus
8. Spikelets pedicellate (fig. 543b), not forming simple spikes (some species with spike-like inflorescences, but spikelets
not sessile); glumes and lemmas awned or not.
9. Plants large, coarse reeds, 1-6 m tall; inflorescence plume-like; rachilla bearded, with abundant long silky hairs
as long as or longer than lemmas (fig. 543c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Phragmites
9. Plants less robust, seldom exceeding 1.5 m tall; inflorescence various, but not plume-like; rachilla lacking beard
or, if bearded, then hairs short.
10. Lemmas densely villous on callus (fig. 544c); ovary pubescent at summit; plants of northern Great Plains
....................................................................... 8. Scolochloa
10. Lemmas not villous on callus; ovary glabrous.
11. Lemmas 3-nerved.
12. Glumes and lemmas rounded dorsally, not keeled; florets lacking cobwebby hairs at base.
13. Glumes and lemmas firm, yellowish-brown to purplish, truncate and irregularly cut at sum-
mit; plants of northern sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. Catabrosa
13. Glumes and lemmas thin and dry, pale to red, acute to acuminate; plants widespread .... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Eragrostis
12. Glumes and lemmas keeled; florets with a tuft of cobwebby hairs at base (except P. annual . . . . . .. 11. Poa
stem
stigma
\\\u\';IIIli+!_ base of
blade
rachilla
sheath
filament}
stamen
+------anther
c d
Fig. 537. Structure of Poaceae (grasses): a. section of stem with leaf base. generalized; b. floret. generalized;
c. spikelet. side view. generalized; d. spikelet. diagram of longitudinal section (F).
253
254 ANGIOSPERMS
1. Arundinaria (Cane)
Perennials, woody; stems 1-5 m tall, much-branched, leafy; leaves with both outer and inner ligule, sheath summit with conspicuous
bristles; inflorescence a raceme or panicle; spikelets several-flowered, 4-7 cm long; reproductive stems dying after fruiting.
1. A. gigantea (Walt.) Muhl. Giant Cane, Wild Cane, Switch bayment; ssp. tecta (Walt.) McClure, predominately ofthe Atlan-
Cane, Wild Bamboo Fig. 538 tic Coast; and the intermediate ssp. macrosperma (Michx.) Mc-
Riverbanks, swamps, sloughs, and bayous, often forming cane- Clure. These three subspecies are extremely difficult to distin-
brakes. Del. and Md. w. to s. Ohio, s. Ind., s. Ill., and s. Mo., s. to guish without considerable experience with this group. The
n. Fla. and se. Tex. In this, our only native bamboo, McClure stems are used in making baskets and for fishing rods.
(1973) recognizes ssp. gigantea, centered on the MisSissippi em-
Perennial herbs, halophytic; stems spreading to ascending; leaves stimy erect-spreading, conspicuously 2-ranked; stems usually hid-
den by overlapping leaf sheaths; plants dioecious; inflorescence a spike-like panicle.
1. D. spicata (L.) Greene Spike Grass Fig. 542 little from our inland populations (nw. Minn., w. Iowa, and w.
Salt marshes and wet alkaline soil. Coastal, P.B.I., e. Que., and Mo.) said to belong to var. stricta (Torr.) Beetle. The species, with
C.B.!. s. to Fla., w. to Tex. and Mex.; Great Plains; West Coast; its conspicuously 2-ranked leaves, is distinctive even in the vege-
Cuba. Our coastal plants belong to var. spicata, which differs tative stage.
3. Zizania (Wild-rice)
Annuals, with short roots (easily pulled up); stems slender to thick, reaching 4 m tall; leaves flat, long, often floating when young;
spil(elets in much-branched panicles; flowers unisexual. spikelets of the upper inflorescence branches pistillate, spikelets of lower
branches staminate, the latter red, greenish, or yellow; lemmas of pistillate spikelets long-awned. Zizania is sometimes confused
with Zizaniopsis.
Both species of Zizania are important food for wildlife, especially wildfowl. Zizania palustris was (and locally still is) an important
food source for Native Americans, especially in Wisconsin and Minnesota, and is now commercially harvested for domestic markets.
REFERENCES: Aiken, 1986; A&en et at., 1988; Chambliss, 1940; Dore, 1969; Duvall and Biesboer, 1988; Fassett, 1924; Terrell et aI.,
1997; Warwick and Aiken, 1986.
1. Pistillate lemmas thin and papery, dull, finely striate; rudimentary (aborted) spikelets very narrow, thread-like (fig. 539f).
2. Plants 1-4 m tall; leaves (5)10-72 mm wide; ligules (6)10-20(25) mm long; awn of pistillate lemma 10-70(90) mm
long ..................................................................... lao Z. aquatica var. aquatica
2. Plants 0.3-1 m tall; leaves 3-12(20) mm wide; ligules ca. 3 mm long; awn of pistillate lemma 1-8 mm long; plants of St.
Lawrence Seaway ............................................................ 1b. Z. aquatica var. brevis
1. Pistillate lemmas firm and tough, lustrous, coarsely ribbed; rudimentary (aborted) spikelets 1.5-2 mm wide (fig. 539d).
3. Plants 0.7-1.5 m tall; leaves 3-15(20) mm wide, Iigules 3-5(10) mm long; lower pistillate branches with 2-8 spikelets
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2a. Z. palustris var. palustris
3. Plants 0.9-3 m tall; leaves (10)20-40 mm wide, Iigules 10-15 mm long; lower pistillate branches with 9-30 spikelets
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2b. Z. palustris var. interior
\;
b c
I ...
~...
d e f
Fig. 539. Zizania palustris var. interior: a. basal portion of plant (Hitchcock); b. leaf and panicle (Hitchcock);
c. pistillate spikelet. fertile. filled with grain (Hitchcock); d. pistillate spikelet. aborted (F).
Zizania aquatica var. aquatica: e. pistillate spikelet. fertile. filled with grain; f. pistillate spikelet.
aborted (F).
257
Fig. 540. Zizaniopsis miliacea: a. habit; b. stam
inat e spikelet; c. pistillate spikelet:
d. ripe grai n (Hitchcock).
258
Fig. 0541. TripslIcum dllciy[oides: a. habit; b. pistillate spikelet; c. pair of staminate spikelets (Hitchcock).
259
c
Perennial herbs; stems 1-3 m or more tall; leaves long. narrow. with rough margins; spikelets unisexual. pistillate and staminate
spikelets on same branches. staminate spikelets on lower half of branch. pistillate spikelets on upper half.
Perennial herbs. with thick creeping rhizomes; stems erect; leaves broad. flat; spikelets in terminal or axillary spikes. segmenting at
the joints at maturity; flowers unisexual, staminate florets in upper portion of spike. pistillate florets in lower portion.
6. Elymus (Wild-rye)
Perennial herbs; inflorescence a dense. bristly spike; spikelets 2 or 3 at each rachis node. each several-flowered. lemma long-awned.
1. Clumes 0.8-2 rum wide. flat; spike erect; leaves 3-13 mm wide ...................................... 1. E. virginicus
1. Clumes 0.4-0.8 rum wide. bristle-like; spike arching; leaves 10-20 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. E. riparius
7. Phragmites (Reed)
Perennial herbs. arising from rhizomes. colonial; stems 2-6 m tall; leaf blades broad; inflorescence large. plume-like; rachis bearded.
with long silky hairs.
Phragmites is sometimes confused with Zizania (Wild-rice). but in the young. vegetative state, it can be distinguished by its thick
rhizomes, which make it difficult to pull up.
1. P. australis (Cav.) Trin. ex Steud. Common Reed Fig. 543 tive because of its large size and colonial habit. its large plume-
Freshwater. brackish. or saline marshy areas, pond margins, like inflorescence. and the tendency of its leaves to be present
ditches, and borders of salt marshes and tidal waters. Que. w. to on only one side of the stem. (P. communis Trin.; P. maximus var.
B.C., s. to Fla .. Tex .. Calif.. and Mex.; C.Am. This plant is distinc- berlandieri (Fourn.) Moldenke of Fassett)
Fig. 543. Phragmites australis: a. habit; b. spikelet; c. floret (Hitchcock).
262
Fig. 544. Scolochloa festucacea: a. habit; b. spike
let; c. floret (Hitchcock).
263
264 ANGIOSPERMS
8. Scolochloa
Perennial herbs. stout. arising from rhizomes. up to l.5 m tall; leaves flat; inflorescence an open panicle; spikelets several-flowered.
with large glumes.
Perennial herbs. arising from rhizomes; stems creeping at base and rooting at nodes; leaves flat; inflorescence a pyramidal panicle;
spikelets relatively short. obtuse. florets loosely overlapping; lemmas 3-nerved. truncate at tip.
l. E. hypnoides (Lam.) BSP. Creeping Love Grass Fig. 546 mud of bottomlands. and ditches; weedy in wet disturbed sites.
Muddy shores and sandbars of streams and rivers. Me. and Que. Me. and sw. Que. w. to N.D. and e. Wash . s. to Fla . Ariz .. Calif..
w. to N.D.. s. to Fla . e. Tex .. and Mex.; w. Wash. s. to Calif.; w.I. and Mex.; rare in western states. (E. caroliniana (Spreng.)
and S.Am. Scribn.; E. diffusa Buck!.)
2. E. reptans (Michx.) Nees Creeping Love Grass Fig. 546 4. E. frankii C. A. Mey. ex Steud. Sandbar Love Grass
River margins and shores. and frequently beds of streams and Fig. 545
ponds after drought. Ky. and s. Ill. w. to S.D. s. to n. Fla .. e. Tex . River sandbars and muddy banks. and seasonally wet soils. N.H.
and e. Mex; S.Am. (Neeragrostis reptans (Michx.) Nicora) and sw. Que. w. to s. Ont. and Minn .. s. to Fla .. Ark.. and Okla.
(E. frankii var. brevipes Fassett)
3. E. pectinacea (Michx.) Nees Small Love Grass Fig. 545
Sandy and gravelly shores. river sandbars. shallow water and
Perennial or annual herbs. arising from rhizomes or tufted; leaves with flat blades ending in boat-shaped apex; spikelets small. often
only 2- or 3-flowered; upper glume shorter than lowest lemma; glumes and lemmas keeled and distinctly nerved. awnless. usually
with a tuft of cobwebby hairs at base.
1. Plants annual, 5-20 cm tall; lemmas lacking cobwebby tuft of hairs at base ................................. 1. P. annua
1. Plants perennial. over 25 cm tall; lemmas with cobwebby tuft of hairs at base.
lemma
266
Poaceae (Gramineae) / Grass Family 267
2. Panicle many-flowered. branches usually 3-5 per node; plants somewhat robust. usually with about 6 leaves ...... 2. P. palustris
2. Panicle few-flowered. branches 2 per node; plants slender. usually with 2 or 3 leaves ...................... 3. P. paludigena
2. P. palustris L. Fowl Bluegrass Fig. 548 Poa autumnalis, P. sylvestris, and P. trivialis tend to grow in drier
Meadows. shores. and thickets. Lab. and Nfld. w. to sw. N.W.T. sites. but may occur in alluvial soils. springy places. or seeps.
and Alask.. s. to N.E . Va .. w. N.C .. Ohio. n. II!.. nw. Mo .. n. N.M ..
and n. Calif. This species is often found only in vegetative condi-
tion. with floating leaves.
Perennial herbs; stems slender, weak. and decumbent or procumbent; panicles contracted. spikelets small. awnless; lemmas dorsally
rounded. This genus is similar to Glyceria, which differs in having closed sheaths.
1. P. fernaldii (Hitchc.) E. G. Voss Fig. 548 3. P. maritima (Huds.) Par!' Fig. 548
Shallow water and wet, open places. Nfld. w. to Mich. and Saline shores and saIt marshes. Coastal. st. P. et Miq. and Que.
Minn.. s. to N.E., L.r., N.Y.. and Pa. (Glyceria fernaldii (Hitchc.) s. to N.B.
st. John; G. pallida var. fernaldii Hitchc.; Torreyochloa fernaldii
(Hitchc.) Church) 4. P. langeana (Ber/.) S0ren. ex Hult.
Saline shores and salt marshes. Coastal. Green!' and Baffin r.
2. P. pallida (Torr.) Clausen Fig. 549 s. to Nfld .. Me .. N.H .. and Conn. S0renson (Hulten. 1950) has
Shallow water, pond margins, sloughs, wet open places. Sw. N.S. determined P. langeana to be the name with priority and recog-
and s. Me. w. to s. Ont. and Mich . s. to Va .. Tenn. and se. Mo. nizes three subspecies. Our material best fits ssp. alaskana
(Glyceria pallida (Torr.) Trin.; G. neogea of Fassett. non Steud.; (Scribn. & Merr.) Soren. (P. paupercula var. alaskana (Scribn. &
Torreyochloa pallida (Torr.) Church) Merr.) Fern. & Weath.)
Perennial herbs. arising from rhizomes; stems thick. erect; leaf blades broad and flat. sheaths closed; spikelets several-flowered.
awnless; lemmas dorsally rounded. conspicuously nerved. Glyceria is unique among the grasses in having closed sheaths.
REFERENCES: Church. 1949; Dore. 1947; Freckmann and Reed. 1979; Gutteridge. 1954.
268
a
1. G. acutiflora Torr. Fig. 550 Minn., s. to N.E., Va., mts. of Tenn., ne. Ohio, and n. Ill. A taller
Shallow water and wet soils. S. N.H. w. to Mich., s. to Va., W.Va., plant with somewhat smaller spike\ets has been treated as var.
Tenn., and Mo. laxa (Scribn.) Hitchc., but it intergrades with var. canadensis and
may not deserve varietal status.
2. G. fluitans (L.) R. Br. Floating Manna Grass Fig. 550
Shallow water. Nfld., Gaspe Pen., Que., and N.S.; Nantucket I., 9. G. striata (Lam.) Hitchc. Fowl Manna Grass, Meadow
Mass.; N.Y.; Black Hills, S.D. Grass Fig. 549
9a. G. striata var. striata
3. G. borealis (Nash) Batchelder Small Floating Manna Grass
Fig. 550 Moist meadows, wet places, and pond margins. Nfld. w. to Alta.,
s. to n. Fla. and e. Tex.
Shallow water and wet places. Nfld. w. to Man., N.W.T., and
Alask., s. to N.E., Pa .. n. Ohio, n. TIl., Iowa, S.D., mts. to Colo.. 9b. G. striata var. stricta (Scribn.) Fern.
Ariz., and Calif. Wet meadows and shallow pond margins. Nfld. w. to Man., sw.
N.W.T., and Alask., s. to n. N.E., n. TIl., Iowa, mts. to N.M., Ariz.,
4. G. septentrionalis Hitchc. Eastern Manna Grass Fig. 550 Oreg., and Calif. (G. striata ssp. stricta (Scribn.) Hult.)
Shallow water, marshy shores, swamps, woodland pools, and
wet places. Mass. w. to s. Ont., Mich., and Minn., s. to n. Ga., 10. G. grandis S. Wats. Reed Manna Grass, Reed Meadow
Ala .. La .. and e. Tex. Grass Fig. 550
Stream banks, marshes, and wet places. Nfld. w. to Man., sw.
5. G. arkansana Fern. Fig. 549 N.W.T., and Alask., s. to Va., mts. of Tenn., ne. Ohio, n. Ill., Neb.,
Wet bottomlands, swamps, sloughs, and shallow water of pond mts. to Colo., N.M., Ariz., and Oreg. This taxon is sometimes
and lake margins. Ill. w. to Okla., s. to Ark. and La.; se. Va. treated as a subspecies of G. maxima. (G. maxima ssp. grandis (S.
Wats.) HulL)
6. G. obtusa (Muhl.) Trin. Fig. 549
Swales, open swamps, shores, and wet peaty and sandy soils. 11. G. maxima (Hartm.) Holmb. Reed Grass, Tall Manna
Usually coastal. N.S. s. to N.C. Grass
Marshes and wet ditches. Local, s. Ont. and Wisc.; intro. from
7. G. melicaria (Michx.) F. T. Hubbard Eurasia. Canadian introductions are more robust than the na-
Wet woods, swamps, and wet peaty soils. N.B. w. to Ohio and tive G. grandis and are said to be reddish on the lower portion of
Ont., s. to N.E., Md., and mts. of N.C., Ky., and Tenn. stems, making them distinctive in the vegetative state (Gut-
teridge, 1954). Freckmann and Reed (1979) note that in Wis-
8. G. canadensis (Michx.) Trin. Rattlesnake Manna Grass consin this species has potential as a troublesome wetland weed.
Fig. 550
Bogs, meadows, swales, shores, and wet places. Nfld. w. to
Perennial herbs, usually tufted; stems solid, often tall, erect; inflorescences lateral and terminal, racemose or spike-like (densely
clustered racemes in ours); spikelets paired, 1 sessile and bisexual. the other pedicellate and sterile (ours), reduced to a glume or
often just a pedicel (fig. 557h), or pedicel bearing staminate floret, pedicel long-villous.
_-- lem ma
1. A. glomerutus (Walt.) BSP. Fig. 557 s. to Fla .. Tex .. and Mex.; W.I. and C.Am. A variable species.
Peaty. boggy. or wet ground. swamps. and marshes. Se. Mass. Campbell (1983) recognizes several varieties. (A. virginicus var.
and R.I. w. to N.J.. Pa . W.Va .. Ky.. Ark. Okla .. N.M.. and Calif., abbreviatus (Hack.) Fern. & Grise.)
Perennial herbs. tall. reed-like; leaves long. flat; inflorescence a terminal panicle; spikelets I-flowered. paired at each joint. 1 sessile
and 1 pedicellate; most species with spikelets subtended by a ring of silky hairs.
1. Inflorescence strongly contracted. branches stilly ascending (fig. 551a); spikelets not subtended by a ring of hairs ...... 1. E. strictus
1. Inflorescence with loosely ascending to spreading branches (fig. 5 SIc); spikelets subtended by a ring of silky hairs (fig. 551b.d).
2. Basal hairs long. longer than spikelets (fig. 551d). giving panicle a plume-like appearance . . . . . . . . . . . . . . . . .. 2. E. giganteus
2. Basal hairs shorter than spikelets (fig. 551b). panicle not plumose ................................. 3. E. brevibarbis
1. E. strictus Baldw. ex Ell. Narrow Plume Grass Fig. 551 ditches. Coastal plain and lower piedmont. N. Y. s. to Fla .. w. to
Swales. marshes. swamp margins. ditches. and wet places. e. Tex.; inland n. to Ark.. Tenn . and Ky. (E. saccharoides Michx.)
Coastal plain. Va. s. to Fla . w. to e. Tex.; inland n. to Ark.. Mo .
and Tenn. 3. E. brevibarbis Michx. Fig. 551
Sandy or peaty swales. swamp margins. ditches. and wet open-
2. E. giganteus (Walt.) Muhl. Sugarcane Plume Grass ings. Coastal plain. Del. s. to n. Fla. w. to La. and Ark. (E. coarc-
Fig. 551 tatus Fern.; E. coarctatus var. elIiottianus Fern.)
Swales. marshes. margins of swamps. wet lowlands. and
Perennial herbs; stems thick and erect. or weak. slender. and usually matted; spikelets I-flowered. in 2 rows on one side of rachis.
forming I-sided spikes.
1. Principal leaves strongly involute. 0.5-2 mm wide; stems 1-6 mm in diameter at base. typically weak and matted down ... 1. S. patens
1. Principal leaves flat. 5-20 mm wide. becoming involute at apex; stems 5-25 mm in diameter at base. erect.
2. Leaf margins smooth; keels of glumes smooth ................................................ 2. S. alterniflora
2. Leaf margins scabrous (often involute); keels of glumes with short. stiff spine-like hairs (fig. 555b).
3. First glume nearly as long as floret; second glume with conspicuous awn. 5-10 mm long; leaves 4-10(15) mm wide
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. S. pectinata
3. First glume about half the length of floret; second glume sharp-pointed. but not awned; leaves 10-25 mm wide . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 4. S. cynosuroides
1. S. patens (Ait.) Muhl. Saltmeadow Grass. Saltmeadow gravels of shores and rivers. Nfld. w. to James Bay. s. Man .. and
Cord Grass Fig. 552 sw. N.W.T.. s. to w. N.C . Ark.. Tex . N.M . and Oreg. This is the
Salt marshes. saline flats. low dunes. and tidal shores; often common inland species of Spartina. (S. pectinata var. suttiei
dominant in upper salt marshes. Coastal. sw. Nfld. and e. Que. (Farw.) Fern.; S. michauxiana Hitchc.)
s. to Va. and Fla .. w. to e. Tex. A purported hybrid. S. Xcaespitosa
A. A. Eat. (S. patens X S. pectinata). forming tufts is occasionally 4. S. cynosuroides (L.) Roth Big Cord Grass. Salt Reed Grass
encountered; its taxonomic status remains unclear (McDonnell Fig. 554
and Crow. 1979). (S. patens var. monogyna (M. A. Curtis) Fern.; Salt marshes. tidal waters. and freshwater coastal marshes.
S. juncea (Michx.) Willd.; S. patens var. juncea (Michx.) Hitchc.) Coastal, Mass. s. to Fla.. w. to e. Tex. (S. cynosuroides var. polysta-
chya (Michx.) Beal; S. polystachya (Michx.) Beauv.)
2. S. aIternifloru Loisel. Saltwater Cord Grass. Smooth Cord
Grass Fig. 553 S. gracilis Trin. (Alkali Cord Grass) was discovered in Minnesota
Salt marshes and tidal creeks. Coastal. Nfld. and e. Que. s. to in 1980 (Coffin and pfannmuller. 1988). In western North
Fla . w. to e. Tex. America the species frequently occurs at margins of alkali lakes.
small streams. and river bottoms. but in Minnesota it occurs in
3. S. pectinata Link Saltwater Cord Grass. Prairie Cord Grass prairie sites.
Fig. 555
Freshwater marshes. margins of salt marshes. wet prairies. and
a
k).
of pl an t (Hitchcoc
: a. upper po rti on
Pig. 55 1. Er ian thu s strictus let s (Hitchcock).
is: b. pa ir of sp ike
Er ian thu s brevibarb sp ike let (Hitchcock).
s: c. habit; d.
Er ian thu s giganteu
27 4
c
275
a b
Fig. 554. Spartina cynosuroides: a. basal portion of plant; b. lower midsection of stem; c. upper section of stem;
d. panicle; e. ligule; f. spikelet (G&W).
277
r/~
~J.'i\
. !
.
".
a
c
Fig. 556. Leersia oryzoides: a. upper portion of plant (Hitchcock); b. base of plant (Hitchcock); c. growth form
with intlorescence nearly hidden by sheathing leaf (F).
Leersia lenticularis: d. portion of intlorescence and leaf (Hitchcock); e. base of plant (F).
279
a b
Annuals. stout. tufted; stems erect; leaf blades flat; spikelets in 2 rows along one side of rachis. forming I-sided spikes.
1. B. syzigachne (Steud.) Fern. Slough Grass Fig. 555 Calif. Our plants are referable to ssp. syzigachne. (B. erudformis
Sloughs. marshes. ditches. and wet ground. W. Que. w. to Man . of Amer. authors; B. syzigachne var. uniflora (Scribn. ex Gray)
sw. N.W.T. and Alask .. s. to N.Y. n. Ohio. n. Ill.. Kan . N.M . and Bovin)
Perennial herbs. arising from rhizomes; stems somewhat weak. often bearing flat leaves with cutting. scabrous margins; inflorescence
a panicle; spikelets I-flowered. bisexual or sterile. with bristly ciliate to hispidulous margins. overlapping in a single row.
1. Spikelets broadly oval to subcircular. 3-4 mm wide. closely overlapping (fig. 55 6d); principal leaf blades usually 10-15 mm
wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. L. lenticularis
1. Spikelets oblong. 1-2 mm wide. loosely overlapping (fig. 557a.b); principal leaf blades 2-10 mm wide.
2. Panicle open. branches spreading and typically lacking spikelets on lower portion (fig. 556a).
3. Lower branches of panicle whorled or appearing whorled; leaf sheaths conspicuously retrorse-scabrous; blade margins
spinulose; rhizomes slender ........................................................... 2. L. oryzoides
3. Lower branches of panicle alternate; leaf sheaths glabrous or weakly scabrous; blade margins weakly scabrous; rhizomes
thick . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3. L. virginica
2. Panicle narrow. branches strongly ascending and bearing spikelets to base (fig. 557a) ...................... 4. L. hexandra
1. L. lenticularis Michx. Catchfly Grass Fig. 556 often bear cleistogamous inflorescences hidden in the leaf
Swamps. marshes. low woods. and wet ditches. Coastal plain. sheaths (fig. 556c).
Md. s. to Fla . w. to e. Tex. and se. Okla.; major river drainages
into Mississippi R. Valley n. to Ind . Ill.. w. Wisc .. se. Minn .. Mo .. 3. L. virginica Willd. White Grass Fig. 557
and e. Neb. Moist to wet or marshy stream banks. swales. swamps. and low
woods. N.B .. c. Me.. and s. Que. w. to s. Ont.. s. Minn. and se.
2. L. oryzoides (L.) Sw. Rice Cutgrass Fig. 556 N.D . s. to n. Fla. and e. Tex. (L. virginica var. ovata (Poir.) Fern.)
Swamps. marshes. shores. and wet places. Que. and Me. w. to
B.C . s. to Fla . Tex . N.M .. Ariz .. and Calif. Our taxon is var. ory- 4. L. hexandra Sw. Fig. 557
zoides. This species often forms zones around ponds and may Shallow water. ditches. and wet places. Coastal plain. Va. s. to
be so abundant as to make walking difficult (the rough leaves Fla. w. to e. Tex.; Trop. Am.
snagging clothes and scraping skin). In late summer the plants
Perennial herbs. arising from rhizomes (ours) or tufted; stems erect. with short blades; inflorescence a panicle; spikelets bronze-
colored. with 1 terminal bisexual floret and 2 staminate florets; glumes as long as and enclosing florets; lemmas hispidulous. espe-
cially along margins.
Perennial herbs. arising from rhizomes (ours). or annuals; stems erect. with flat leaves; inflorescence a dense. spike-like panicle;
spikelets with 1 fertile floret and 2 sterile lemmas; lemmas with 2 tufts of silky hairs at base.
Fig. 558. Phalaris arundinacea: a. habit; b. ligule; c. spikelet; d. florets; e. grain (Reed).
282
Poaceae (Gramineae) / Grass Family 283
1. P. arundinacea 1. Reed Canary Grass Fig. 558 forming large colonies. Although native. some populations rep-
Meadows, swales, shores, and wet ground. Nfld. w. to Man., sw. resent introductions from Eurasia. A variegated form with white
N.W.T., and Alask., s. to Va., w. N.C., Ky., Ill. , Mo., Okla., N.M., striped leaves is sometimes cultivated and known as 'Ribbon
Ariz., and ne. Calif. This is a common grass of wetlands, often Grass' or 'Gardener's-Garters.'
1. Awn inserted near middle oflemma, straight, barely (if at all) longer than glumes (fig. 557c) ................... 1. A. aequalis
1. Awn inserted near base oflemma. geniculate, 2-4 mm longer than glumes (fig. 557d,e).
2. Plants perennial. stems often decumbent. rooting at nodes; spikelets 2.5(3) mm long .................... 2. A. geniculatus
2. Plants annual. tufted. stems erect; spikelets 2-2.5 mm long ...................................... 3. A. carolinianus
1. A. aequalis Sobo!. Short-awn Foxtail Fig. 557 B.C., s. to N.E., Va .. Pa., Wisc., Minn .. sporadic s. to Okla .. Wyo ..
Shallow water, shores. ditches, and wet ground. Green!' and Ariz., and Calif. Our plants mayor may not represent introduc-
Lab. w. to Man . w. N.W.T. and Alask . s. to N.E.. Md .. Ohio, sw. tions from Eurasia. (A. pallescens Piper & Beetle)
Ky., Ill., Mo., N.M., Ariz., and Calif. A growth form with lax.
floating or creeping stems has been called var. natans (Wah- 3. A. carolinianus Walt. Annual Foxtail Fig. 557
lenb.) Fern. This is our most widespread species of the genus. (A. Shores, ditches, low ground, and waste places. 1.r., N.Y., and
aristulatus Michx.; A. geniculatus var. aristulatus (Michx.) Torr.) N.J. w. to s. Mich. and N.D., s. to Fla. and Tex.; sporadic. Sask.
w. to s. B.C . s. to N.M . Ariz . and Calif.
2. A. geniculatus 1. Water Foxtail Fig. 557
Shallow water and wet places. Se. Lab. and Nfld. w. to Sask. and
1. C. pickeringii Gray Fig. 559 and wet tundra meadows. Green!' and Lab. w. to Alask.. s. to
Sphagnum bogs. wet meadows. wet alpine peats. and margins N.S . n. Me.. n. N.H .. Mich .. Wisc .. Minn .. N.D. mts. to Colo..
of mountain streams and ponds. Nfld.. N.S.. and mts. of Me . Utah. and Calif. (C. neglecta var. stricta (Timm) Griseb.; C. neglecta
N.H .. and n. N.Y.; ne. Mass .. 1.r.. N.Y., and N.J. (C. pickeringii var. var. micrantha (Kearney) Stebbins)
debilis (Kearney) Fern. & Wieg.)
2b. C. stricta ssp. inexpansa (Gray) C. W. Greene Fig. 559
Wet meadows, marshes, and shores of lakes and streams. Lab.
2. C. stricta (Timm) Koe!.
and Nfld. w. to Hudson Bay and Alask.. s. to N.E.. N.Y.. Ont..
2a. C. stricta ssp. stricta Ind . n. Iowa. Neb.. Colo.. and Calif. Greene (1984) recognizes
Peaty and sandy soils. marshes. shores of lakes and streams. two apomictic variants within ssp. inexpansa for North America.
d
Fig. 559. Calamagrostis canadensis: a. habit. with glumes and floret (Hitchcock).
Calamagrostis cinnoides: b. panicle with glumes and floret (Hitchcock).
Calamagrostis stricta ssp. inexpansa: c. panicle with glumes and floret (Hitchcock).
Calamagrostis pickeringii: d. panicle with glumes and floret (Hitchcock).
284
Poaceae (Gramineae) / Grass Family 285
but does not regard them as meriting nomenclatural recogni- America, all of which occur in wet habitats. In the southern and
tion. One, which has been known as C. inexpansa, has straight inland portions of the range var. canadensis is more common;
or weakly twisted awns; the other, previously known as C. lac- var. macouniana (Vasey) Stebbins represents a smaller extreme,
ustris, has geniculate, strongly twisted awns. (C. inexpansa Gray; with less densely flowered panicles and smaller spikelets, and is
C. stricta var. brevior Vasey; C. inexpansa var. brevior (Vasey) Steb- best represented on the Great Plains; var. langsdorjii (Link) In-
bins; C. stricta var. robusta Vasey; C. inexpansa var. robusta (Va- man is a far-northern taxon and tends to occur. at high eleva-
sey) Stebbins; C. lacustris (Kearney) Nash; C. femaldii Louis- tions in our range. (C. canadensis var. robusta Vasey; C. canadensis
Marie) var. scabra (Presl) Hitchc.; C. nubila Louis-Marie)
3. C. canadensis (Michx.) Beauv. Biuejoint Fig. 559 4. C. cinnoides (Muhl.) Bart. Fig. 559
Wet meadows, open swamps, thickets, bog margins, and wet Wet peaty or sandy soils, open swamps, and wet woods. Chiefly
montane soils. Green!' and Lab. w. to Alask., s. to N.E., N.J., mts. coastal states; N.S., s. Me., and s. N.H. sw. to s. N.Y., Pa., and
to N.C., n. Ohio, n. Ind., sw. Ky., n. Mo., Neb., mts. to Colo. and Ohio, s. to Va., N.C., and Tenn.; Ga.
Calif. Greene (1980) recognizes three varieties in eastern North
Perennial (ours) or annual herbs; stems slender (ours), often decumbent; inflorescence an open panicle with slender branches;
spikelets chiefly I-flowered, glumes acute; lemma membranous, acute.
1. M. uniflora (Muhl.) Fern. Fig. 560 Muhlenbergia capiIIaris, M. expansa, M. frondosa, M. glomerata, M.
Bogs, swales, sandy shores, and damp ground. Nfld. and Que. schreberi, and M. sylvatica are occasionally found in wet sites
w. to Ont., s. to N.E., N.J., N.Y., n. Mich., and ne. Minn. This is a and may occur in alluvial soils.
small delicate plant that could be mistaken at first glance for a
species of Panicum or Agrostis.
286
Fig. 561. Paspalum fluitans: a. habit; b. ligule; c. ligule, side view at junction of blade and sheath; d. portion of
raceme; e, f. spikelet, two views (G&W, as P. repens).
287
288 ANGIOSPERMS
1. P. dissectum (L.) 1. Mudbank Paspalum Fig. 562 Mass. w. to s. Ohio. s. Mo .. and Kans .. s. to c. Fla. and se. Tex.
Shallow water and wet places. marshy shores. frequently abun- (P. laeve var. circulare (Nash) Fern.; P. laeve var. pilOS1ll1l
dant after water receeds. S. N.J. and e. Md. w. to Tenn .. sw. Ky.. Scribn.)
s. IlL. and s. Mo . s. to c. Fla. and e. Tex.
5. P. pubiflorum Rupr. ex Fourn. Hairy Paspalum Fig. 563
2. P. fluitans (Ell.) Kunth Floating Paspalum Fig. 561 Open low. wet places. stream banks. margins of ponds and lakes.
Ponds. lakes. swamps. quiet waters of streams. and muddy wet meadows. Va. w. to s. Ohio. s. Ind .. s. Ill.. s. Mo .. and se.
stream banks. Se. Va. s. to Fla . w. to se. Tex.; inland n. to Ky.. Kans . s. to Fla. and e. Tex. Plants with glabrous spikelets have
Ind.. IlL. Mo .. and e. Okla.; Trop. Am. This is our most aquatic been called var. glabrum Vasey ex Scribn. and are more likely to
species of the genus. the plants often occuring submersed. with be encountered in our range. (P. gemillum Nash; P. laevigllllllis
elongate. spongy stems and developing numerous dark. feath- Scribn.)
ery roots at nodes. Pohl (1980) regards this species as distinct
from P. repens Berg. which is strictly tropical. 6. P. praecox Walt. Early Paspalum Fig. 502
Swamps. wet pine savannas. and flatwoods. Coastal plain. se.
3. P. boscianulII Fliigge Fig. 563 Va. s. to c. Fla .. w. to se. Tex.: inland in w. Tenn. and w. Ky. Our
Shores. wet sands. peaty or mucky sites. and low woods. Va. w. plants are referable to var. curtisianuIII (Steud.) Vasey.
to Tenn. and Ark .. s. to s. Fla. and e. Tex.; Trop. Am.
PaspalulII bijiduIII. P. distichuIII. P. floridanum. and P. urvillei may
4. P. laeve Michx. Fig. 562 sometimes occur in wet sites in the southeastern portion of
Wet sands. sandy peats or mucks. shores. thickets. and savannas. our range.
1. Plants perennial. with creeping. knotty rhizomes; stems slender. 0.4-1 m tall; spikelets subtended by involucre of 8-12 bristles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. S. geniculata
1. Plants annual. lacking rhizomes; stems thick (up to 2 cm in diameter). 3-4 m tall; spikelets subtended by involucre of 1-3
bristles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. S. maglla
1. S. geniculata (Lam.) Beauv. Knotroot Bristlegrass. 2. S. magna Griseb. Giant Foxtail. Giant Bristlegrass
Perennial Foxtail Fig. 563 Fig. 563
Freshwater or saline marshes. shores. swales. and seasonally Swamps. marshes. fresh and saline swales. shores. and bayous.
wet sites. Se. N.B. w. to Pa. WVa . s. Ill.. and Kans .. s. to Fla . Coastal plain. N.J. s. to Fla .. w. to Tex.: inland n. to Ark.: WI.
Tex . N.M . Ariz . and Calif.: WI. and S.Am. According to Ker- andC.Am.
guelen and colleagues (1987). our taxon may be conspecific
with S. parvifIora (Poir.) Kerguelen. in which this latter name
would have nomenclatural priority.
27. Sacciolepis
Perennial (ours) or annual herbs. tall; stems often decumbent: leaves flat. broad: inflorescence a contracted panicle: spikelets gibbous
because of round swelling at base of second glume. 2-flowered. lower floret sterile or staminate. upper floret bisexual.
Fig. 562. Paspalum Iaeve: a. habit; b. spikelet, two views; c. upper floret (Hitchcock).
Paspalum dissectum: d. inflorescence; e. spikelet, two views; f. upper floret (Hitchcock).
Paspalum praecox: g. inflorescence; h. spikelet, two views; i. upper floret (Hitchcock).
289
d
h
g
Fig. 563. Paspalum pUbiflorum: a. inflorescence; b. spikelet. two views; c. upper floret (Hitchcock).
Paspalum boscianum: d. inflorescence; e. spikelet. two views; f. upper floret (Hitchcock).
Setaria geniculata: g. habit (Hitchcock).
Setaria magna: h. portion of inflorescence with leaf (Hitchcock).
290
J
I
e
b
Fig. 564. Sacciolepis striata: a. habit; b. spikelet. two views; c. upper floret (Hitchcock).
Echinochloa walteri: d. inflorescence; e. leaf sheath; f. spikelet (Beal).
291
292 ANGIOSPERMS
Perennial or annual herbs; leaves flat; inflorescence a panicle with spike-like lateral branches; spikelets spiny-hispid, with 1 fertile
floret.
REFERENCES: Fassett, 1949; Gould, Ali, and Fairbrothers, 1972; Wiegand, 1921.
1. Leaf sheaths with bristly hairs (fig. 564e); spikelets elliptic, about 3 times as long as wide; second (larger) glume awned ....
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. E. walteri
1. Leaf sheaths glabrous; spikelets ovate, about twice as long as wide; second (larger) glume lacking awn.
2. Lateral inflorescence branches simple (fig. 565d), 1-2 cm long; spikelets crowded into about 4 rows on one side of rachis;
panicle pale green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. B. colona
2. Lateral inflorescence branches compound (fig. 565a), more than 2 cm long; spikelets crowded and irregularly clustered;
panicle dark g'reen to purple.
3. Smooth, lustrous portion of upper (fertile) lemma sharply differentiated from dark, dull, wrinkled tip, with a ring of
minute setae present just below junction of body and tip; second glume and lower (sterile) lemma usually lacking stiff
papillate-based hairs .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3. B. crusgalli
3. Smooth, lustrous portion of upper (fertile) lemma gradually blending to the dull, wrinkled tip, lacking a ring of setae on
upper portion of lemma at junction of body and tip; second glume and lower (sterile) lemma covered with stiff papillate-
based hairs ...................................................................... 4. B. muricata
1. B. walteri (Pursh) Heller Fig. 564 and disturbed sites; weedy. Widely natzd. throughout the U.S.;
Alkaline and saline marshes, swamps, and shallow water of intro. from Old World.
ponds and ditches. Que. and Ont. w. to s. Mich., Wisc., and Iowa,
s. to Fla., e. Tex., and e. Mex.; Cuba. The spikes of this species 4. B. muricata (Beam>;) Fern.
often have a more bristly aspect than our other species. Low ground, ditches, and margins of swales and ponds. N.B.
and Que. w. to Mich., Minn., and Alta., s. to Fla., Tex., N.M.,
2. E. colona (L.) Link Jungle Rice Fig. 565 and Calif. This species is quite variable and several varieties have
Moist to wet open places, shores, sand and gravel bars, ditches, been described; these are not differentiated here. Voss (1972)
cultivated fields, and disturbed sites; weedy. Va. w. to Mo., s. to has pointed out that the name B. pungens (Poir.) Rydb., ap-
Fla., Tex., N.M., 'Ariz., and s. Calif.; sporadic elsewhere; intro. pearing in many manuals for this species, is incorrect.
from Old World.
Perennial and annual herbs; inflorescence typically an open panicle, sometimes contracted; spike lets 2-flowered, lower floret sterile,
consisting of a lemma only, upper floret bisexual, with lemma and palea hard, cartilaginous, enclosing the grain.
The first three species are treated by some authors in the segregate genus Dichanthelium (Hitchc. & Chase) Gould.
REFERENCES: Gould and Clark, 1978; Lelong, 1984; Stephenson, 1984; Webster, 1988.
1. Basal leaves distinctly different from cauline leaves, forming a winter rosette.
2. Ligular hairs conspicuous, 3-5 mm long ................................................... 1. P. acuminatum
2. Ligular hairs obscure, less than 1 mm long, or lacking.
3. Spikelets pubescent, elliptic, obtuse, 2-2.2 mm long. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. P. boreale
3. Spikelets glabrous or obscurely puberulent, ovate, acute, 2.3-2.6 mm long ........................ 3. P. scabriusculum
1. Basal leaves similar to cauline leaves, not forming a winter rosette.
4. Spikelets with wart-like tubercles on second glume and lower (sterile) lemma (fig. 566h) .................. 4. P. verrucosum
4. Spikelets lacking tubercules.
5. PlantsperenniaJ, rhizomatous or densely tufted, with dead leaves from previous year present.
6. Panicle contracted, spike-like (fig. 568a); ligule membranous ................................. 5. P. hemitomon
6. Panicle open; ligule a fringe of cilia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6. P. longifolium
Fig. 565. Echinochloa crusgaIli: a. habit; b. spikelet. two views; c. upper floret (Hitchcock).
Echinochloa colona: d. inflorescence and portion of plant (Hitchcock).
293
a
0 1
I
I
.'1
:'
spiKelets
0.: ...
i
h
Fig. 566. Panicum philadelphicum: a. spikelet, two views, top, and upper floret, bottom (Hitchcock).
PanicumjIexile: b. inflorescence (B&B); c. spikelet, two views, left, and upper floret, right (Hitchcock).
Panicum acuminatum: d. inflorescence; e. spikelet, two views; f. upper floret (Hitchcock).
Panicum verrucosum: g. habit (B&B); h. spikelet, two views (Hitchcock); i. upper floret (Hitchcock).
294
Poaceae (Gramineae) / Grass Family 295
If)
~
a
(()
c d
Fig. 567. Panicum boreale: a. spikelet, two views, left, and upper floret, right (Hitchcock).
Panicum longifolium: b. spikelet, two views, left, and upper floret, right (Hitchcock).
Panicum scabriusculum: c. spikelet, two views, left, and upper floret, right (Hitchcock).
Panicum implicatum: d. spikelet, two views, left, and upper floret, right (Hitchcock).
5. Plants annual, lacking rhizomes, stems solitary or branched at base, lacking dead leaves from previous year.
7. Spikelets with long, tapering tips (fig. 566c), 3-3.5 mm long; internodes of stem pilose ................... 7. P. flexile
7. Spikelets with short-pointed tips (fig. 566a), 1.7-2 mm long; internodes of stem glabrous (except sometimes
papillose-hispid below nodes) .................................................... 8. P. philadelphicum
Perennial herbs. arising from rhizomes; stems and leaves erect. floating. or submersed; flowers unisexual. in globose heads. staminate
heads borne above pistillate heads; staminate heads falling off early; pistillate heads appearing burr-like; perianth parts of pistillate
flowers 3-6. scale-like; fruit a drupe with 1 or 2 locules. obovoid to fusiform. frequently with a slight median constriction.
Sparganium is a difficult group. and mature fruit is necessary for the identification of most species. with the exception of S. eurycar-
pum. which is our only species with 2 stigmas.
REFERENCES: Beal. 1960a; Cook. 1985; Cook and Nicholls. 1986. 1987; Crow and Hellquist. 1981; Fernald. 1922; Harms. 1973;
Lakela. 1941; Reveal. 1970; Voss. 1966.
Fig. 568. Panicum hemitomon: a. habit; b. ligule; c, d. spikelet, two views (G&W).
297
298 ANGIOSPERMS
l. S. eurycarpum Englem. Large-fruited Bur-reed Fig. 569 In most references this taxon is treated as S. chlorocarpum.
Shallow water. damp shores. swales. and marshes. Nfld. and which Cook and Nicholls (1986) regard as conspecific with S.
Que. w. to n. Alta. and s. B.C . s. to Va .. n. Ky.. Mo .. Okla .. N.M .. emersum. The most common phase of the species in the East is
Ariz .. and Calif. referable to ssp. acaule Cook and Nicholls. which is distinguished
by congested pistillate heads. somewhat smaller fruits. and
2. S. glomeratum (Beur!. ex Laestad.) Neum. lower bracts conspicuously longer than the inflorescence. How-
Bogs and shallow water. Reported from s. Saguenay Co . Que .. ever. we have noted that such plants frequently occur in mixed
ne. Minn .. Alta .. Yuk.. and B.C.; Cook and Nicholls (1986) con- populations with plants referable to ssp. emersum (Crow and
firm it only from Labrador. Hellquist. 1981). Hence we are inclined to agree with Voss
(1966) in recognizing these plants as representing two growth
3. S. natans L. Northern Bur-reed Fig. 5 70 forms. (S. chlorocarpum Rydb.; S. chlorocarpum var. acaule
Lakes. ponds. streams. and ditches. Ntld. w. to Alask.. s. to n. (Beeby) Fern.)
N.J .. n. Pa .. Mich .. Wisc .. Utah. and Calif. Although this taxon
is typically referred to as S. minimum. Cook (1985) has deter- 7. S. fluctuans (Engelm. ex Morong) Robins. Fig. 573
mined S. natans to be the correct name. (S. minimum (Hartm.) Lakes and ponds oflow alkalinity. Nfld. w. to s. Que. and n. Alta ..
Fries) and B.C . s. to N.E.. n. Pa .. n. Mich .. and Minn. Vegetative plants
may be confused with S. angustifolium; the leaf width in S. fluc-
4. S. hyperboreum Beur!. ex Laestad. Fig. 575 tuans ranges from 3-11 mm. but is generally wider than 5 mm.
Cold ponds. streams. and ditches. Arctic regions s. to Nfld .. n. whereas that of S. angustifolium ranges from 1.5-5 mm. but is
N.S .. Que .. Man .. and B.C. typically less than 5 mm wide.
pistillate ---w~'\
head
c
a
Fig. 569. Sparganium eurycarpum: a. upper portion of plant; b. flowering head; c. fruiting head; d. fruit
(NHAES).
299
sta mi nat e hea d
it
c. fruiting head; d. fru
sub me rse d pla nt; b. habit. emersed plant;
habit.
Fig. 570. Sparga
(NH S). IIl natans: a.
AEniU
30 0
supra-axillary
bract
c
b
Fig. 571. Sparganium angusti!olium: a. habit; b. upper portion of plant with fruiting heads; c. fruit (NHAES).
301
c
a
.
. b h
sparganlU b .t ",0 (0
Fig. 5 7 rn ernersu '" " " c.
(,ul"'"
2. rn. a. .
~1 Maa, a
. ( " , , , lN
l'Al'S)'
302
Fig. 573. Sparganiumj1uctuans: a. habit; b. upper portion of plant; c. fruit (NHAES).
303
Fig. 574. Sparganium androcladum: a. habit; b. upper portion of plant; c. fruiting head; d. fruit (NHAES).
304
c
Perennial herbs, ca. 1-3 m tall, arising from creeping rhizomes, colonial; leaves long and narrow, erect and twisted, sheathing at
base; flowers unisexual, borne in a dense cylindric terminal spike-like inflorescence, pistillate portion below staminate portion; fruit
a minute thin-walled achene, raised on a slender stalk bearing numerous long hairs, style persistent.
Cattails may appear in almost any wet place, often being the first invaders in excavated ponds or ditches. The underground stems
spread extensively, so that a large stand of cattails may actually consist of but a few individual plants. A good field character for
recognizing Typha in the vegetative state is the distinctive twisting of the leaves.
REFERENCES: Crow and Hellquist, 1981; Hotchkiss and Dozier, 1949; Lee, 1975; Smith, 1967, 1986, 1987; Yeo, 1964.
1. Staminate and pistillate parts of inflorescence usually contiguous (fig. 576a,b) (occasionally separated by a gap of up to 5 mm,
or even up to 8 mm in some clones), mature fruiting spike at least 2.5 cm in diameter; pistillate bractlets absent; stigma
lanceolate to ovate-Ianceo late, flattened, persistent in fruit; shoots and leaves relatively broad . . . . . . . . . . . . . . . . . . . . 1. T. latifolia
1. Staminate and pistillate parts of inflorescence usually separated by a gap of 0.5-8 cm (figs. 577a,c, 578a,b), mature fruiting
spike less than 2.5 cm in diameter; pistillate bractlets present, numerous (fig. 577r) (minute in T. Xglauca); stigmas slender,
elongate, often deciduous in fruit; shoots and leaves relatively narrow.
2. Leaf sheaths with auricles (fig. 577d) (usually diSintegrating with age); fruiting spike dark brown; pistillate bractlet tips
blunt and dark brown, or narrower than stigmas and very pale.
3. Dried leaf blades 3-8 mm wide; spike gap about 1-8 cm; pistillate bractlet tips conspicuous (fig. 577s,t), blunt, dark
brown; stigmas capillary, usually deciduous in fruit .. , ... , .... , . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. T. angustifolia
3. Dried leaf blades about 6-15 mID wide; spike gap about 0.5-4 cm; pistillate bractlet tips minute (require high magnifica-
tion), pale; stigmas linear-lanceolate, partly persistent " ... " .. ,............................... 3. T. Xglauca
2. Leaf sheaths tapered to blade, lacking auricles; fruiting spike bright cinnamon-brown or orange-brown; pistillate bractlet
tips usually pointed, usually paler and wider than stigmas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. T. domingensis
1. T. latifolia L. Common Cattail Fig. 576 species. This taxon is a hybrid between T. latifolia and T. angusti-
Damp shores, marshes, swamps, marshy river margins, and folia, and although plants usually form few seeds, they may
roadside ditches; often forming extensive stands. Nfld. w. to persist for many years, spreading widely by rhizomes and some-
Alask., s. to Fla., Tex., Calif., and Mex. Plants with a small sepa- times becoming ecologically very important. Plants intermedi-
ration of the staminate and pistillate portions of the spikes have ate between T. Xglauca and the parents, especially T. angustifolia,
been described as Typha latifolia f. ambigua (Sonder) Kronf. and are locally common. Hybrids of T. domingensis X T. latifolia are
may be confused with T. angustifolia or T. Xglauca. very similar and might also be expected on the southeastern
and southwestern edges of our range. (T. angustifolia var. elon-
2. T. angustifolia L. Narrowleaf Cattail Fig. 577 gata (Dud!.) Wieg.)
Coastal salt marshes and inland in nutrient-rich waters. P.E.I.
and N.S. w. to s. Que., Ont., s. Sask., s. Mont., Wyo., and se. 4. Typha domingensis Pers. Southern Cattail Fig. 578
Oreg., s. to Fla., Ga., Mo., s. Tex., Calif., and Mex. Coastal marshes, swamps, and pools, and brackish and saline
sites. Coastal plain, De!. and e. Md. s. to Fla., w. to La. and Tex.;
3. T. Xglauca Godr. Hybrid Cattail inland to s. Kan., Okla., Utah, N.M., Nev., s. Oreg., Calif., and
Occasional, often locally abundant, usually with the parental Mex.; C.Am. and S.Am.
e
pistillate
spike
c d m n
Fig. 576. Typha lati/alia: a. upper portion of plant with contiguous staminate (above) and pistillate (below)
spike; b. older spike; c, d. pistillate spike variations; e. portion of pistillate spike axis. showing com-
pound pedicels; f. single compound pedicel; g. young pistillate flowers with hair-like bractlets;
h. i. anthers on branched filaments; j. pollen tetrad variations; k. staminate bractIets. commonly
white-tipped; I. stigma; m. sterile pistillate flower with ellipsoidal aborted ovary; n. aborted ovary;
o. fertile pistillate flower (Mason).
307
a
-;-';'0'
~*':':
,,-.
r
' 'f
b
o
q
s u
Fig. 577. Typha angustifolia: a-c. upper portion of plant with separate staminate (above) and pistillate (below)
spikes; d. auricled leaf sheath; e. portion of pistillate spike axis, showing compound pedicels; f. single
compound pedicel; g. cluster of young anthers surrounded by bractlets, filament not yet elongated;
h-j. mature stamens; k-m. staminate bractlets; n. pollen grains; o. sterile pistillate flower with
aborted ovary; p. aborted ovary; q. cluster of bractlets, some with enlarged tips; r. swollen tip of
pistillate bractlet; s. pistillate bractlets; t, u. fertile pistillate flowers (Mason).
308
,,
!
..1
. ~:.
Fig. 578. Typha domingensis: a. b. inflorescence. staminate above. pistillate below; c. pistillate spike; d. portion
of pistillate spike axis. showing compound pedicels; e. compound pedicels; f. g. bractlets of staminate
flowers; h. pollen grains; i. sterile pistillate flower with aborted ovary; j. pistillate bractlet with swol-
len tip; k. l. aborted ovary; m. fertile pistillate flower (Mason).
309
Subclass Zingiberidae
Marantaceae / Arrowroot Family
1. Thalia (Thalia)
Perennial herbs, large, erect, arising from thick rhizomes; leaves large, alternate, basal, long-petioled; inflorescence a narrow to
spreading panicle; flowers bisexual, in pairs, subtended by 2 bracts; sepals 3, small; petals 3, llanceolate, 2 obovate, purple; ovary
inferior; fruit a fleshy bluish-purple utricle.
310
Fig. 579. Thalia dealbata: a. habit; b. individual bract with drying apex; c. portion of flower showing outer
staminode, stamen, calloused staminode, and hooded staminode; d. pistil and stigmatic surface;
e. ovary with bracts at summit; f. outer staminode; g. stamen separated from flower; h. hooded
staminode (C&C).
311
Subclass Liliidae
Pontederiaceae / Pickerel-weed Family
1. Plants free-floating; leaves with blades sub circular to broadly elliptic. base widely cuncatc. petioles usually inflated (fig. 580a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. EichllOrnia
1. Plants rooted. emersed or submersed; leaves with blades ovate to lanceolate (figs. 581. 582) or reniform (fig. 584c). base
cordate (fig. 582c) to truncate (fig. 582e.f). petioles not inflated. or leaves linear and ribbon-like (fig. 583a.b).
2. Stamens 6; flowers 2-lipped. in a dense spike-like inflorescence. blue-violet; leaves cordate. hastate. or narrowly to broadly
lanceolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Pontederia
2. Stamens 3; flowers regular. solitary. yellow. blue. or rarely white; leaves reniform. lanceolate. or linear and ribbon-like.
3. Leaves lax. long-linear. ribbon-like. sessile (fig. 583a.b). usually submersed; flowers yellow .................. 3. Zosterella
3. Leaves erect. reniform to rcniform-cordate. or ovate to elliptic to elliptic-Ianceolate. petiolate. usually emersed; flowers
white to purplish-blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Heteranthera
1. Eichhornia (Water-hyacinth)
Perennial and annual herbs; leaves subcircular. ovate to broadly elliptiC. petioles often inflated; inflorescence spike-like; flowers violet-
blue with yellow streaks. subtended by a spathe-like bract; fruit a capsule.
2. Pontederia (Pickerel-weed)
Perennial herbs. arising from creeping rhizomes; leaves basal. erect. long-petioled. usually cordate; flowers violet-blue. in a dense
spike subtended by a spathe-like bract; fruit a utricle. crested with 6 toothed ridges.
1. P. cordata L. Pickerel-weed Figs. 581. 582 and S.Am. The seedlings consist of submersed rosettes of linear
Margins of lakes. ponds. and streams. marshes. swamps. and leaves and often have 1 or 2 petiolate. slightly laneeolate leaves.
ditches. P.E.I. and N.S. w. to s. Que .. Ont.. Mich .. Wisc.. and The range of variability in the leaves of this species is consider-
Minn .. s. to Fla .. Ala .. Miss .. La .. and Tex.; Cuba. Belize. able. (P. cordata var. lancifolia (Muhl.) Torr.; P. lanceo/ata Nutt.)
312
Fig. 580. Eichhornia crassipes: a. habit; b. leaf with inflated petiole; c. flower; d. flower. longitudinal section;
e. ovary, cross-section (Mason).
313
Fig. 581. Pontederia cordata: a. habit; b. flower; c. withering perianth, revolute-coiled, exposing shape of devel-
oping ovary (C&C).
314
,;.,
f~
Fig. 582. Pontederia cordata: a. habit. submersed form; b. habit. submersed juvenile
plant; c-f. leaf variations
(NHAES).
315
Fig. 583. Zosterella dubia: a. habit (NHAES); b. portion of filiform growth form (NHAES); c. flower (NHAES);
d. habit, terrestrial form (F).
316
Haemodoraceae I Bloodwort Family 317
1. Z. dubia (Jacq.) Small Water Star-grass Fig. 583 with the conspicuous midvein in Potamogeton. Plants stranded
Alkaline waters or muddy shores of ponds, lakes, and streams. in the mud or floating in dense mats where the foliage reaches
N.B .. Me., and sw. Que. w. to s. Ont .. Minn., Ida., B.C., and Oreg., the surface tend to produce conspicuous yellow flowers; c1eis-
s. to Fla., Tex., Ariz., Calif., and Mex. This is often confused with togamous flowers often occur on submersed stems. (Heteranth-
a few of the linear-leaved species of Potamogeton, but the pres- era dubia (Jacq.) MacM.)
ence of an obscure midvein in Zosterella is distinctive in contrast
4. Heteranthera (Mud-plantain)
Perennial or annual herbs; leaves in a basal rosette; flowers blue or white, emerging from sheathing leaf bases; fruit a capsule.
1. H. limosa (Sw.) Willd. Fig. 584 Va., w. N.C., n. Ga., and w. Fla, w. to Tex. and Mex.; WI., C.Am.,
Muddy shores and quiet waters of canals and ponds. Ky. and s. and S.Am.
Ill. w. to sw. Minn., s. S.D., Neb., and Colo., s. to Fla., Okla., Tex.,
N.M., Ariz., and Mex.; C.Am. and S.Am. 3. H. multiflora (Griseb.) Horn
Muddy shores and shallow water. N.J. s. to ne. N.C.; sw. Ill. and
2. II. renijormis R. & P. Fig. 584 Mo. w. to Neb .. s. to sw. Tenn., Miss., and Tex.; S.Am.
Muddy shores and shallow water. S. Conn. and c. N.Y. s. to se.
1. Rhizomes red; outer perianth parts shorter than inner parts (fig. 585d); ovary inferior; stamens 3; seeds brown ..... 1. Lachnanthes
1. Rhizomes white to brown; outer and inner perianth parts of equal length (fig. 585c); ovary half-inferior; stamens 6; seeds
yellow-white ............................................................................ 2. Lophiola
1. Lachnanthes (Red-root)
Perennial herbs, arising from slender rhizomes that exude a red juice; leaves basal and scattered along stem; inflorescence a com-
pound cyme; perianth yellow, tomentose; fruit a globose capsule; seeds brown, flat.
2. Lophiola (Golden-crest)
Perennial herbs, arising from slender to thickened rhizomes; leaves alternate, 2-ranked, linear, basal and cauline; inflorescence a
solitary, terminal, dichotomous cyme; perianth yellow, tomentose; fruit an ovoid, beaked capsule; seeds ellipsoidal, yellow-white.
318
Fig. 585. Lachnanthes caroliniana: a. base of plant; b. upper portion of plant; c. flower; d. flower, longitudirial
section (G&W).
Lophiola americana: e. upper portion of plant; f. flower (Gleason).
319
320 ANGIOSPERMS
1. Stems thick. leafy to top. leaves numerous (fig. 586a); plants 60-200 em tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. Veratrum
1. Stems thin. leaves basal or near base. or if scattered along stem. then 4 or fewer (typically 3); plants 2-60 em tall.
2. Leaves basal or near base; flowers scapous or subscapous.
3. Filaments wooly-pubescent; perianth yellow ................................................ 2. Nartheciwn
3. Filaments glabrous; perianth white. green. or pink.
4. Perianth pink; scape thick. hollow; leaves not 2-ranked ........................................ 3. Helonias
4. Perianth white or green; scape slender. solid; leaves 2-ranked (fig. 587d) ............................ 4. Tofieldia
2. Leaves scattered along stem; flowers not scapous ................................................ 5. Smilacina
2. Narthecium (Bog-asphodel)
Perennial herbs. arising from stout rhizomes; leaves basal. erect. narrowly linear; inflorescence a raceme. bracteate; perianth yellow;
ovary 3-locular; fruit a capsule; seeds fusiform. 2-tailed.
Perennial herbs. arising from short rhizomes; leaves elongate. basal; scape hollow. with reduced bract-like leaves; inflorescence a
spike-like raceme. bracts absent; perianth pink; fruit a 3-lobed loculicidal capsule; seeds 2-tailed.
Perennial herbs. arising from short rhizomes; leaves linear. 2-ranked. basal; inflorescence a terminal spike or raceme; flowers sub-
tended by an involucre of bracts; perianth green or white; ovary superior. 3-locular; fruit a septicidal capsule.
1. Plants glabrous; fruiting raceme 5-8 mm wide; seeds not appendaged ..................................... 1. T. pusilla
1. Plants glandular or pubescent (fig. 587b); fruiting raceme 10-20 mm wide; seeds appendaged.
2. Capsule as long as or slightly shorter than perianth; seed appendages short. not contorted ................... 2. T. racemosa
2. Capsule longer than perianth; seed appendages contorted ......................................... 3. T. giutinosa
1. T. pusilla (Michx.) Pers. Fig. 586 2. T. racemosa (Walt.) BSP. Fig. 587
Wet rocks and calcareous alpine or arctic marshes. Green!.. Wet sand. clay. peat. pine savannas. and seeps. Coastal plain. s.
Lab .. and Nfld. w. to Alask .. s. to Gaspe Pen .. Que .. n. Mich .. N.J. s. to Fla . w. to se. Tex. (Triantha racemosa (Walt.) Sm.)
n. Minn .. nw. Mont.. and s. B.C. (T. paiustris Huds.)
k
r h
Perennial herbs, arising from short, thick rhizomes or slender, creeping rhizomes (ours); leaves alternate, sessile or subsessile, nar-
rowing to a subpetiolate sheath; flowers borne in a panicle or raceme (ours); perianth white, 3-merous (ours) or 2-merous; ovary
superior, 3-locular; fruit a berry.
1. S. trifolia (L.) Desf. False Solomon's-seal Fig. 586 1986b) has presented evidence for treating Smilacina and Mai-
Wet mossy woods, bogs, coniferous swamps, and peaty shores. anthemum as congeneric; the name Maianthemum would then
S. Lab. and Nfld. w. to Man., N.W.T., and Alask., s. to N.E., n. have nomenclatural priority. (Maianthemum trifolium (1.)
N.J., n. Ohio, n. Ill., Minn., Alta., and B.C. La Frankie (1986a, Sloboda)
1. 1. fulva Ker. Red Iris Fig. 588 3. l. brevicaulis Raf. Lamance Iris Fig. 591
Swamps, marshes, wooded swamps, and ditches. Ga. w. to La., Swamps, wet woods, and bottomlands. S. Ont. and Ohio w. to
n. along the Mississippi embayment to se. Mo., sw. Ky., and sw. Ill. and e. Kans., s. to Ala. and e. Tex. (1. foliosa Mackenz. & Bush)
Ill.
4. 1. prismatica Pursh Slender Blue Flag
2. 1. pseudacorus 1. Yellow Iris Fig. 589 Brackish to saline marshes, freshwater marshes, shores, and
Marshes, meadows, streams, and lakeshores. Nfld. and N.S. w. meadows. Coastal, N.S. s. to Del., ne. Md., and Ga.; inland in
to Ont., Minn., and Man., s. to N.C., W.Va., Ohio, n. Ky., and Ill.; mts. of Tenn., N.C., and Ga. (1. prismatica var. austrina Fern.)
se. Tex.; nw. Mont., n. Ida., s. B.C., nw. Wash., and n. Calif.;
intro. from Eurasia.
b
~
:'
~;; .:':r" . ,
h
,".,'
',-
.:
sepal
Fig. 590. Iris virginica var. virginica: a. basal portion of plant (G&W); b. flowering stem (G&W); c. flower (F);
d. sepal (G&W); e. petal (G&W); f. style branches (G&W); g. fruiting stem (G&W); h. capsule, cross-
section (G&W); i. seed (G&W).
Iris versicolor: j. flower (F).
326
a
5. I. versicolor L. Blue Flag Figs. 590, 591 Que. w. to s. Mich., Minn., and e. Kans., s. to e. Va., N.C., Fla.,
Marshes, meadows, stream banks, lake and pond margins, and and e. Tex. Two varieties have been recognized: the southern
ditches. Lab. and Nfld. w. to Man., s. to w. Va., n. Ohio, Mich., coastal var. virginica, characterized by capsules about as wide as
Wisc., and Minn. long, and the more northern var. shrevei (Small) Anderson, with
capsules much longer than wide. The latter taxon has also been
6. I. virginica 1. Southern Blue Flag Figs. 590, 591 treated as a variety of I. versicolor (var. shrevei (Small) Boivin) or
Marshes, ditches, wet shores, swamps, and shallow water. Sw. as a distinct species (I. shrevei Small).
Perennial or annual herbs, small; roots thread-like; leaves alternate, small, bract-like or scale-like; flowers bisexual, solitary, or borne
in capitate, racemose, or cymose inflorescences; perianth 6-merous, borne at summit of a floral tube; ovary inferior; fruit a 3-valved
capsule, seeds minute.
1. Floral tube obtusely 3-angled; flowers greenish-white, cream, or light blue, sessile, few to several in a capitate cluster . .. 1. B. capitata
1. Floral tube 3-winged; flowers violet, on short pedicels borne in 2-branched cymes, or solitary. . . . . . . . . . . . . . . . . . .. 2. B. biflora
The orchids are the largest family of flowering plants and the most advanced family of the Monocots, numbering nearly 20,000
species. Because of the beauty of many of the species, members of the Orchid Family are also vulnerable to collection. Many states
have conservation laws protecting their native species of orchids. The family is easily recognized by the following characteristics:
flowers bilaterally symmetrical; sepals 3; petals 3, lateral 2 similar, third modified into a distinct labellum, typically in a lower position
because of 180-degree twisting of the ovary/pedicel; stamens adnate to style, forming column; anthers 1 (2 in Cypripedium), with
pollen usually densely aggregated into a waxy mass, the pollinium; anther and stigmatic lobes separated by rostellum and its vis-
cidium (sticky pad detaching with pollinium in pollen transfer); ovary inferior; fruit a capsule, with very numerous, tiny, dust-
like seeds.
REFERENCES: Brackley, 1985; Case, 1987; Correll, 1950; Dressler, 1981; Fuller, 1933; Henry, Buker, and Pearth, 1975; Luer, 1975;
Morris and Eames, 1929; Sheviak, 1974; Whiting and Catling, 1986.
1. Labellum of flower pouch-like (fig. 593); leaves plicate; fertile anthers 2 ................................. 1. Cypripedium
1. Labellum of flower various, but not pouch-like; leaves flat, or if ribbed, not plicate; fertile anthers 1.
2. Leaves whorled at stem tip (fig. 594a); sepals extending far beyond petals (fig. 594a) .......................... 2. Isotria
2. Leaves basal, or basal and cauline, never whorled; sepals about the same length as petals.
3. Flowers solitary, pink (rarely albino); leaves solitary (rarely 2), or undeveloped at time of flowering.
4. Leaf linear, basal; labellum 3-lobed, lateral lobes short, middle lobe long, arching downward (fig. 594c) ...... 3. Arethusa
4. Leaf ovate, borne about midway on stem; labellum unlobed, not arching ............................ 4. Pogonia
3. Flowers 2 or more, of various colors, inflorescence a raceme or spikc; leaves 2 or more.
5. Leaves cauline.
6. Leaves several, alternate; flowers spurred (fig. 596c), of various colors .......................... 5. Platanthera
6. Leaves 2, opposite (fig. 601c); flowers not spurred, green ...................................... 6. Listera
5. Leaves basal or appearing basal, or absent at time of flowering.
7. Labellum uppermost, column below it, flowers thus appearing inverted (fig. 602b) .................. 7. Calopogon
7. Labellum lowermost, column above it.
8. Leaves linear to linear-Ianceolate; flowers white to ivory, spiraled in a spike (figs. 603a, 604a) . . . . . .. 8. Spiranthes
e
b
Fig. 592. Burmannia capitata: a. habit; b. flowers with developing capsule; c. flower, spread open; d. style and
stigmas (C&C).
Burmannia biflora: e. habit (C&C); f. flower, with ovary longitudinally sectioned (G&W); g. flower,
spread open (C&C); h. style and stigmas (C&C); i. ovary, cross-section, with thin wings (G&W);
j. seed (G&W).
329
330 ANGIOSPERMS
1. Cypripedium (Lady's-slipper)
Perennial herbs. with coarse, fibrous roots; leaves basal or cauline. plicate. sheathing at base; flowers solitary or few, large. showy;
labellum inflated, pouch-like; fertile stamens 2; fruit a capsule.
1. C. acaule Ait. Pink Lady's-slipper; Moccasin-flower 3. C. candidum Muhl. ex Willd. Small White Lady's-slipper
Pig. 593 Pens, swampy meadows, prairies, and mossy glades. Rare;
Acid soils of dry woodlands. in sphagnum bogs, and wet. mossy c. N.Y. w. to s. Ont., Man., and Sask.. s. to n. N.J .. e. Pa., Md.,
coniferous woods. Nfld. w. to Alta. and N.WT., s. to N.C., w. S.C., Ky.. Mo., and N.D.
n. Ga .. n. Ala., e. Tenn .. n. Ind., n. Ill., Wisc., and Minn.
4. C. reginae Walt. Showy Lady's-slipper Fig. 593
2. C. parvijlorum Salisb. Small Yellow Lady's-slipper Chiefly calcareous coniferous swamps, bogs, wet slopes, and
Fig. 593 shores. Nfld. w. to Ont. and e. Sask.. s. to n. N.J.. mts. to Ga. and
Chiefly calcareous swamps, bogs, mossy woods, and wet shores. Tenn., Ohio, Ill., and Mo.
Nfld. and N.S. w. to Ont., Mich .. Wisc .. and e. Minn., s. to N.J.,
Pa .. mts. to N.C .. S.C .. and Tenn., Ohio, Ind .. and n. Ill. The typi- 5. C. arietinum R. Br. Ram's-head Lady's-slipper Fig. 593
cally larger-flowered C. pubescens Willd., with greenish-yellow Swamps and bogs, especially with Thuja. and dry. rich, wooded
sepals and petals and a larger lower labellum, tends to occur in hillsides. N.S. and Que. w. to Man. and Sask., s. to N.B., N.Y.,
dry to moist sites. (C. calceolus var. parvijlorum (Salish) Pern.) Mich., Wisc., and n. Minn.
3. Arethusa (Arethusa)
Perennial herbs, arising from a bulb; stem bearing a single grass-like leaf; flowers solitary. erect or arching. sepals and petals similar;
labellum 3-lobed. middle lobe long, arching; fruit a capsule.
Fig. 593. Cypripedium reginae: a. habit (Henry, Buker, and Pearth, 1975).
Cypripedium acaule: b. habit (WVA).
Cypripedium arietinum: c. habit; d. flower (Crow).
Cypripedium parviflorum: e. habit (WVA).
331
d
332
c
spur----''t-
e
a
Fig. 595. Pogonia ophioglossoides: a. habit; b. flower, side view; c. capsule (C&C).
Platantherapsycodes: d. inflorescence (G&W).
Platanthera leucophaea: e. upper portion of plant; f. flower (NYS Museum).
333
334 ANGIOSPERMS
4. Pogonia (Pogonia)
Perennial herbs, arising from a short rhizome; stems usually bearing a single, ovate leaf; flowers 1 or 2; sepals and petals similar;
labellum unlobed, bearded on upper surface; fruit a capsule.
1. P. ophioglossoides (1.) Ker-Gawl. Rose Pogonia Fig. 595 to N.C. and Tenn., Ohio, and n. Iil.; disjunct to se. Mo. (P. ophio-
Sphagnum bogs, peaty swales, and swamps. Nfld. w. to Ont. and glossoides var. brachypogon Fern.)
Minn., s. chiefly along the coastal plain to Fla. and e. Tex., mts.
Perennial herbs, with elongate, tuberous roots; leaves scattered along stem; flowers in racemes, subtended by bracts; labellum vari-
ous, entire to strongly 3-parted, often toothed to fringed; spur present, conspicuous, often longer than labellum; fruit a capsule.
This temperate group of terrestrial orchids is treated in the genus Habenaria (sensu lato) in older literature, but Luer (1975) notes
that no true Habenaria occur north of the subtropical region of the United States.
1. P. grandij10ra (Bigel.) Lindl. Large Purple Fringed Orchid 3. P. leucophaea (Nutt.) Lindl. Prairie Orchid, Prairie White
Swamps, wet meadows, thickets, and wet woods. Nfld. w. to s. Fringed Orchid Fig. 595
Ont., Wisc., and e. Minn., s. to n. N.J., mts. to N.C. and Tenn., Wet prairies, marly meadows, margins of lakes. and fens. Very
and Ohio. This species appears in most manuals as Habenaria rare; w. N.Y. (extinct?) and s. Ont. w. to Mich., s. Wisc., s. Minn.,
(Platanthera) fimbriata, but Stoutamire (1974) has determined s. Man., and se. N.D., s. to Ohio, Ill., ne. Kans., se. Okla. (ex-
that the correct name is P. grandij1ora. The taxon is sometimes tinct?), and n. La.; disjunct to single stations in n. Me., c. N.B.,
treated as a variety of the smaller-flowered, but very similar, P. and Va. (Habenaria leucophaea (Nutt.) Gray)
psycodes. (Habenaria grandij10ra (Bigel.) Torr.; H. fimbriata (Dry-
ander) R. Br.; Platantherafimbriata (Dryander) Lindl.) 4. P. lacera (Michx.) G. Don Ragged Fringed Orchid, Green
Fringed Orchid Fig. 596
2. P. psycodes (L.) Lindl. Small Purple Fringed Orchid Marshes, sedge swamps, bogs, wet woods, wet to dry open fields
Fig. 595 and prairies, and ditches. Nfld., N.S., and N.B. w. to Ont. and se.
Wet meadows, stream margins, margins of bogs, and open, wet Man., s. to S.C., Ga., Miss., Ark., ne. Tenn., and e. Okla. Our
woods. Nfld. w. to Ont. and e. Minn., s. to n. N.J., mts. to N.C., taxon is var. lacera. (Habenaria lacera (Michx.) R. Br.)
Ky., and Tenn., Ohio, n. Iil., and ne. Iowa; disjunct to Mo. and
Ark. White color forms are occasionally encountered. (Habena-
ria psycodes (L.) Spreng.)
labellum
Fig. 596. Platanthera lacera: a. lower portion of plant; b. inflorescence; c. flower. side view; d. petal; e. upper
sepal; f. labellum; g. h. flowers. hybrids (C&C).
335
e
Fig. 597. Platanthera blephariglottis: a. habit; b. flower. side view (more southerly variant). c. flower. side view
(occasional variant); d. flower. side view (more northerly variant); e. labellum (G&W).
Platanthera ciliaris: f. inflorescence (C&C).
336
Fig. 598. Platanthera cristata: a. basal portion of plant; b. upper portion of plant; c, d. flower, two views (G&W).
337
Fig. 599. Platanthera nivea: a. habit; b. flower; c. flower, side view (C&C).
Platanthera clavellata: d. habit; e. flower; f. flower, side view (C&C).
338
~~
'~V-d
340
Orchidaceae I Orchid Family 341
5. P. blephariglottis (Willd.) Lind!. White Fringed Orchid naria clavellata (Michx.) Spreng.; H. clavellata var. ophioglossoides
Fig. 597 Fern.)
Bogs, marshes, and wet peaty soils. NfId. w. to s. Que., s. Ont.,
and Mich., s. to N.J., Pa., and n. Ohio; c. Ill. A taxon with very 9. P. nivea (Nutt.) Luer Snowy Orchid Fig. 599
long spurs has been described as var. conspicua (Nash) Luer and Wet peaty soils, meadows, and savannas. Chiefly coastal plain,
occurs on the coastal plain and piedmont from s. N.J. to Fla., w. s. N.J. s. to Fla., w. to e. Tex. (Habenaria nivea (NutL) Spreng.)
to Tex. (Habenaria blephariglottis (Willd.) Hook.)
10. P. dilatata (Pursh) Lind!. ex Beck White Bog Orchid, Bog
6. P. ciliaris (L.) Lind!. Yellow Fringed Orchid Fig. 597 Candles Fig. 601
Bogs, marshes, swamps, wet meadows, pcaty or sandy woods, Wet meadows, swales, bogs, and mossy swamps. Green!' and
and dryish swales. S. N.B. w. to s. Ont., s. Mich., and ne. Ill., Lab. w. to Alask., s. to n. N.J., nw. Pa., n. Ill., Minn., mts. to N.M.,
s. to Fla., se. Mo., and e. Tex. (Habenaria ciliaris (L.) R. Br.) Utah, and Calif. Our plants belong to var. dilatata. (Habenaria
dilatata (Pursh) Hook.)
7. P. cristata (Michx.) Lind!. Crested Fringed Orchid, Orange-
crest Orchid Fig. 598 11. P. hyperborea (L.) Lind!. Northern Green Orchid
Meadows, sedge and sphagnum bogs, flatwoods, pine savannas, Fig. 600
margins of streams, and cypress swamps. Se. Mass., Conn., and Wet meadows, bogs, swales, and peaty thickets. N.S. and Que.
se. N.Y. w. to e. Pa., Va., se. Ky., Tenn., and Ark., s. to Fla. and w. to Ont. and Minn., s. to N.B., nw. Pa., n. Ohio, n. Ind., n.
e. Tex. (Habenaria cristata (Michx.) R. Br.) II!., and n. Iowa. Plants in our range are the more robust var.
huronensis. (Habenaria hyperborea (L.) R. Br.)
8. P. clavellata (Michx.) Luer Green Woodland Orchid
Fig. 599 Platanthera integra and P. peramoena may also be encountered in
Wet meadows, shores, stream banks, mossy woods, thickets, wet sites.
and ditches. Nfld. w. to Ont. and Minn., s. to Ga. and La. (Habe-
6. Listera (Twayblade)
Perennial herbs, with fibrous roots; stems bearing a single pair of leaves near middle; flowers in racemes; labellum elongate, entire
to notched to deeply cleft (ours) at tip; fruit a capsule.
1. L. cordata (L.) R. Br. Heart-leaved Twayblade Fig. 601 Ohio, Ont., n. Mich., n. Minn., Man., Alta., mts. to Colo. and
Sphagnum bogs and mossy coniferous swamps. Green!' and N.M., Ida., and n. Calif. Our taxon is var. cordata.
Lab. w. to N.W.T. and Alask., s. to N.B., n. N.J., mts. to N.C., ne.
7. Calopogon (Grass-pink)
Perennial herbs, arising from a bulb-like tuber; leaves solitary, basal; flowers pink, in racemes, appearing inverted, labeIIum upper-
most, column below it; fruit a capsule.
l. C. tuberosus (L.) BSP. Grass-pink Fig. 602 Calopogon pallidus Chapm., a paler-flowered species, may be en-
Sphagnum bogs, peaty meadows, swamps, marshes, and wet countered in the damp pine barrens of southeastern Virginia.
shores. Nfld. w. to Ont. and se. Man., s. to Fla. and e. Tex.; w.I.
(c. pulchellus (Salisb.) R. Br.; C. pulchellus var. latifolius (St.
John) Fern.)
Perennial herbs, with a cluster of thick roots; stem bearing leaves at or near base; inflorescence more or less densely spike-like,
flowers white to ivory, spiraled; labeIIum oblong to ovate, margin wavy or recurved, rarely lobed; column short; fruit a capsule.
REFERENCES: Case and Catling, 1983; Catling 1981, 1982; Sheviak, 1982.
Fig. 602. Calopogon tuberosus: a. basal portion of plant; b. inflorescence; c. labellum. longitudinal section.
showing papillae; d. portion of column; e. pollen tetrads; f. pollinia; g. capsule; h. seed. highly magni-
fied (C&C).
342
Fig. 603. Spiranthes romanzoffiana: a. basal port
ion of plan t and inflorescence; b. flow
er. side view; c. labellum
(Mason).
Spiranthes cerlma: d. habit; e. flower.
side view; f. labellum (C&C).
343
c
Fig. 604. Spiranthes odorata: a. habit; b, c. flowers, side views; d. labellum; e. flower, front view (NYS Museum).
344
)1( {
I
. I
1
~.\
Fig. 605. Spiranthes praecox: a. habit; b. flower, side view; c. labellum, side view; d. labellum, front view (C&C).
Liparis loeselii: e. habit, with flowers and with previous year's stem and fruits; f. column and portion
of labellum, side view; g. flower, front view (G&W).
345
Fig. 606. Malaxis brachypoda: a. habit; b. flower (PB).
Malaxis paludosa: c. habit (PB).
346
Orchidaceae I Orchid Family 347
1. Flowers of spike twisted, usually into 2 I-ranked groups, secund (fig. 605a).
4. Labellum widest at base; flowers white to cream, with yellow center; spike pubescent ...................... 4. S. Iaciniata
4. Labellum widest at tip; flowers white, with green veins; spike nearly glabrous or only sparsely pubescent .......... 5. S. praecox
1. S. romanzoffiana Cham. Hooded Ladies' -tresses Fig. 603 gins of streams. Coastal plain, Va. s. to Fla., w. to e. Tex.; disjunct
Meadows, swamps, swales, margins of ponds, and alkaline bogs; in Ky. (S. cernua var. odorata (Nutt.) Correll; Ibidium odoratum
often calcareous sites. Lab. and Nfld. w. to Alask., s. to Mass., w. (Nutt.) House; Triorchis odorata (Nutt.) Nieuwl.)
Pa., n. Ohio, Mich., Iowa, Neb., N.M., Ariz., and Calif. (Ibidium
romanzoffianum (Cham.) House; Triorchis romanzoffiana (Cham.) 4. S. Iaciniata (Small) Ames Lace-lip Ladies'-tresses
Nieuwl.); Spiranthes stricta (Rydb.) Nels.) Wet meadows, marshes, shallow ponds, and wet pinelands.
Coastal plain, s. N.J. s. to Fla., w. to e. Tex. (Ibidium Iaciniatum
2. S. cernua (L.) Richard Nodding Ladies'-tresses Fig. 603 (Small) House; Triorchis laciniata (Small) Nieuwl.)
Wet to dryish meadows and ditches, swales, lakeshores, and
bogs. P.E.I., N.S., and s. N.B. w. to s. Ont., Mich., Minn., and 5. S. praecox (Walt.) S. Wats. Giant Ladies'-tresses Fig. 605
Neb., s. to n. Fla. and e. 'leX. (Ibidium cernuum (L.) House; Tri- Wet meadows, swamps, swales, and open wet woods. Coastal
orchis cernua (L.) Nieuwl.~ plain, s. N.J. s. to Fla., w. to e. Tex. (Limodorum praecox Walt.;
Ibidium praecox (Walt.) House; Triorchis praecox (Walt.) Nieuwl.)
3. S. odorata (Nutt.) Lindl. Fragrant Ladies'-tresses Fig. 604
Shallow water and muddy shores, swamps, marshes, and mar-
9. Liparis (Twayblade)
Perennial herbs, arising from bulbs or tubers; stems with pair of broad, basal leaves; flowers few, in a raceme; labellum entire, widest
at tip; column elongate; fruit a capsule.
1. L. loeselii (L.) RichardBog Twayblade, Fen Orchid Md., mts. to N.C., Ohio, Ill., Iowa, and N.D.; disjunct to e. Ala.,
Fig. 605 s. Mo., and ne. Kan.
Bogs. fens, peaty meadows, and mountain ravines. N.S. and
Gaspe Pen., Que., w. to s. Ont., Man. and e. Sask., s. to n. N.J.,
Small perennials, arising from tubers; sterns bearing 1 or few broad leaves; flowers small, in dense racemes; petals smaller than
sepals, spreading or recurved; labellum auriculate at base, tapering toward tip; column very short; fruit a capsule.
1. Leaves solitary (rarely 2); stem not swollen at base .............................................. 1. M. brachypoda
1. Leaves 2-5; stem swollen at base, forming a pseudobulb ........................................... 2. M. paludosa
1. M. brachypoda (Gray) Fern. White Malaxis Fig. 606 2. M. paludasa (L.) Sw. Bog Malaxis, Bog Adder's-mouth
Damp calcareous gravels, wet meadows, peats, swales, and Fig. 606
bogs. S. Lab. and Nfld. w. to Ont., c. Sask., c. Alta., B.C., and s. Sphagnum bogs. Rare; n. Ont. and n. Minn., w. sporadically to
Alask., s. to n. N.J., Pa., Mich., ne. Ill., and ne. Minn. (M. mano- Alask.
phyllas var. brachypada (Gray) Morris & Eames)
Abaxial. Pertaining to the side of an organ away from the axis, Aphyllous. Lacking leaves.
such as the lower surface of a leaf; cr. Adaxial. Apical. Pertaining to the apex.
Abscise. To separate from. Apiculate. Tapering to an abrupt short, pointed tip.
Acaulescent. Stemless or apparently so, or with the stem sub- Apocarpous. Gynoecium with separate carpels.
terranean. Apomictic. Typically, setting seed without fertilization.
Achene. A hard, dry, indehiscent, one-seeded, one-locular Appressed. Lying close and pressed against; adpressed.
fruit. Aquatic. Living in water.
Actinomorphic, Descriptive of a flower or set of flower parts Arborescent. Approaching the size and habit of a tree.
which can be cut through the center into equal and similar Arcuate. Arching; bowed; moderately curved.
parts along two or more planes; radially symmetrical; regular. Areolate. Marked out into small spaces.
Acuminate. Long-tapering to a pointed apex, the sides of the Aril. An appendage or outer covering of a seed, growing out
angle somewhat concave. from the hilum or funiculus; often appears as a fleshy seed
Acute. Sharply angled, the sides of the angle essentially covering.
straight, the angle less than 90 degrees. Arillate. Pertaining to a seed having an aril.
Adaxial. Pertaining to the side of an organ toward the axis, Aristate. Bearing a stiff bristle-like awn or seta; tapered to a
such as the upper surface of a leaf; cf. abaxial. very narrow, much elongate apex.
Adnate. Pertaining to the fusion of one structure to another, Armed. Provided with any kind of strong or sharp defense
such as the stipules to the petioles in some species of Potamo- structures such as thorns, spines, prickles, barbs.
geton. Articulate. Jointed; provided with joints or places where sepa-
Adventitious. Pertaining to organs arising from an irregular ration may naturally take place.
or unusual position, such as roots from the stem, or leaves or Ascending. Rising upward; produced somewhat obliquely or
buds from a root. indirectly upward.
Aerenchyma. Type of plant tissue consisting of thin-walled Attenuate. Gradually narrowed to a point at apex and/or
cells with numerous, large intercellular spaces. base.
Agamospermy. Pertaining to the development of seeds with- Auricle. An ear-shaped part of appendage, such as the projec-
out fertilization; a type of apomixis. tions at the base of some leaves.
Aggregate. Crowded into a dense cluster, but remaining sep- Auriculate. Having ear-like lobes; having auricles.
arate. Awl-shaped. Narrow and sharp-pointed, gradually tapering
Alternate. Leaves or other parts borne with but one at each from the base to a slender, stiff point; subulate.
level on the stem. Awn. A stilI, bristle-like appendage, often at the end of a
Ament. A catkin; a lax, usually pendulous, spike-like inflores- structure.
cence, usually bearing apetalous, unisexual flowers. Axil. The angle between an organ and the axis which bears it,
Anastomosing. Pertaining to veins or ridges branching and such as between the leaf and the stem.
merging so as to form a network. Axillary. Located in the axil.
Androecium. A collective term for the stamens. Axis. The main line of development of a plant or organ, such
Androgynous, Pertaining to an inflorescence having both sta- as the main stem.
minate and pistillate flowers, the staminate flowers in the api- Barbed. Having rigid points or short bristles, usually reflexed.
cal, or upper, portion; cr. gynaecandrous. Beaked. Ending in a firm, prolonged slender tip, such as beaks
Annual. A plant which completes its life history within one of fruits or pistils; rostrate.
year. Bearded. Bearing long stiff hairs or bristles; pertaining to a
Anther. The pollen-bearing part of the stamen. tuft or zone of pubescence.
Anthesis. Flowering, or strictly the time of expansion of a Berry. A fleshy fruit with few to many seeds, but lacking
flower when pollination may take place; often used to desig- stones.
nate the flowering period. Biconvex. Lenticular; having both surfaces rounded.
Antrorse. Having hairs or processes directed upward or for- Biennial. A plant which requires two years to complete its life
ward; cr. Retrorse. cycle, the first year growing only vegetatively, often producing
Apetalous. Without petals. only a basal rosette, then flowering, fruiting, and dying the
Apex. The tip or distal end. second year.
353
354 Glossary of Plant Terms
Bifid. Two-cleft; split into two segments. such as at the apex of Carpel. A segment or unit of a compound pistil or the single
some leaves or petals. unit of a simple pistil; the ovule-bearing units of an ovary.
Bilabiate. Two-lipped. Cartilaginous. Firm and tough. but flexible.
Bilaterally symmetrical. Descriptive of a flower or set of Catkin. An ament; a lax. usually pendulous. spike-like inflo-
flower parts which can be cut through the center into two rescence. usually bearing apetalous. unisexual flowers.
equal and similar parts along only one plane; zygomorphic; Caudate. Having a tail-like appendage; as at leaf apices.
irregular. Caudex. A persistent base of stem. often woody. from which
Bipinnate. Twice. or doubly, pinnately compound. new aerial herbaceous stems arise each year.
Bipinnatifid. Twice pinnatifid. Cauline. Pertaining to the stem; as opposed to basal.
Bisexnal. Having both stamens (male parts) and pistils (fe- Cespitose. Growing in tufts; in dense clumps; having several
male parts) present and functional in one flower. as in bisex- to many stems in a close basal tuft.
ual flowers. Chaff. A thin. dry. membranous scale or bract. particularly the
Biternate. Pertaining to leaves which are twice divided into bracts subtending flowers in flower heads of the Asteraceae.
threes (three main divisions themselves divided into three Chaffy. Having or resembling chaff.
parts; nine leaflets); cf. Triternate. Channeled. Having a deep longitudinal groove; furrowed.
Bladder. A thin. sac-like structure; as in Utricularia (Blad- Chartaceous. Thin but stiff; papery texture; usually not green.
derwort). Chlorophyllous. Containing chlorophyll. thus typically giv-
Blade. The expanded. usually flattened. portion of a leaf. bract. ing a green or greenish appearance; as opposed to achloro-
petal. or other structure. phyllous. lacking chlorophyll.
Bloom. A whitish. usually waxy covering on the surface. Ciliate. Having hairs extending from the margins or edges.
Bract. A reduced leaf or leaf-like structure. particularly sub- Ciliolate. The diminutive of ciliate.
tending a flower or an inflorescence. Clasping. Partly or completely surrounding the stem.
Bracteate. Bearing bracts. Clavate. Club-shaped. gradually thickened toward the top.
Bracteole. The diminutive of bract; bractlet. Cleft. Deeply cut; having divisions to or nearly to the middle.
Bractlet. A very small bract. or sometimes applied to bracts in such as a palmately or pinnately cleft leaf.
a secondary position; bracteole. Cleistogamous. Fertilized in bud without the flower opening.
Branchlet. Ultimate division or divisions of a branch. Collar. Outer side of grass leaf at the junction of sheath and
Bristle. A stiff hair or trichome. blade.
Bud. An embryonic axis with its appendages. such as an unex- Column. In the Orchidaceae. a coalescence of stamens and
panded flower or a bud in the axil of a leaf. style into a single. somewhat elongate body.
Bulb. A short underground stem surrounded by succulent Commissure. The face or plane along which two carpels join.
leaves or scales. such as the face by which one mericarp joins another in a
Bulblet. A small bulb-like structure often borne in the axil of schizocarp.
a leaf. inflorescence. or other unusual area. Comose. Bearing a tuft or tufts of soft hairs at one end.
Bulbose. Having bulbs; the structure of a bulb. Compound. Composed of several parts united as a common
Bulbous. Bulb-like. whole. such as a compound pistil. or a compound leaf com-
Callous. Having the texture of a callus. posed of two or more distinct leaflets.
Callus. A thick. leathery, or hardened protuberance; in Compressed. Flattened laterally.
grasses. the tough swelling at the base. or point of insertion. Concave. Pertaining to a surface or margin rounded inwardly;
of the lemma or palea. cf. Convex.
Calyx. Collective term for sepals; the outer. usually green. Cone. A structure consisting of seed-bearing scales arranged
whorl of floral parts; outer series of perianth parts. frequently on a usually elongate axis. such as a pine cone.
enclosing the rest of the flower in the bud stage; sometimes Congeneric. Belonging to the same genus.
colored or petal-like. or sometimes greatly reduced or lacking. Conglomerate. Densely clustered.
Campanulate. Bell-shaped. Conic. Cone-shaped.
Canescent. Gray-pubescent; surface covered with a gray or Connate. United; especially applied to like structures. such as
whitish pubescence. the bases of two opposite leaves.
Capillary. Thread-like or hair-like. very slender. Conspecific. Belonging to the same species.
Capitate. Aggregated into a dense. compact cluster; formed Contiguous. Touching. but not fused.
like a head. Contorted. Twisted; bent or twisted on itself.
Capitulum. A dense inflorescence. consisting of an aggrega- Convex. Pertaining to a surface or margin that is rounded out-
tion of usually sessile flowers; a head. wardly; cf. Concave.
Capsule. A dry dehiscent fruit resulting from the maturation Convolute. Rolled or twisted together in bud stage; rolled lon-
of a compound ovary (of one or more fused carpels). gitudinally in winter buds of some Potamogeton species.
Glossary of Plant Terms 355
Cordate. Heart-shaped; with a sinus and rounded lobes at the Dentate. Toothed. with teeth directed outward. not forward.
base; often applied to the basal portion rather than the en- Denticulate. Minutely or finely dentate.
tire organ. Depauperate. Stunted. much reduced and imperfect in struc-
Coriaceous. Leathery in texture, such as the tcxture of many ture and development.
evergreen leaves. Depressed. Somewhat flattened from above; pressed down.
Corm. A thick, solid, bulb-like stem surrounded by papery Determinate. A defincd number or growth; an inflorescence
scales, usually underground. in which the terminal or central flower opens first. thus the
Corolla. A collective term for petals; the inner whorl of a peri- axis prolongation is arrested.
anth, usually colored or otherwise differentiated; frequently Diadelphous. Having stamens which are united in two
the showy part of the flower, but sometimes reduced or lack- groups by their filaments; as in Fabaceae.
ing; if petals are united, the corolla may be divided into a co- Diaphragmed. Having cross-partitions; as in the pitch of
rolla tubc and lobes or lips, and is then said to be sympetalous Nyssa.
or gamopetalous. Dichotomous. Forking in pairs.
Corona. A crown or inner petal-like appendage bctween the Diffuse. Loosely branching or spreading; descriptive of open
petals and stamens, such as in Asclepias. growth.
Corrugated. Having a surface that is wrinkled or in folds. Digitate. Having finger-like divisions; a compound with mem-
Corymb. A flat-topped, racemose type of inflorescence in bers arising from one point.
which the lower pedicels are successively elongate, the flow- Dimorphic. Occurring in two forms. such as two types of
ering sequence indeterminate. with the marginal flowers leaves or two types of seeds.
opening first. Dioecious. Flowers that are unisexual. with staminate and
Corymbose. Pertaining to flowers or fruits arranged in or re- pistillate flowers on separate plants.
sembling a corymb. Diploid. Having two full sets of chromosomes in each cell.
Creeper. A trailing shoot, usually rooting at most of the nodes. Discoid. Having only disk flowers, as in some Asteraceae;
Creeping. Pertaining to a shoot running along at or ncar thc disk-shaped.
surface of the ground and often rooting at the nodes; repent. Disk. A development of the receptacle at or around the base of
Crenate. Having a margin with low, rounded lobes or teeth. the pistil; in the Asteraceae the central portion of the flow-
Crenulate. Finely crenate; having very small rounded lobes ering head. bearing tubular flowers.
or teeth. Disk flower. Applied to the tubular flowers in the central por-
Crested. Having a ridge-like projection on the surface or across tion of the flowering heads on many Asteraceae (distin-
the top. guished from the ray flowers).
Crisped. Having a wavy surface toward the margin. Dissected. Divided or cut into many small. slender segments.
Cruciform. Cross-shaped, such as tlowers of Brassicaceae. such as dissected leaves.
Cucullate. Hooded or hood-shaped. Distal. At or toward the end farthest from the point of attach-
Culm. The aerial stem of a grass. sedge, or rush. ment; c[ Proximal.
Cuneate. Wedge-shaped; narrowly triangular, tapering toward Distichous. Two-ranked. on opposite sides of an axis and in
the point of attachment. the same plane.
Cuspidate. Tipped with a short, tooth-like rigid point. Distylous. Pertaining to flowers of a given spccies having two
Cyme. A broad, flat-topped inflorescence in which the different style lengths and anthers borne at corresponding
branches arise from the same point, and the flowering se- levels; as in the Lythraceae; cf. Tristylous.
quence is determinate, with the central flowers opening first. Divergent. Spreading broadly; inclining away from each
Deciduous. Not persistent, falling at the end of one season of other.
growth; not evergreen. Divided. Lobed or separated to the base; often referring to the
Decompound. More than once compound. leaf blade when it is cut into distinct divisions to or almost to
Decumbent. Growing reclining on the ground, but with the the midvein.
apical portion ascending. Dorsal. Pertaining to or located on the back, or upper (adax-
Decurrent. Extending downward from the point of insertion. ial). surface; cf. Ventral.
usually applied to leaves apparently prolonged downward Dorsiventral. Flattened with definite upper or front (adaxial)
along the stem, often as wings. and lower, or back (abaxial). surfaces.
Decussate. Pertaining to organs, especially leaves. borne op- Downy. Pubescent; pertaining to a surface covered with fine.
pOSitely on the stem, each pair alternating at right angles soft hairs.
with the pairs above and below. Drupe. A fleshy. indehiscent fruit with the seed enclosed in a
Dehiscent. Opening and shedding contents; applied to sta- stony endocarp or inner layer of fruit wall (stone); stone fruit.
mens and fruits as opening by valves. slits. pores, etc. Druplet. One small drupe in an aggregate fruit made up of
Deltoid. Pertaining to shapes that are triangular in outline. drupes; as in Rubus (blackberries. raspberries).
356 Glossary of Plant Terms
Elator. Band like structure which is sensitive to changes in hu- Floccose. Surface covered with irregular tufts of soft wooly or
midity, and is an aid in dispersal of the spores, as in Equisetum. cottony hairs which usually rub off readily.
Ellipsoid. A solid with an elliptic outline. Floral tube. Tubular structure in the flower derived by the fu-
Elliptic. Oval or oblong in outline, with the ends equally nar- sion of the floral envelopes and the androecium, on which are
rowed or rounded. borne the calyx, corolla, and stamens; sometimes referred to
Emarginate. Having a shallow notch at the apex. as a hypanthium.
Embryo. The rudimentary plant within the seed. Floret. A small flower, usually one of a dense cluster; in
Emergent. Extending out of the water; emersed. grasses, a unit consisting of the pistil and stamens sur-
Emersed. Extending out of the water; emergent. rounded by the lemma and palea.
Endocarp. The inner layer of a fruit wall, sometimes becoming Flower. An axis bearing stamens or pistils or both, typically sub-
hard or bony, as in a drupe. tended by a perianth, although the perianth can be lacking.
Entire. Having a margin without any indentations, teeth, or Foliaceous. Leaf-like in appearance or texture.
lobes (mayor may not be hairy, or ciliate). -foliolate. Suffix pertaining to the number ofleaflets in a com-
Epicalyx. A series of bracts closely sub tending and resembling pound leaf; cf. Trifoliolate.
sepals or a calyx. Follicle. A dry dehiscent fruit splitting along only one suture;
Epigynous. Pertaining to a flower in which the calyx, corolla, the product of a simple pistil.
and stamens are borne at the summit of the ovary, with the Forked. Divided into nearly equal segments.
ovary fused to an expanded receptacle or floral tube, the Foveolate. Pitted.
ovary thus said to be inferior. Frond. In the Lemnaceae, the green thallus-like stem which
Epipetalous. Descriptive of stamens when they are inserted on functions as foliage; the leaf in ferns.
the corolla. Fruit. The seed-bearing organ; the matured pistil.
Epiphyte. A plant which grows upon another plant, but with- Funnelform. Having the shape of a funnel. the tube gradually
out parasitizing it. widening into a limb; usually applied to the shape of the co-
Equitant. Descriptive of leaves overlapping in two ranks; as rolla.
in Iris. Furrowed. Having longitudinal channels or grooves; chan-
Erect. In an upright position; ascending. neled.
Erose. Having an uneven margin, appearing gnawed, or having Fusiform. Tapering at both ends; spindle-shaped.
a margin with very small teeth of irregular shape and size. Gemma. An asexual propagule, sometimes appearing as a
Estuarine. Growing in estuaries. vegetative bud.
Evergreen. Remaining green through the dormant season Geniculate. Bent abruptly.
and functional the following growing season. Gibbous. Swollen on one side, such as on a calyx or corolla
Exfoliate. To shed or come off in thin scale-like or flake-like tube.
layers. Glabrate. Becoming glabrous with age.
Exocarp. The outermost layer of the fruit wall. Glabrous. Smooth, lacking pubescence; not hairy.
Exserted. Extending beyond; sticking out; such as stamens Gland. A secreting structure, often a protuberance or append-
from the perianth. age, frequently the surface sticky, waxy, or glistening.
Falcate. Curved and flat. like a sickle. Glanduliferous. Bearing glands.
Farinose. Surface covered with a mealy powder. Glandular-pubescent. Having hairs, or trichomes, termi-
Fascicle. A bundle or cluster, such as the leaves of Pinus or the nated by pinhead-like glands, often sticky.
stamens of Hypericum. Glaucous. Covered with a whitish waxy bloom or powdery
Fasciculate. Congested in close clusters or bundles. coating.
Feather-veined. Having veins all branching from the sides of Globose. Shaped like a sphere or globe.
a midrib. Glochidia. Barbed or hooked hairs or bristles.
Fetid. Having a disagreeable odor. Glomerate. Densely or compactly clustered; often referring to
Filament. The portion of the stamen bearing the anther. flowers occurring in dense clusters.
Filiform. Thread-like, long, and very slender. Glomerulatc. Having small, dense clusters of flowers.
Fimbriate. Fringed; cut into regular segments giving a Glomerule. A small cluster of flowers, usually consisting of a
fringed appearance. densely compacted cyme.
Fimbrillose. Abounding with fringe. Glume. One of a pair of bracts lowermost on a grass spikelet.
Flabellate. Fan-shaped or broadly wedge-shaped. Glutinous. Covered with a sticky exudate; viscid.
Flaccid. Limp, lax, and weak; floppy. Grain. The seed-like fruit of a grass; the small hard structure
Flexuous. Having a wavy appearance; especially applied to in the fruiting structure of Rumex.
submersed aquatics where plants are not rigid, thus they Granulose. Composed of or appearing covered with minute
readily bend in flowing water. grains.
Glossary of Plant Terms 357
Gynaecandrous. Pertaining to an inflorescence with stami- Hypogynous. Pertaining to a !lower having the calyx, corolla,
nate and pistillate flowers in the same spike, the pistillate and stamens attached below the ovary, the ovary thus said to
flowers in the apical, or upper, portion; especially used in the be superior.
Cyperaceae (Carex); cf. Androgynous. Imbricate. Overlapping, vertically or spirally, with the upper
Gynoecium. Collective term for the pistil or pistils of a flower. portion covering the base of the next higher.
Gynostegium. A complex structure in the flowers of the As- Immersed. Growing wholly under water; submersed.
clepiadaceae whereby the upper portion of the gynoecium is Imperfect. Pertaining to !lowers having functional stamens or
united with the androecium. pistils, but not both; unisexual.
Habit. The general growth form of a plant, such as herbaceous Incised. Having deeply cleft margins.
or woody. Indehiscent. Pertaining to fruits that remain closed.
Habitat. The type of environmental conditions under which a Indusium. A thin, epidermal outgrowth covering the sori on
plant grows. ferns.
Halophyte. A plant of saline soil. Inferior ovary. The calyx, corolla, and stamens arise from the
Hastate. Arrow-shaped, with basal lobes that spread or extend summit of the ovary because of the fusion of tissue to the
downward and outward, sometimes nearly at right angles; ovary wall; d. Superior ovary.
halberd-shaped. Inflated. Bladdery.
Head. An inflorescence consisting of a dense cluster of sessile Inflorescence. An aggregation of flowers occurring together
or nearly sessile flowers on a very short axis; a capitulum; as in a particular arrangement.
in Astcraceae. Infrudescence. An inflorescence when the fruits develop.
Helicoid. Spiraled or coiled like a helix. Inserted. Attached to or growing out of.
Herb. A plant with no persistent woody stem above ground. Internode. The portion of a stem between the nodes or points
Herbaceous. Having the structure or texture of an herb, not where leaves are borne.
woody; dying back to ground level each year; sometimes ap- Introduced. Brought from another region.
plied to bracts which are not stiff and cartilaginous. Involucral. Pertaining to an involucre.
Heteromorphic. Having different growth forms. Involucre. A cluster of bracts in an inflorescence subtending
Heterophyllous. Having different types of leaves. or surrounding a flower cluster, or head, or a single flower.
Heterosporous. Producing spores of two different sizes, the Involute. Having margins that are rolled inward (toward the
larger (megaspore) developing into a female gametophyte, the upper, or adaxial, surface); d. Revolute.
smaller (microspore) developing into a male gametophyte; cf. Irregular. Descriptive of a flower or set of !lower parts which
Homosporous. can be cut through the center into two equal and similar
Hibernaculum. Winter bud. parts along only one plane; bilaterally symmetrical; zygo-
Hip. Pertaining to a fruiting structure which consists of a morphic.
fleshy, expanded vase-like hypanthium enclosing the achenes; Jointed. Having one or more constrictions marking a point of
used in Rosa. articulation; as in Equisetum.
Hirsute. Surface covered with long, rather coarse or stiff hairs. Keel. A central ridge, like the keel of a boat; as in some Spar-
Hirtellous. Minutely or softly hirsute. ganium.
Hispid. Surface covered with stiffish hairs that are sometimes Labellum. Lip; the distinctive lower (by twist of the pedicel!
bristly or spike-like. ovary) petal of the Orchidaceae.
Hispidulous. Minutely hispid; with short, stiff hairs. Labiate. Lipped, as applied to an unequally divided calyx or co-
Homomorphic. Having one growth form. rolla.
Homosporous. Producing spores which are all of one size; d. Lacerate. Irregularly cleft.
Heterosporous. Lacunae. Air spaces within a tissue, especially in the leaves,
Hyaline. Having a thin, membranous, transparent or translu- stems, and roots of submersed aquatic plants.
cent texture, Lamina. The blade or expanded part of a leaf, bract, petal, etc.
Hybrid. A plant resulting from a cross between parents that Lanceolate. Shaped like a lance, several times longer than
are genetically unlike; in taxonomy, the offspring of two differ- wide, broadest toward the base and narrowed to the apex.
ent species or infraspecific units. Lateral. Pertaining to the sides.
Hydrophyte. An aquatic plant. Latex. The milky juice or sap of some plants, such as Asclepias.
Hypanthium. A cup-like receptacle derived from the fusion of Lax. Loose or somewhat limp; not rigid or tight.
floral envelopes and androecium, and on which are borne the Leaflet. A division of a compound leaf.
calyx, corolla, stamens; sometimes applied to a floral tube de- Legume. Fruit in Fabaceae produced from a simple pistil, with
rived from the fusion of floral envelopes and androecium. many seeds and splitting along two sutures.
Hypogynium. The perianth-like structure sub tending the Lemma. The lower of the two bracts surrounding the pistil and
ovary in the Cyperaceae, especially Sc1eria. stamens in a flower in the grasses.
358 Glossary of Plant Terms
Lenticel. Corky spots on young bark of stems and roots, in- Nodulose. Having little knobs or knots.
volved in gaseous exchange; corresponding to epidermal Nut. A hard, indehiscent one-seeded fruit.
stomata. Nutlet. A small, hard, indehiscent one-seeded fruit.
Lenticular. Having both surfaces lens-shaped; biconvex. Obconic. Conic or cone-shaped, with the point of attachment
Ligule. The collar-like appendage, membranous or composed at the apex.
of hairs, at the junction of the blade and the sheath in the Obcordate. Heart-shaped, but inverted, with the point at the
leaf of grasses and a few Carex. base.
Linear. Long and narrow, with parallel or nearly parallel Oblanceolate. Lanceolate, but broadest above the middle,
margins. long-tapered to the base.
Lip. The upper or lower part of an unequally divided or bilabi- Oblique. Unequal-sided or slanting.
ate calyx or corolla; the labellum of Orchidaceae. Oblong. Longer than broad, with the sides nearly paralleL
Lobe. Any segment of an organ, especially if rounded. Obovate. Ovate, but inverted, broadest above the middle; d.
Locule. The cavity of an ovary or anther. ovate.
Loculicidal. Pertaining to the dehiscence of a capsule along Obovoid. Ovoid, but inverted, with the broader end toward the
the back of the locule, not along the septum. apex; cf. ovoid.
Loment. A specialized legume which is constricted between Obtuse. With the sides or margins meeting at more than a 90-
the seeds and breaks apart into one-seeded segments rather degree angle.
than splitting lengthwise along the sutures. Ocrea. A stipular sheath at the nodes in the Polygonaceae.
Massulae. A group or mass of cohering pollen grains, as in Ocreolae. The smaller or secondary stipular sheaths, as in the
the Orchidaceae, or microspores, as in Azolla. inflorescences in the Polygonaceae.
Median. Pertaining to the middle. Olivaceous. Olive-green.
Megasporangium. A structure in which megaspores are pro- Opaque. Not translucent.
duced. Opposite. Pertaining to leaves or bracts occurring two at a
Megaspore. The female or larger type of spore in Isoetes, Mar- node on opposite sides of the stem; in flower parts when one
silea, AzolIa, and Salvinia. part occurs in front of another.
Membranous. Thin, rather soft, and more or less translucent Orbicular. Circular in outline.
and pliable; membranaceous. Oval. Broadly elliptic.
Mericarp. A segment of a schizocarp, such as in the Apiaceae. Ovary. The part of the pistil in which the seeds develop.
-merous. Suffix pertaining to parts or their number, typically Ovate. Egg-shaped in outline, with the broader end toward
referring to the perianth only; as a 5-merous (pentamerous) the base.
flower having corolla parts in fives. Ovoid. A solid with an ovate outline.
Microsporangium. A structure in which microspores are Ovuliferous. Bearing ovules; in ovuliferous cones in Gymno-
produced. sperms, the seed-bearing structures.
Microspore. The smaller type of spore in Isoetes, Marsilea, Palate. A rounded projection or prominence of the lower lip of
AzolIa, and Salvinia. a bilabiate corolla, causing the throat of the corolla to be
Microstrobilus. A strobilus, or cone-like structure, bearing closed or nearly so.
microsporangia or pollen sacs; the male cones in Gymno- Palea. The tiny, often very thin, upper bract of a grass floret,
sperms; androstrobilus. enclosed by the lemma.
Midrib. The main or central vein of a leaf or leaflet or similar Palmate. Radiately lobed or divided.
structure. Paniculate. Pertaining to flowers or fruits arranged in or re-
Monadelphous. Having stamens which are united in a single sembling a panicle.
tube or column by their filaments. Panicle. A loose, irregularly branched inflorescence, the flow-
Monoecious. Flowers that are unisexual, with staminate and ering sequence indeterminate, with the lower flowers open-
pistillate flowers on the same plant. ing first.
Mucilaginous. Slimy. Papilla. A minute nipple-shaped projection.
Mucronate. Having a short, sharp, abrupt tip on a rounded Papillate. Having one or more papillae; papillose.
or truncate apex. Papillose. Bearing minute nipple-shaped projections; papil-
Naturalized. Having a foreign origin, but established and re- late.
producing without cultivation as though native. Pappus. The bristle-like, capillary, scaly, or chaffy structure
Nectary. Any area or appendage where nectar is secreted. (modified calyx) at the summit of the achene in the
Nerve. A simple or unbranched vein or rib. Asteraceae.
Netted. Reticulate; veins connected to form a network. Papule. A relatively large papilla, or nipple-like projection.
Node. The level of a stem at which one or more leaves or bracts Parallel-veined. Descriptive of an organ in which the veins
are borne. run toward the apex in approximately parallel lines.
Glossary of Plant Terms 359
Parted. Cleft to below the middle. ovary. style. and stigma. may be simple or compound (with
Pectinate. Descriptive of an organ which is cleft into divisions two or more carpels).
such as to resemble a comb; pinnatifid. with narrow. closely Pistillate. Pertaining to a flower that has a pistil or pistils. but
set segments. not stamens; unisexual. with pistils.
Pedicel. The stalk of an individual flower or fruit. Pith. The spongy central portion of a stem.
Pedicellate. Borne on a pedicel. Placentation. Pertaining to the arrangement of the placenta.
Peduncle. The primary stalk of an entire inflorescence. or the the interior area of the ovary bearing the ovules.
stalk of a flower when borne singly. Plaited. Plicate; with folds. usually lengthwise.
Pedunculate. Borne on a peduncle. Plano-convex. Having one surface rounded and the other
Pellucid-punctate. Having translucent dots. surface flat.
Peltate. Pertaining to a flat structure that is attached by its Plicate. Plaited; with folds. usually lengthWise.
lower surface. often toward the center. not at its margin; as Plumose. Having fine. elongate hairs arranged like the plume
in Brasenia and Nelumbo. of a feather.
Pendulous. More or less hanging. Podogyne. A stalk bearing the ovary or fruit. sometimes re-
Perennate. To overwinter; to survive from year to year. sembling a pedicel. but arising as an elongation at the base
Perennial. Living three or more years; herbaceous perennials of the ovary. within the calyx; gynophore; as in Ruppia.
usually dying back each winter. Pollinium. A mass of waxy pollen or of coherent pollen-
Perfect. Pertaining to a single flower having both functional grains. as in Asclepiadaceae and Orchidaceae.
stamens and a pistil or pistils. with a calyx and corolla either Polygamodioecious. Chiefly dioecious. but bearing some bi-
present or absent; bisexual. sexual flowers.
Perfoliate. Pertaining to leaves or bracts that apparently have Polygamomonoecious. Chiefly monoecious. but bearing some
the stem passing through them; opposite or whorled leaves bisexual flowers.
that are united in a collar-like structure around the stem. Polymorphic. Exhibiting more than one form; variable as to
Perianth. General term for a calyx and corolla together. or for habit.
either one if the other is absent. Pome. A fleshy. indehiscent fruit developed from an inferior
Perigynium. A special sac-like structure which encloses the ovary. with the seeds enclosed within a cartilaginous ovary
pistil and ultimately the fruit in Carex. wall. the fleshy part developed from the hypanthium.
Perigynous. Pertaining to a flower having a calyx. corolla. Prickle. A small and more or less slender sharp outgrowth
and stamens borne on a hypanthium. but without these tis- from the epidermis; internodal thorn.
sues being fused to the ovary. the ovary thus superior. Procumbent. Trailing or lying on the ground. but not typi-
Persistent. Pertaining to an organ that remains attached after cally rooting at the nodes.
completing its normal biological function. Prophyllum. The bracteole at the base of an individual flower.
Petal. A unit of the corolla. as in 'uncus.
Petaloid. Having the appearance of petals. such as petaloid Prostrate. Lying flat upon the ground.
sepals. Proximal. At or toward the base or the point of attachment;
Petiolate. Having a petiole. cr. Distal.
Petiole. The stalk of a leaf. Pubernlent. Covered with a pubescence of very short hairs.
Phyllary. An involucral bract in the Asteraceae. not densely spaced.
Phyllodial. Pertaining to a more or less flattened. expanded. Pubescent. Often a general term referring to a surface being
bladeless petiole that serves as a leaf blade; often linear or covered with hairs. in contrast with glabrous; a type of pubes-
sometimes resembling a ribbon-like leaf. cence with short. soft. downy hairs.
Pilose. Having long. soft hairs. Punctate. Having depressed dots scattered over the surface.
Pinna. Leaflet or primary division of a pinnately compound Raceme. A simple inflorescence of pediceled flowers along a
leaf. common axis. the flowering sequence indeterminate. with the
Pinnate. Compound. with leaflets arranged along the two lower flowers opening first.
sides of an axis. Racemose. Pertaining to flowers or fruits arranged in or re-
Pinnate-pinnatifid. A pinnately compound leaf with the sembling a raceme.
leaflets then cleft pinnately. such that the sinuses do not quite Rachilla. A secondary axis; specifically in the Cyperaceae and
reach the midvein. Poaceae. the floral axis. in contrast with the axis of the spike
Pinnatifid. Pinnately cleft. the sinuses not quite reaching the or spikelet.
midvein. Rachis. The central axis of an inflorescence or of a com-
Pinnule. The secondary unit of a twice pinnately compound pound leaf.
leaf. Radially symmetrical. Descriptive of a flower or set of flower
Pistil. The seed-bearing organ of the flower. consisting of the parts which can be cut through the center into equal and
360 Glossary of Plant Terms
similar parts along two or more planes; actinomorphic; Scale. A small, thin, flattened structure, usually a modified
regular. leaf, arising from a stem or other organ.
Radiate. Spreading from a common center; in the Asteraceae, Scape. A leafless or nearly leafless stem bearing flowers.
bearing ray, or Iigulate, flowers. Scapous. Bearing or resembling a scape.
Ray. The branch of an umbelliform inflorescence. Scarious. Thin, dry, and membranous, not green.
Ray flower. The strap-like, or Iigulate, flower in a head of Schizocarp. A fruit which splits into one-seeded segments
flowers of many Asteraceae. (mericarps), such as in the Apiaceae.
Receptacle. The more or less expanded portion of an axis Scorpioid. An inflorescence coiled in bud, as in Drosera and
which bears the parts of the flower, or the cluster of flowers Myosotis.
of a head. Scurfy. Pertaining to a surface covered with scale-like or bran-
Receptacular. Pertaining to the receptacle; as in receptacular like particles.
bracts sub tending the cluster of flowers in a head in the Secund. Having a part or element directed to one side only.
As1eraceae. Seed. A ripened ovule.
Recurved. Curved downward or backward. Segment. A subdivision or lobe of a deeply cleft or divided leaf
Reflexed. Sharply bent downward or backward from the point or similar organ.
of attachment. Sepal. A unit of the calyx.
Regular. Descriptive of a flower or set of flower parts which can Septate. Divided by partitions.
be cut through the center into equal and similar parts along Septicidal. Pertaining to the dehiscence of a capsule along
two or more planes; actinomorphic; radially symmetrical. the septum.
Reniform. Kidney-shaped. Septum. Any kind of partition.
Repent. Creeping; prostrate and rooting at the nodes. Sericeous. Silky; covered with soft, silky hairs which usually
Reticulate. Forming a network. point in one direction.
Retrorse. Having hairs or processes directed backward or Serrate. Having sharp teeth pointing forward.
downward; cf. Antrorse. Serrulate. Having very small marginal teeth; minutely ser-
Revolute. Pertaining to margins that are rolled backward (to- rate.
ward the lower, or abaxial, surface); cf. Involute. Sessile. Joined directly by the base, lacking a stalk, pedicel, or
Rhizoid. A simple, root-like structure. petiole.
Rhizome. Any prostrate or subterranean stem. Seta. A bristle or bristle-shaped structure.
Rhomboid. Having a rhombic outline; an equilateral oblique- Setaceous. Bearing bristles.
angled shape; four-sided, with a somewhat diamond outline. Setulose. Bearing minute bristles.
Rib. Prominent veins or ridges on structures such as some Sheath. The basal part of a leaf wrapped about the stem; a
fruits, seeds, bracts, perigynia, or stems. collar-like outgrowth at a node.
Riparian. Growing by rivers or streams. Sigmoid. Doubly curved or S-shaped.
Rootstock. An underground stem; rhizome. Silicle. A short silique. usually wider than long. in the Brassi-
Rosette. A cluster of leaves or other organs radiating from an caceae.
apparent common point of attachment. Silique. An elongate, two-celled fruit in the Brassicaceae in
Rostellum. A small beak; a slender extension between the which at dehiscence the two valves fall away. exposing a per-
stigmatic cavity and the anther in the Orchidaceae. sistent septum.
Rostrate. Having a beak. Silky. Covered with soft, straight hairs, usually pointing in one
Rotate. Pertaining to a sympetalous corolla having a short direction; sericeous.
tube and flaring into a broad. flat, and circular limb. Simple. Consisting of a single, or unbranched, unit, sometimes
Rufescent. Becoming reddish or reddish-brown. cleft, but not divided into distinct parts; not compound.
Rugose. Having a wrinkled surface, with sunken veins. Sinuate. Having a strongly wavy margin in the plane of the
Runner. A very slender stolon; a horizontal stem with long blade.
internodes trailing along the surface of the ground. Sinus. The cleft or angle between two lobes.
Saccate. Sac- or pouch-shaped. Smooth. Glabrous. or lacking hairs; without roughness.
Sagittate. Shaped like an arrowhead, with the basal lobes di- Sori. Clusters of sporangia in ferns.
rected downward or backward. Spadix. A spike with a fleshy axis, as in Araceae.
Salverform. Descriptive of a corolla having a slender tube that Spathe. A large, sheathing bract, often open on one side, en-
abruptly expands into a flat limb. closing an inflorescence.
Samara. An indehiscent winged fruit. Spatulate. Oblong. with the basal portion narrow. Widening
Scaberulous. Descriptive of a surface that is slightly rough to above, and rounded at the apex.
the touch. Spicate. Pertaining to flowers or fruits arranged in or resem-
Scabrous. Descriptive of a surface that is rough to the touch. bling a spike.
Glossary of Plant Terms 361
Spicule. A small, slender, sharp-pointed structure, usually on Succulent. Fleshy; soft and thickened in texture.
a margin or surface. Superior ovary. Occurs in a flower whose calyx, corolla, and
Spiculose. Having a margin or surface covered with fine points. stamens arise from the receptacle below the ovary; cf. Infe-
Spike. An elongate inflorescence with flowers sessile or ap- rior ovary.
pearing sessile. Supra-axillary. Borne at a distance above the axil, as in Spar-
Spikelet. A secondary spike in some sedges; the segment of ganium.
the inflorescence of grasses enclosed by the glumes. Suture. A line of opening or dehiscence.
Spine. A firm, slender, sharp-pointed structure; a sharp woody Sympetalous. Having the petals of the corolla united, at least
or rigid outgrowth from the stem, usually a modified stem. at the base; gamopetalous.
Spinulose. Having small spines over the surface or along the Syncarpous. Pertaining to a gynoecium with carpels united
margin. in one pistil; a compound pistil.
Sporangium. A structure in which spores are produced and Synsepalous. Having the sepals of the calyx united; gamosep-
contained. alous.
Spore. A microscopic, simple reproductive body, the product of Taproot. The primary descending root; a direct continuation
meiosis, which is capable of developing into a new individual, of the radicle of the embryo.
particularly in Pteridophytes and lower plants. Taxon. A general term applied to any taxonomic element or
Sporocarp. An organ enclosing sporangia and spores; as in group.
Marsilea. Isoetes, Azolla. and Salvinia. Tendril. A slender, twisting or clasping outgrowth by which
Sporophyll. A leaf bearing sporangia and spores; sometimes the plant climbs or clings for support; may be a modified
aggregated into a strobilus. stem, leaf, leaflet, or stipule.
Spur. A more or less elongate tubular or sac-like extension of Tepal. A term applied to the units of the perianth when the
some part of a flower, usually nectariferous. calyx and corolla are of similar form and not readily differen-
Stamen. The pollen-bearing organ of the flower. tiated.
Staminate. Pertaining to a flower that has stamens but no pis- Terete. Round in outline; circular in cross-section.
til; unisexual, with stamens. Terminal. At the end; apical; distal.
Staminode. A sterile stamen, or any structure resembling sta- Ternate. Borne in threes.
mens in the stamina I part of the flower; sometimes stami- Tetrad. A group of four structures forming a unit, as pollen or
nodes are petaloid. spores in tetrads.
Stellate. Star-shaped. Tetraploid. Having four full sets of chromosomes in each cell.
Sterile. Lacking functional reproductive organs. Thalloid. Resembling a thallus.
Stigma. The portion of a pistil which is receptive to pollen for Thallus. A vegetative body without differentiation into stem
effective fertilization. and leaf.
Stipe. A little stalk; a leaf stalk of a fern frond. Thorn. Spine; an internodal thorn is the same as a prickle.
Stipitate. Having a stipe. Throat. The opening into a sympetalous corolla or perianth;
Stipulate. Having stipules. the place where the corolla tube and limb join.
Stipule. Appendages at the base of a leaf, sometimes fused to Thyrse. A contracted cylindrical or ovoid and usually com-
the petiole. pact panicle.
Stolon. A stem which trails on the ground. often rooting at Thyrsoid. Pertaining to flowers or fruits arranged in or resem-
nodes; a runner; term often used to describe slender rhizomes bling a thyrse.
near the surface of the ground. Tomentose. Pertaining to a densely pubescent surface with
Stoloniferous. Producing stolons. matted, soft wooly hairs; densely wooly.
Stramineous. Straw-colored. Trailing. Prostrate but not rooting.
Striate. Marked with fine longitudinal lines or streaks. Translucent. Thin and membranous, such that light can be
Strigose. Having a surface covered with appressed sharp. transmitted through the tissue; partly transparent.
straight, and stiff hairs. Triadelphous. Having stamens which are united in three
Strobilus. A cone-like structure bearing the reproductive or- groups.
gans of one or both sexes, as in Equisetum. Lycopodium, and Trichome. Any hair-like outgrowth of the epidermis.
conifers. Trifoliate. Having three leaves.
Style. The tapered upper portion of a pistil. Trifoliolate. Pertaining to a compound leaf having three
Stylopodium. The disk-like expansion at the base of the style, leaflets.
as in Apiaceae. Trigonous. Three-angled.
Sub-. A prefix meaning under, or almost, or not quite. Tripinnate. Three times pinnately compound.
Subulate. Awl-shaped, narrow and tapering from base to Triploid. Having three full sets of chromosomes in each cell.
apex, often somewhat stiff. Tristylous. Pertaining to flowers of a given species having
362 Glossary of Plant Terms
three different style lengths and anthers borne at correspond- Valve. Each of the units resulting from the splitting of a
ing levels; in the Lythraceae, flowers with a long style having capsule.
midlength and short stamens, flowers with a midlength style Vascular. Pertaining to the presence of conductive tissues, xy-
have long and short stamens, and flowers with short styles lem and phloem, in a plant.
have long and midlength stamens. Veins. Threads of fibrovascular tissue in leaves and other
Triternate. Pertaining to leaves which are three times divided organs.
into threes; typically 27 leaflets; cf. Biternate. Velum. The membranous indusium or covering in Isoetes.
Truncate. Ending abruptly, appearing as if cut square across Velutinous. Velvety; covered with fine, soft, short, spreading
the end; pertaining to a base or apex that is nearly straight pubescence.
across. Venation. The arrangement of the veins.
Tuber. A thickened, short, subterranean stem; sometimes a Ventral. Pertaining to the lower, or abaxial. surface; cf.
short thickening on a root. Dorsal.
Tubercle. A swollen appendage, especially on the fruits of Verticil. A whorl.
some Cyperaceae; a small rounded protruding body. Verticillate. Having structures such as leaves, flowers, or
Tuberculate. Bearing tubercles. fruits arranged in whorls.
Tuberous. Bearing or producing tubers. Vestigial. Rudimentary.
Tubular. Descriptive of the corolla forming a slender tube. Villous. Covered with long, silky, straight hairs.
Turion. A winter bud; hibernaculum. Viscid. Sticky; glutinous.
Tussock. A dense tuft of grass or a grass-like plant. Viscidium. The sticky pad detaching with the pollinium in
Umbel. An inflorescence in which the peduncles and pedicels pollen transfer, as in the Orchidaceae.
radiate from the same level, often flat-topped; in the Apiaceae Weak. Not strong, such as some stems at the base.
the umbels are usually compound. Weedy. Troublesome or aggressive, intruding where not
Umbelliform. Pertaining to an inflorescence resembling an wanted.
umbel. Whorled. Arranged in a circle at one level on the stem.
Undulate. Having a wavy surface or margin. Wing. Any thin or membranous expansion bordering or sur-
Unisexual. Pertaining to flowers that are either staminate or rounding an organ.
pistillate; pertaining to plants bearing either staminate or pis- Winter bud. A small vegetative shoot which survives the win-
tillate flowers, but not both. ter and renews growth in spring; turion; hybernaculum; as
Urceolate. Urn-shaped; hollow and cylindrical or ovoid with a in Utricularia, MyriophyIIum, and Potamogeton.
constriction at or below the mouth. Zygomorphic. Descriptive of a flower or set of flower parts
Utricle. A small bladdery I-seeded fruit; any small bladder- which can be cut through the center into two equal and simi-
like body. lar parts along only one plane; bilaterally symmetrical; ir-
Valvate. Opening by valves; in buds, meeting by the edges regular.
without overlapping.
Glossary of Habitat Terms
This glossary provides definitions of habitat terms that may not Cedar. Thuja occidentalis. or Atlantic Coast Cedar. Chamaecy-
be in everyday usage. Definitions of habitats such as ponds. paris thyoides. See Swamp.
lakes. streams. brooks. temporary pools. and many others Cobble. An area characterized by numerous small to medium-
which are readily understood by everyone are not included sized stones.
here. It must be realized that some terms have regional mean- Conifer swamp. A swamp dominated by Tamarack. Larix lari-
ings. Thus. what is called a pond in Newfoundland is certainly cina. and/or Black Spruce. Picea mariana. See Swamp.
large enough to be called a lake anywhere else. Likewise the Cranberry bog. A bog managed as an agricultural area for
term "bog" in the South describes a somewhat different habitat the cultivation of cranberries. Vaccinium macrocarpon.
from that in the North. yet there are strong enough similarities Cypress swamp. A swamp with much standing water for the
(in acidity. poor drainage. brown waters. and some of the same growing season. and characterized by the deciduous conifer
species) that the term "bog" is readily understood by a north- Taxodium.
erner visiting a southern bog. Ditch. A manmade channel that is often seasonally wet.
Estuary. Shallow to deep saline tidal waters and adjacent sa-
Acid water. Water characterized by a pH value below 7. but line wetlands where river systems meet the ocean. Salinity
more often below a pH of 6. levels are lower than oceanic levels because of freshwater run-
Alkaline water. Water that is high in calcium carbonate con- off from the land and incoming rivers and streams.
tent. This water is also typically correlated with a pH above 7. Eutrophic water. Water that is high in nutrients. particularly
Alluvial plain. Flat. often broad. river margins with an accu- nitrates and phosphates.
mulation of silt caused by seasonal flooding. Also called a Fen. A peat-accumulating wetland that receives some of its nu-
floodplain. trients from the substrate and from runoff waters. It usually
Alluvial tidal freshwater shore. Flat shore with an accumu- supports marsh-like vegetation and shrubs.
lation of silty soil due to diurnal tidal flooding. caused by oce- Flats. Usually applied to broad. flat areas. especially behind
anic tides backing up the flow of freshwater. thus upstream dunes or along broad. sandy lake shores that arc occasionally
from the area of penetration of saltwater. flooded. and where the water table is close to the surface.
Alpine bog. A bog found above the timber line at high eleva- Floodplain. See Alluvial plain.
tions. or somewhat lower elevations at high latitudes. Fresh tidal flat. A mud flat that is subjected to fluctuation but
Bayou. A slow-moving creek or stream usually near the coast; is upstream of the area penetrated by saltwater.
primarily in the Gulf coastal plain region. Freshwater tidal marsh. A marshy area along a tidal river
Bog. In the North. an acid peat-accumulating wetland that flooded twice a day by freshwater. upstream of the area pene-
has no Significant inlets or outlets and supports various aci- trated by saltwater. and characterized by sedge vegetation.
dophilic plants. Sphagnum moss is typically present. and the Glade. An open area in a woods or forest. often wet and
habitat is nutrient poor. There is frequently a floating mat of marshy and dominated by sedges and grasses.
vegetation on the margin of an open bog pond. In the South. Impoundment. A manmade body of water. This is often sub-
bogs are open or semi-open areas characterized by grass and jected to great water-level fluctuations because of drawdown
sedge flora. and spring recharge.
Bottomland. Low land found adjacent to a river or stream Mangrove swamp. A tropical and subtropical forested wet-
that experiences periodic flooding. land characterized by the presence of tides and saltwater,
Brackish water. Non-coastal water with a moderate to high typically developing at the mouths of river systems.
concentration of dissolved inorganic salts; salty. Marl. Calcium carbonate (lime) that is precipitated by certain
Calcareous fen. Low marshy ground covered wholly or partly aquatic plants such as Cham and Potamogeton in the process
with standing water over a limestone bedrock. and character- of photosynthesis.
ized by lime-lOVing plants. See Fen. Marly sand. Damp to wet sand rich in marl.
Canal. A waterway originally dug for navigation. irrigation. or Marly wet panne. Barren marl flat that is covered with a de-
transporting water. posit of marl because of flooding and subsequent evaporation.
Canebrake. A thicket of Canegrass. Arundinaria gigantea; Marsh. A frequently or continuously inundated tract of land
found in the South. characterized by emersed. chiefly herbaceous wetland vege-
Cedar swamp. A swamp dominated by either Northern White tation.
363
364 Glossary of Habitat Terms
Moderately alkaline water. Here referring to water with a only from precipitation and capillary action; characteristic of
bicarbonate range of approximately 20-50 mg HCO, - litec 1 coastal plains of Newfoundland, the Maritimes, and eastern
or with a carbonate range of approximately 15-40 mg/l Maine.
CaCO,. Rill. A tiny stream, or rivulet.
Mossy talus. Wet, moss-covered area at the base of a cliff that Rivulet. A tiny stream.
is associated with springs or constant seeps. See Talus slope. Saline marsh. A coastal marsh influenced by salt water, with
Muck. Wet, heavy soil or well-composted peat. waterlogged soil during the growing season, and often cov-
Muskeg. A widely used term for peatlands of Canada and ered with up to 1 meter of water.
Alaska with the characteristics of northern bogs; especially Saline water. Salty water.
common in boreal and subarctic regions. Salt marsh. A coastal marsh flooded by tidal salt water, the
Panne (Pan). A highly saline depression or pool in a salt uppermost portions with waterlogged soil, but flooded only by
marsh. Also used to describe a wet depression associated with monthly spring tides.
sand dunes or sandy shores. Seep (seepage). An area of slow leaking or oozing water.
Peat barren. A large flat area dominated by a peaty substrate, Slough. A marshy, swampy area, very shallow lake system, or
appearing "barren" because of the absence of forest. sluggish channel with deep mud or accumulating peat; espe-
Peat. Soil produced by incomplete decay of sedges and grasses cially of the prairie regions.
or sphagnum moss or other vegetation. Sphagnum bog. A bog with a deep. often floating mat of
Peaty slope. Peatland that occupies shallow depressions on sphagnum moss. See Bog.
gentle gradients that are often associated with springs, seeps Spring. A flow of water from the ground; frequently serves as
of small brooks, or cold maritime coasts. a source of a stream.
Pine barren bog. A highly acidic boggy area along the Atlan- Swale. A natural, open, treeless hollow, meadow, or depression
tic and Gulf coastal plains consisting of pine flatlands domi- which is at least seasonally wet.
nated by such species as Pitch Pine, Pinus rigida, and Shortleaf Swamp. A wooded area with waterlogged soil, covered by sur-
Pine, Pinus palustris. face water much of the year. The term is often used loosely as
Pine savanna. An area of waterlogged substrates or inun- a synonym for "marsh"; thus the term "wooded swamp" is
dated with shallow surface water during seasons of heavy sometimes employed to clarify the forested or wooded nature
precipitation. This area may become quite dry during ex- of the area.
tended periods of droughts. Talus slope. Coarse rock debris formed at the base of a cliff or
Pineland. Land dominated by pines, especially along the a steep slope, caused chiefly by gravitational rock slides.
southern coastal plain. Thicket. An area of very dense growth of shrubs, usually
Pocosin. A broad, flat, evergreen shrub bog along the south- found along streams or on margins of lakes and ponds.
ern coastal plain that is generally water-logged, acidic, and Vernal pool. An area of standing water for a period in the
nutrient poor, with a sandy or peaty substrate. spring, but drying each summer.
Prairie. An area of the Midwest and Plains states dominated Waste area. Any area that is greatly disturbed by human ac-
by grasses growing on a thick, rich, loamy soil. Depressions tivities and is no longer actively used.
on rolling prairie land tend to be wet. Wet meadow. Low grassland often waterlogged and/or with
Quagmire. Very soft, muddy or boggy ground. the water table near the surface, but without standing water
Raised Bog. A large, generally flat area of peat, accumulated most of the year.
to a level above the original basin, that receives its water Wet montane soil. Wet soil at high elevation.
References
Ackerman. J. D. 1986. Mechanistic implications for pollination and light microscopy in Kansas funcus (Juncaceae). Brittonia
in the marine angiosperm Zostera marina. Aquatic Bot. 24: 3S: 201-209.
343-353. Browning. J. K.. D. Gordon-Gray. and S. G. Smith. 1995. Achene
Adams. P.. and R. K. Godfrey. 1961. Observations on the Sagitta- structure and taxonomy of North American Bolboschoenus
ria subulata complex. Rhodora 63: 247-266. (Cyperaceae). Brittonia 47: 433-445.
Aiken. S. G. 1986. The distinct morphology and germination of Bruederle. L. P.. and D. E. Fairbrothers. 1986. Allozyme varia-
the grains of two species of Wild Rice (Zizania. Poaceae). Ca- tion in populations of the Carex crinita complex (Cyperaceae).
nad. Field-Nat. 100: 237-240. Syst. Bot. 11: 583-594.
Aiken. S. G.. P. F. Lee. D. Punter. and J. M. Stewart. 1988. Wild Buell, M. F. 1935. Acorus calamus in America. Rhodora 37:
rice in Canada. Agric. Canad. Publ. 1830. 367-369.
Anderson. D. E. 1961. Taxonomy and distribution of the genus Campbell. C. S. 1983. Systematics of Andropogon virginicus com-
Phalaris. Iowa State J. Sci. 36: 1-96. plex (Gramineae). J. Arnold Arbor. 64: 171-254.
Anderson. E. 1928. The problem of species in Northern Blue Campbell. C. S. 1985. The subfamilies and tribes of Gramineae
Flags. Iris versicolor L. and Iris virginica L. Ann. Missouri Bot. (Poaceae) in the southeastern United States. J. Arnold Arbor.
Gard. 15: 241-332. 66: 123-199.
Anderson. E. 1936. The species problem in Iris. Ann. Missouri Case. F. w.. Jr. 1987. Orchids of the Western Great Lakes Region.
Bot. Gard. 23: 457-509. Rev. ed. Cranbrook Inst. Sci.. Bloomfield Hills. Mich.
Anderson. L. C. C. D. Zeis. and S. F. Alam. 1974. Phytogeogra- Case. F. w.. Jr.. and P. M. Catling. 1983. The genus Spiranthes in
phy and possible origins of Butomus in North America. Bull. Michigan. Michigan Bot. 22: 79-92.
Torrey Bot. Club 101: 292-296. Catling. P. M. 1981. Taxonomy of autumn-flowering Spiranthes
Beal, E. O. 1960a. Sparganium (Sparganiaceae) in the southeast- species of southern Nova Scotia. Canad. J. Bot. 59: 1253-
ern United States. Brittonia 12: 176-181. 1270.
Beal, E. O. 1960b. The Alismataceae of the Carolinas. J. Elisha Catling. P. M. 1982. Breeding systems of northeastern North
Mitchell Sci. Soc. 76: 68-79. American Spiranthes taxa (Orchidaceae). Canad. J. Bot. 60:
Beal, E. 0 .. J. W. Wooten. and R. B. Kaul. 1982. Review of the 3017-3039.
Sagitta ria engelmanniana complex (Alismataceae) with envi- Catling. P. M.. and I. Dobson. 1985. The biology of Canadian
ronmental correlations. Syst. Bot. 7: 417-482. weeds. 69. Potamogeton crispus L. Canad. J. PI. Sci. 65:
Beetle. A. A. 1947. Scirpus. North American Flora 18: 4S1-504. 655-668.
Bj6rkquist. I. 1968. Studies in Alisma L. II. Chromosome stud- Catling. P. M.. and W. G. Dore. 1982. Status and identification
ies. crossing experiments and taxonomy. Opera Bot. 19: of Hydrocharis morsus-ranae and Limnobium spongia (Hydro-
1-138. charitaceae) in northeastern North America. Rhodora 84:
Blake, S. F. 1912. The forms of Peltandra virginica. Rhodora 14: 523-545.
102-106. Catling. P. M., and Z. S. Porebski. 1995. The spread and current
Bogin. C. 1955. Revision of the genus Sagittaria (Alismataceae). distribution of European Frogbit. Hydrocharis morsus-ranae L..
Mem. New York Bot. Gard. 9: 179-233. in North America. Canad. Field-Nat. 109: 236-241.
Brackley. F. E. 1985. The orchids of New Hampshire. Rhodora Chambliss. C. E. 1940. The botany and history of Zizania aquat-
87: 1-117. ica L. (Wild Rice). J. Washington Acad. Sci. 30: 185-205. (Re-
Braun. E. L. 1967. The Vascular Flora of Ohio. Vol. 1. Monocot- printed in Annu. Rep. Smithsonian Inst., 1940.)
yledonae. Ohio State Univ. Press. Columbus. Ohio. Chase. A. 1964. First Book of Grasses. Smithsonian Inst. Press.
Brayshaw. T. C. 1985. Pondweeds and Bur-reeds. and Their Rel- Washington. D.C.
atives: Aquatic Families of Monocotyledons in British Colum- Church. C. L. 1949. A cytotaxonomic study of Glyceria and Puc-
bia. Occas. Pap. British Columbia Provo Mus. 26: 1-167. cinellia. Amer. J. Bot. 36: 155-165.
Britton. N. L. and A. Brown. 1896-1898. An Illustrated Flora Church. G. L. 1967. Taxonomic and generic relationships of
of the Northern United States. Canada. and the British Pos- eastern North American species of Elymus with setaceous
sessions. Charles Scribner's Sons. New York. [Vol. 1. 1896; glumes. Rhodora 69: 121-162.
Vol. 2. 1897; Vol. 3. 1898]. Clark. H. L.. and J. W. Thieret. ] 968. The duckweeds of Minne-
Brooks. R. E.. and C. Kuhn. 1986. Seed morphology under SEM sota. Michigan Bot. 7: 67-76.
365
Clausen, R. T. 1936. Studies in the genus Najas in northern Czerepanov, S. K. 1981. Plantae Vasculares URSS. Komarov
United States. Rhodora 38: 334345. Botanical Inst. Leningrad. (In Russian).
Clausen, R. T. 1937. A new species of Najas from the Hudson Daubs, E. H. 1962. The occurrence of Spirodela oligorrhiza in the
River. Rhodora 39: 5760. United States. Rhodora 64: 8385.
Clausen, R. T. 1952. Suggestion for the assignment of Torrey- Daubs, E. H. 1965. A monograph of Lemnaceae. Illinois Biol.
ochloa to Puccinellia. Rhodora 54: 4245. Monogr. 34:1118.
Clayton, W. D. 1968. The correct name of the Common Reed. Delahoussaye, A. J., and J. W. Thieret. 1967. Cyperus subgenus
Taxon 17:168169. Kyllinga (Cyperaceae) in the continental United States. Sida
Clements, S. E. 1990. Juneaceae (Rush Family) of New York 3: 128136.
State. Contr. to a Flora of New York State VII. New York State Dore, W G. 1947. Glyceria maxima in Canada. Canad. Field-Nat.
Mus. Bull. No. 475. 61: 174.
Cody, W. J. 1961. Iris pseudaeorus L, escaped from cultivation in Dore, W G. 1954. Frog-bit (Hydrocharis morsus-ranae L.) in the
Canada. Canad. Field-Nat. 75: 139142. Ottawa River. Canad. Field-Nat. 68: 18018 1.
Cook, C. D. K. 1985. Sparganium: Some old names and their Dore, W G. 1957. Wolffia in Canada. Canad. Field-Nat. 71:
types. Bot. Jahrb. Syst. 107: 269276. 1016.
Cook, C. D. K. 1988. Wind pollination in aquatic angiosperms. Dore, W G. 1968. Progress of the European Frog-bit in Canada.
Ann. Missouri Bot. Gard. 75: 768777. Canad. Field-Nat. 82: 7682.
Cook, C. D. K., and R. Liibnd. 1982. A revision of the genus Dore, W G. 1969. Wild Rice. Canad. Dept. Agric. Res. Publ.
Hydrocharis (Hydrocharitaceae). Aquatic Bot. 14: 177204. 1393: 184.
Cook, C. D. K., and M. S. Nicholls. 1986. A monographic study Dore, W. G., and J. McNeill. 1980. Grasses of Ontario. Agric. Ca-
of the genus Sparganium (Sparganiaceae). Part 1. Subgenus nad. Monogr. 26.
Xanthosparganium Holmberg. Bot. Helv. 96: 213267. Dressler, R. L. 1981. The Orchids. Natural History and Classifi-
Cook, C. D. K., and M. S. Nicholls. 1987. A monographic study cation. Harvard Univ. Press, Cambridge, Mass.
of the genus Sparganium (Sparganiaceae). Part 2. Subgenus Duvall, M. R., and D. D. Biesboer. 1988. Nonreciprocal hybrid-
Sparganium. Bot. Helv. 97: 144. ization failure in crosses between annual wild-rice species
Cook, C. D. K., and K. Urmi-Knig. 1983. A revision of the ge- (Zizania palustris Z. aquatica: Poaceae). Syst. Bot. 13: 229
nus Limnobium including Hydromystria (Hydrocharitaceae). 334.
Aquatic Bot. 17: 127. Eleuterius, L. N. 1975. The life history of the Salt Marsh Rush,
Cook, C. D. K., and K. Urmi-Knig. 1985. A revision of the ge- Juncus roemerianus. Bull. Torrey Bot. Club 102: 135140.
nus Elodea (Hydrocharitaceae). Aquatic Bot. 21: 111156. Fairey, J. E., III. 1967. The genus Scleria in the southeastern
Cope, T. A., and C. A. Stace. 1978. The Juncus bufonius L. aggre- United States. Castanea 32: 3771.
gate in western Europe. Watsonia 12: 113128. Fassett, N. C. 1924. A study of the genus Zizania. Rhodora 26:
Core, E. L. 1941. Butomus umbellatus in America. Ohio J. Sci. 153160.
41: 7985. Fassett, N. C. 1949. Some notes on Echinochloa. Rhodora 51: 13.
Correll, D. S. 1950. Native Orchids of North America. Stanford Fedchenko, B. A. 1934. Butomaceae. Pp. 228230 in V. L.
Univ. Press, Stanford, Calif. Komarov (ed.), Flora of the U.S.S.R. Vol. 1. (English transla-
Correll, D. S., and H. B. Correll. 19 72. Aquatic and Wetland tion, IPST Press, Jerusalem.)
Plants of Southwestern United States. U.S. Env. Protect. Fernald, M. L. 1903. Some variations in Triglochin maritima,
Agency, U.S. Gov. Print. Off., Washington, D.C. Rhodora 5: 174175.
Crins, W. J., and P. W. Ball. 1989a. Taxonomy of the Carex flava Fernald, M. L. 1918. The diagnostic character of Vallisneria
complex (Cyperaceae) in North America and northern Eur- americana. Rhodora 20: 108110.
asia. I. Numerical taxonomy and character analysis. Canad. Fernald, M. L. 1922. Notes on Sparganium. Rhodora 24: 2634.
J. Bot. 67: 10321047. Fernald, M. L. 1923. Notes on the distribution of Najas in north-
Crins, W. J. and P. W. Ball. 1989b. Taxonomy of the Carex flava eastern America. Rhodora 25: 105109.
complex (Cyperaceae) in North America and northern Eur- Fernald, M. L. 1932. The linear-leaved North American species
asia. II. Taxonomic treatment. Canad. J. Bot. 67: 10481065. of Potamogeton section Axillares. Mem. Amer. Acad. Arts 17:
Crow, G. E. 1993. Species diversity in aquatic angiosperms: 1183.
Latitudinal patterns. Aquat. Bot. 44: 229258. Fernald, M. L., and K. M. Wiegand. 1914. The genus Ruppia in
Crow, G. E. and C. B. Hellquist. 1981. Aquatic Vascular Plants eastern North America. Rhodora 16: 11912 7.
of New England. Part 2. Typhaceae and Sparganiaceae. New Ford, B. A., and P W. Ball. 1988. A reevaluation of the Trig-
Hampshire Agric. Exp. Sta. Bull. 517. lochin maritimum L. complex (Juncaginaceae) in eastern and
Crow, G. E. and C. B. Hellquist. 1982. Aquatic Vascular Plants central North America and Europe. Rhodora 90: 313337.
of New England. Part 4. Juncaginaceae, Scheuchzeriaceae, Freckmann, R. W, and D. M. Reed. 1979. Glyceria maxima, a
Butomaceae, Hydrochariotaceae. New Hampshire Agric. Exp. new, potentially troublesome wetland weed. Bull. Bot. Club
Sta. Bull. 520. Wisconsin 11: 3035.
Frieland, S. 1941. The American species of Hemicarpha. Amer. Harms, V L., D. E Hooper, and L. Baker. 1986. Plantago maritima
J. Bot. 28: 855861. and Carex mackenziei new for Saskatchewan: Additional rare
Fuller, A. M. 1933. Studies on the flora of Wisconsin. Part I. The inland stations for two seacoast salt marsh species. Rhodora
orchids; Orchidaceae. Bull. Pub. Mus. Milwaukee 14: 1284. 88: 315323.
Gaiser, L. 0. 1949. Further distribution of Butomus umbellatus Hartog, C. den, and E van der Plas. 1970. A synopsis of the
in the Great Lakes region. Rhodora 51: 387390. Lemnaceae. Blumea 18: 355368.
Gale, S. 1944. Rhynchospora, section Eurhynchospora, in Canada, Haynes, R. R. 1974. A revision of North American Potamoge-
the United States and the West Indies. Rhodora 46: 89134, ton subsection Pusilli (Potamogetonaceae). Rhodora 76:
159197, 207249, 255278. 564649.
Gleason, H. A. 1952. The New Britton and Brown Illustrated Haynes, R. R. 1978. The Potamogetonaceae in the southeast-
Flora of the Northeastern United States and Adjacent Can- ern United States. J. Arnold Arbor. 59: 170191.
ada. 3 vols. Lancaster, Press, Lancaster, Pa. Haynes, R. R. 1979. Revision of North and Central America
Godfrey, R. K., and J. W. Wooten. 1979. Aquatic and Wetland Najas (Najadaceae). Sida 8: 3456.
Plants of Southeastern United States. Monocotyledons. Univ Haynes, R. R. 1980. Aquatic and marsh plants of Alabama. I. -
Georgia Press, Athens, Ga. Alismatidae. Castanea 45: 3151.
Gomez, L. D. 1984. Las plantas acuticas y anfibias de Costa Haynes, R. R. 1985. A revision of the clasping-leaved Potamoge-
Rica y Centroamrica. 1. Liliopsida. Editorial Universidad ton (Potamogetonaceae). Sida 11: 173188.
Estatal a Distancia, San Jose, Costa Rica. Haynes, R. R. 1988. Reproductive biology of selected aquatic
Gould, F. W. 1975. The Grasses of Texas. Texas A & M Univ. plants. Ann. Missouri Bot. Gard. 75: 805810.
Press, College Station, Tex. Haynes, R. R., and C. B. Hellquist. 1996. New combinations in
Gould, E W, and C. A. Clark. 1978. Dichanthelium (Poaceae) in North American Alismatidae. Novon 6: 370371.
the United States and Canada. Ann. Missouri Bot. Gard. 65: Haynes, R. R. and C. B. Hellquist. 2000. Potamogetonaceae.
10881132. Pp. 4774 in: Flora of North America North of Mexico. Vol.
Gould, E W, and R. B. Shaw. 1983. Grass Systematics. 2nd ed. 22. Convening ed. N. T. Morin and Editorial Committee.
Texas A & M Univ. Press, College Station, Tex. Oxford Univ. Press, New York.
Gould, E W, M. A. Ali, and D. E. Fairbrothers. 19 72. A revision Haynes, R. R., and L. B. Holm-Nielsen. 1987. The Zannichellia-
of Echinochloa in the United States. Amer. Midl. Nat. 87: ceae in the southeastern United States. J. Arnold Arbor. 68:
3659. 259268.
Grayum, M. H. 1987. A summary of evidence and arguments Haynes, R. R., and D.C. Williams. 1975. Evidence for the hy-
supporting removal of Acorns from the Araceae. Taxon 36: brid origin of Potamogeton longiligulatus (Potamogetonaceae).
723729. Michigan Bot. 14: 94100.
Grear, J. W, Jr. 1966. Cytogeography of Orontium aquaticum. Hellquist, C. B. 1980. Correlation of alkalinity and the distri-
Rhodora 68: 2534. bution of Potamogeton in New England. Rhodora 82: 331
Greene, C. W 1980. The systematics of Calamagrostis (Grami- 344.
neae) in eastern North America. Ph.D. diss., Harvard Univ., Hellquist, C. B. 1984. Observations of Potamogeton hillii Morong
Cambridge, Mass. in North America. Rhodora 86: 101111.
Greene, C. W 1984. Sexual and apomictic reproduction in Cala- Hellquist, C. B. 1993. Taxonomic considerations in aquatic veg-
magrostis (Gramineae) from eastern North America. Amer. J. etation assessments. Lake and Reserv. Managem. 7: 175183.
Bot. 71: 285293. Hellquist, C. B., and G. E. Crow. 1980. Aquatic Vascular Plants
Greene, C. W 1987. Calamagrostis pickeringii in Maine. Rhodora of New England. Part 1. Zosteraceae, Potamogetonaceae,
89: 333336. Zannichelliaceae, Najadaceae. New Hampshire Agric. Exp.
Gutteridge, R. L. 1954. Glyceria maxima on the Mississippi River, Sta. Bull. 515.
Ontario, 1953. Canad. Field-Nat. 68: 133135. Hellquist, C. B., and G. E. Crow. 1981. Aquatic Vascular Plants
Hagstr6m, J. 0. 1916. Critical researches on the Potamogetons. of New England. Part 3. Alismataceae. New Hampshire
Kungl. Svenska Vetenskapskad. Handl. 55: 1281. Agric. Exp. Sta. Bull. 518.
Hmet-Ahti, L. 1980. Juncus alpinoarticulatus: The legitimate Hellquist, C. B., and G. E. Crow. 1982. Aquatic Vascular Plants
name for Juncus alpinus. Ann. Bot. Fenn. 17: 341342. of New England. Part 5. Araceae, Lemnaceae, Xyridaceae, Er-
Hmet-Ahti, L. 1986. North American races of Juncus alpinoarti- iocaulaceae, and Pontederiaceae. New Hampshire Agric. Exp.
culatus (Juncaceae). Ann. Bot. Fenn. 23: 277281. Sta. Bull. 523.
Harms, V L. 1973. Taxonomic studies of North American Spar- Hellquist, C. B., and G. E. Crow. 1986. Potamogeton haynesii
ganium. I. S. hyperboreum and S. minimum. Canad. J. Bot. 51: (Potamogetonaceae), a new species from northeastern North
16291641. America. Brittonia 38: 415419.
Harms, V L., and G. E Ledingham. 1986. The Narrow-leaved Hellquist, C. B., and R. L. Hilton. 1983. A new species of
Cat-tail, T. glauca, newly reported from Saskatchewan. Ca- Potamogeton (Potamogetonaceae) from northeastern United
nad. Field-Nat. 100: 107110. States. Syst. Bot. 8: 8692.
Hellquist, C. B., C. T. Philbrick, and R. L. Hilton. 1988. The tax- Kral, R. 1983. The Xyridaceae in the southeastern United
onomic status of Potamogeton lateralis Morong (Potamoget- States. J. Arnold Arbor. 64: 421429.
onaceae). Rhodora 90: 1519. Kral, R. 1988. The genus Xyris (Xyridaceae) in Venezuela and
Hendricks, A. J. 1957. A revision of the genus Alisma (Dill.) L. contiguous northern South America. Ann. Missouri Bot.
Amer. Midl. Nat. 58: 470493. Gard. 75: 522722.
Henry, L. K., W. E. Buker, and D. L. Pearth. 1975. Western Kugel, A. R. 1958. Variation in the Spiraea alba-latifolia com-
Pennsylvania orchids. Castanea 40: 93168. plex. Ph.D. diss., Univ of Michigan, Ann Arbor, Mich.
Hermann, E J. 1975. Manual of the Rushes (Juncus spp.) of the La Frankie, J. V, Jr. 1986a. Morphology and taxonomy of the
Rocky Mountains and Colorado Basin. U.S.D.A. Forest Serv. New World species of Maianthemum (Liliaceae). J. Arnold
Gen. Tech. Rep. RM-18. Arbor. 67: 371439.
Hess, W J. 1986. Wolffia papulifera Thompson (Lemnaceae), new La Frankie, J. V, Jr. 1986b. Transfer of the species of Smila-
to Michigan. Sida 11: 407411. cina Desf. to Maianthemum Wigg. (Liliaceae). Taxon 35:
Hillman, W S. 1961. The Lemnaceae, or duckweeds: A review 584589.
of the descriptive and experimental literature. Bot. Rev. 27: Lakela, 0. 1941. Sparganium glomeratum in Minnesota. Rhodora
221287. 43: 8385.
Hillman, W S., and D. D. Culley, Jr. 1978. The uses of duck- Landolt, E. 1980. Key to determination of taxa within the fam-
weeds, Amer. Sci. 66: 442451. ily of Lemnaceae. Veroff. Geobot. Inst. ETH Stiftung Rbel,
Hitchcock, A. S. 1950. Manual of the Grasses of the United Zurich 70: 1321.
States. 2nd ed., revised by A. Chase. U.S.D.A. Misc. Publ. 200. Landolt, E. 1981. Distribution pattern of the family Lemnaceae
U.S. Gov. Print. Off., Washington, D.C. in North Carolina. Veroff. Geobot. Inst. ETH Stiftung Rbel,
Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. Zurich 77: 112148.
19551969. Vascular Plants of the Pacific Northwest. 5 Landolt, E. 1986. The family of Lemnaceaea monographic
parts. Univ. Washington Press, Seattle, Wash. study. (Vol. 1.) Veroff. Geobot. Inst. ETH Stiftung Rbel, Zu-
Holub, J. 1977. Stuckenia Burner 1912the correct name for Co- rich 71: 1566.
leogeton (Potamogetonaceae). Preslia, Praha 69: 361366. Lee, D. W. 1975. Population variation and introgression in
Horn, C. N. 1983. The annual growth cycle of Heteranthera North American Typha. Taxon. 24: 633641.
dubia in Ohio. Michigan Bot. 23: 2934. Lelong, M. G. 1984. New combinations for Panicum subgenus
Horn, C. N. 1986. Typifications and a new combination in Het- Panicum and subgenus Dichanthelium (Poaceae) of the south-
eranthera (Pontederiaceae). Phytologia 59: 290. eastern United States. Brittonia 36: 262273.
Hotchkiss, N., and H. L. Dozier. 1949. Taxonomy and distribu- Les, D. H. 1983. Taxonomic implications of aneuploidy and
tion of North America cattails. Amer. Midi. Nat. 41: 237 polyploidy in Potamogeton (Potamogetonaceae). Rhodora 85:
254. 301323.
Hultn, E. 1950. Flora of Alaska and Yukon. X. Lunds Univ Les, D. H., and R. R. Haynes. 1996. Coleogeton (Potamogetona-
Arsskrift N. E, Sweden, Avd. 2, Bd. 46. ceae), a new genus of pondweeds. Novon 6: 389391.
Hunt, G. S., and R. W Lutz. 1959. Seed production by Curly- Les, D. H., and D. J. Sheridan. 1990a. Hagstroms concept of
leaved Pondweed and its significance to waterfowl. J. Wildl. phylogenetic relationships in Potamogeton L. (Potamogetona-
Managem. 23: 405408. ceae). Taxon 39: 4158.
Jacobs, D. L. 1947. An ecological life history of Spirodela poly- Les, D. H., and D. J. Sheridan. 1990b. Biochemical heterophily
rhiza. Ecol. Monogr. 17: 437469. and flavonoid evolution in North American Potamogeton.
Kerguelen, M., G. Bosc., and J. Lambinon. 1987. Donnes taxo- Amer. J. Bot. 77: 458465.
nomiques, nomenclaturales et chorologiques pour une rvi- Les, D. H., M. H. Cleland, and C. T. Philbrick. 1995. Taxonomic
sion de la flore de France. Lejeunia, n.s., 120: 1263. realignments in the Potamogetonaceae: evidence from mo-
Kessler, J. W 1988. A treatment of Scleria (Cyperaceae) for lecular data. Abstract. Amer. J. Bot. 82: 144.
North America north of Mexico. Sida 12: 391407. Les, D. H., L. J. Mehrhoff, M. A. Cleland, and J. D. Gabel. 1997.
Koyama, T. 1962. The genus Scirpus Linn., some North Ameri- Hydrilla verticillata (Hydrocharitaceae) in Connecticut. J.
can aphylloid species. Canad. J. Bot. 40: 913937. Aquat. Pl. Manag. 35: 1014.
Koyama, T. 1963. The genus Scirpus Linn., critical species of the Lve, A., and D. Lve. 1957. Drug content and polyploidy in
section Pterolepis. Canad. J. Bot. 40: 913937. Acorus. Proc. Genet. Soc. Canad. 2: 1417.
Kral, R. 1966a. Eriocaulaceae of continental North America Lve, A., and D. Lve. 1958. Biosystematics of Triglochin mari-
north of Mexico. Sida 2: 285332. timum agg. Naturaliste Canad. 85: 156165.
Kral, R. 1966b. Xyris (Xyridaceae) of the continental United Lve, D., and H. Lieth. 1961. Triglochin gaspense, a new species
States and Canada. Sida 2: 177260. of Arrow Grass. Canad. J. Bot. 39: 12611272.
Kral, R. 1971. A treatment of Abildgaardia, Bulbostylis and Fim- Lowden, R. M. 1973. Revision of the genus Pontederia L. Rho-
bristylis (Cyperaceae) for North America. Sida 4: 57227. dora 75: 426487.
Kral, R. 1978. A synopsis of Fuirena (Cyperaceae) for the Ameri- Lowden, R. M. 1982. An approach to the taxonomy of Vallis-
cas north of South America. Sida 7: 309354. neria (Hydrocharitaceae). Aquatic Bot. 13: 269298.
Lowden, R. M. 1992. Floral variation and taxonomy of Limnob- Padgett, D. J. and G. E. Crow. 1994a. Foreign plant stock: con-
ium L. C. Richard (Hydrocharitaceae) Rhodora 94: 111134. cerns for wetland mitigation. Rest. Managem. Notes 12: 168
Luer, C. A. 1975. The Native Orchids of the United States and 171.
Canada Excluding Florida. New York Botanical Garden, Padgett, D. J. and G. E. Crow. 1994b. A vegetation and floristic
Bronx, N.Y. analysis of a created wetland in southeastern New Hamp-
McClure, E. A. 1973. Genera of Bamboos Native to the New shire. Rhodora 96: 129.
World (Gramineae: Bambusoideae). Smithsonian Contr. Bot. Penfound, W. T., and T. T. Earle. 1948. The biology of the Water
No. 9. Hyacinth. Ecol. Monogr. 18: 447472.
McDonnell, M. J., and G. E. Crow. 1979. The typification and Philbrick, C. T. 1983. Aspects of floral biology in three species
taxonomic status of Spartina caespitosa A. A. Eaton. Rhodora of Potamogeton (Pondweeds). Michigan Bot. 23: 3538.
81: 123129. Philbrick, C. T. 1988. Evolution of underwater outcrossing from
Mackenzie, K. K. 19311935. Cyperaceae. North American aerial pollination systems: A hypothesis. Ann. Missouri Bot.
Flora 18: 1478. Gard. 75: 836841.
Mackenzie, K. K. 1940. North American Cariceae. 2 vols. New Pohl, R. W. 1980. Flora Costaricensis. Family no. 15, Grami-
York Botanical Garden, Bronx, N.Y. neae. Fieldiana, Bot. Ser., n.s., 4: 1608.
Malme, G. O. K. 1937. Xyridaceae. North American Flora 19: Pyrah, G. L. 1969. Taxonomic and distributional studies in
315. Leersia (Gramineae). Iowa State J. Sci. 44: 215270.
Marie-Victorin, F. 1931. LAnacharis canadensis. Histoire et so- Raymond, M. 1951. Two new Eriophorum hybrids from north-
lution dun imbroglio taxonomique. Contr. Lab. Bot. Univ. eastern North America. Sv. Bot. Tidsk. 15: 523531.
Montral 18: 143. Raymond, M. 1958. Additional notes on some S.E. Asiatic Scir-
Marie-Victorin, F. 1943. Les Vallisneries americaines. Contr. pus. Nat. Canad. 86: 225242.
Inst. Bot. Univ. Montral 46: 138. Reed, C. F. 1970. Selected Weeds of the United States. U.S.D.A.
Mason, H. L. 1957. A Flora of the Marshes of California. Univ. Agric. Res. Ser., Agric. Handb. 336.
California Press, Berkeley, Calif. Reeder, J. R. 1953. Affinities of the grass genus Beckmannia
Merilinen, J. 1968. Najas minor All. in North America. Rho- Host. Bull. Torrey Bot. Club 80: 187196.
dora 70: 161175. Reinking, M. 1981. Juncus stuckeyi (Juncaceae), a natural hy-
Miller, M., and E. O. Beal. 1972. Scirpus validus and S. acutus brid from northern Ohio. Brittonia 33: 170178.
a question of distinctness. J. Minnesota Acad. Sci. 38: 2123. Reveal, J. L. 1970. Sparganum simplex Huds., a superfluous
Mitchell, R. S., and C. J. Sheviak. 1982. Rare Plants of New York name. Taxon 19: 796797.
State. New York State Mus. Bull. No. 445. Reveal, J. L. 1990. The neotypification of Lemna minuta Humb.,
Mobberly, D. G. 1956. The taxonomy and distribution of the ge- Bonpl. & Kunth, an earlier name for Lemna minuscula Herter
nus Spartina. Iowa State J. Sci. 30: 471574. (Lemnaceae). Taxon 39: 328330.
Moldenke, H. N. 1937. Eriocaulaceae. North American Flora Reznicek, A. A. 1990. Evolution in sedges (Carex, Cyperaceae).
19: 1750. Canad. J. Bot. 68: 14091432.
Moore, E. 1915. The Potamogetons in relation to pond culture. Reznicek, A. A., and P. W. Ball. 1974. The taxonomy of Carex
Bull. Bur. Fisheries 33: 251291. series Lupulinae in Canada. Canad. J. Bot. 52: 23872399.
Morris, F., and E. E. Eames. 1929. Our Wild Orchids. Charles Reznicek, A. A., and P. W. Ball. 1980. The taxonomy of Carex
Scribners Sons, New York. section Stellulate in North America north of Mexico. Contr.
Muenscher, W. C. 1936. The germination of seeds of Potamo- Univ. Michigan Herb. 14: 153203.
geton. Ann. Bot. (London) 50: 805821. Reznicek, A. A., and R. S. W. Bobbette. 1976. The taxonomy of
Naczi, R. F. C., R. J. Driskill, E. L. Pennell, N. E. Seyfried, A. O. Potamogeton subsection Hybridi in North America. Rhodora
Tucker, and N. H. Dill. 1986. New records of some rare Del- 78: 650673.
MarVa sedges. Bartonia 52: 4957. Reznicek, A. A., and P. M. Catling. 1986. Carex striata, the cor-
Netherland, M. D. 1997. Turion ecology of Hydrilla. J. Aquat. rect name for C. walteriana (Cyperaceae). Rhodora 88: 405
Plant Manage. 35: 110. 406.
Ogden, E. C. 1943. The broadleaved species of Potamogeton of Rhoades, R. W. 1962. The aquatic form of Alisma subcordatum
North America north of Mexico. Rhodora 45: 57105, 119 Rat Rhodora 64: 227229.
163, 171214. Richardson, F. D. 1980. Ecology of Ruppia maritima L. in New
Padgett, D. J. and G. E. Crow. 1993a. Some unwelcome additions Hampshire (U.S.A.) tidal marshes. Rhodora 82: 403439.
to the flora of New Hampshire. Rhodora 95: 348351 Richardson, F. D. 1983. Variation, adaptation and reproductive
Padgett, D. J. and G. E. Crow. 1993b. A comparison of floristic biology in Ruppia maritima L. populations from New Hamp-
composition and species richness within and between cre- shire coastal and estuarine tidal marshes. Ph.D. diss., Univ. of
ated and natural wetlands of southeastern New Hampshire. New Hampshire, Durham, N.H.
Pp. 171186 in F. J. Webb, Jr., ed. Proceedings of the Twen- Roberts, M. L. 1972. Butomus umbellatus in the Mississippi
tieth Annual Conference on Wetlands and Restoration and watershed. Castanea 37: 8385.
Creation. Hillsborough Community College. Tampa, Fla. Roberts, M. L., R. L. Stuckey, and R. S. Mitchell. 1981. Hydro-
370 References
charis morsus-ranae (Hydrocharitaceae): New to the United Gunckel (ed.), Current Topics in Plant Science. Academic
States. Rhodora 83: 147-148. Press. New York.
Rominger. J. M. 1962. Taxonomy of Setaria (Gramineae) in Smith. S. G. 1986. The cattails (Typha): Interspecific ecological
North America. Illinois BioI. Monogr. 29: 1-132. differences and problems of identification. Lake and Reserv.
Rosatti. T. J. 1987. The genera of Pontederiaceae in the south- Managem. 2: 13-16.
eastern United States. J. Arnold Arbor. 68: 35-71. Smith. S. G. 1987. Typha: Its taxonomy and the ecological sig-
st. John. H. 1920. The genus Elodea in New England. Rhodora nificance of hybrids. Arch. Hydrobiol. 27: 129-138.
22: 17-29. Smith. S. G. 1995. New combinations in North American
st. John. H. 1965. Monograph of the genus Elodea. Part 4. Sum- Schoenoplectus. Bolboschoenus. Isolepis. and Trichophorum
mary. Rhodora 67: 1-35. 155-180. (Cyperaceae). Novon 5: 97-102.
Schrenk. W. J. 1978. North American Platantheras: Evolution Smith. S. G. in press. Schoenoplectus. In: Flora of North America
in the making. Amer. Orchid Soc. Bull. 47: 429-437. North of Mexico. Vol. 23. Convening ed. N. T. Morin and Edi-
Schuyler. A. E. 1962a. A new species of Scirpus in the north- torial Committee. Oxford Univ. Press. New York.
eastern United States. Rhodora 64: 43-49. Snogerup. S. 1963. Studies in the genus Juncus. III. Observa-
Schuyler. A. E. 1962b. Sporadic culm formation in Scirpus lon- tions on the diversity of chromosome numbers. Bot. Not.
gii. Bartonia 32: 1-5. 116: 142-156.
Schuyler. A. E. 1964a. Notes on five species of Scirpus in eastern Snogerup. S. 1980. Juncus. pp. 102-111 in Tutin et al.. Flora
North America. Bartonia 33: 1-6. Europaea. Vol. 1. Cambridge Univ. Press. Cambridge.
Schuyler. A. E. 1964b. A biosystematic study of the Scirpus Soderstrom. T. R. K. W. Hilu. C. S. Campbell. and M. E. Bark-
cyperinus complex. Proc. Acad. Nat. Sci. Philadelphia 115: worth (eds.). 1987. Grass Systematics and Evolution. Smith-
283-311. sonian Inst. Press. Washington. D.C.
Schuyler. A. E. 1966. The taxonomic delineation of Scirpus lin- Standley, L. A. 1983. A clarification of the status of Carex crinita
eatus and Scirpus pendulus. Notul. Nat. Acad. Nat. Sci. Phila- and C. gynandra (Cyperaceae). Rhodora 85: 229-241.
delphia 390: 1-3. Standley, L. A. 1987a. Anatomical and chromosomal studies of
Schuyler. A. E. 1967. A taxonomic revision of the North Ameri- Carex section Phacocystis in eastern North America. Bot. Gaz.
can leafy species of Scirpus. Proc. Acad. Nat. Sci. Philadelphia (Crawfordsville) 148: 507-518.
119: 295-323. Standley, L. A. 1987b. Taxonomy of the Carex lenticularis com-
Schuyler. A. E. 1968. A new status for an eastern North Ameri- plex in eastern North America. Canad. J. Bot. 65: 673-686.
can Scirpus. Rhodora 69: 198-202. Staniforth, R. J., and K. A. Frego. 1980. Flowering Rush (Buto-
Schuyler, A. E. 197 4a. Typification and application of the mus umbellatus) in the Canadian prairies. Canad. Field-Nat.
names Scirpus americanus Pers., S. olneyi Gray, and S. pungens 94: 333-336.
Vahl. Rhodora 76: 51-52. Stephenson. S. N. 1984. The genus Dichanthelium (Poaceae) in
Schuyler, A. E. 1974b. Scirpus cylindricus: An ecologically re- Michigan. Michigan Bot. 23: 107-119.
stricted eastern North American tuberous bulrush. Bartonia Stoutamire, W. P. 1974. Relationships of the Purple-fringed Or-
43: 29-37. chis Platanthera psycodes and P. grandiflora. Brittonia 26:
Setchell, W. A. 1929. Morphological and phenological notes on 42-58.
Zostera marina 1. Univ. California Publ. Bot. 14: 389-452. Stuckey, R. L. 1968. Distributional history of Butomus umbella-
Setchell, W. A. 1933. A preliminary survey of the species of tus (Flowering Rush) in the western Lake Erie and Lake st.
Zostera. Proc. Natl. Acad. U.S.A. 19: 810-817. Clair region. Michigan Bot. 7: 134-142.
Shaffer-Fehre, M. 1991. The position of Najas within the Alis- Stuckey. R. L. 1979. Distributional history of Potamogeton crispus
matidae (Monocotyledones) in the light of new evidence from (Curly Pondweed) in North America. Bartonia 46: 22-42.
seed coat structures in the Hydrocharitoideae (Hydrocharita- Stuckey. R. 1. 1980. The migration and establishment of Juncus
ceae). Bot. J. Linn. Soc. 107: 189-209. gerardii (Juncaceae) in the interior of North America. Sida 8:
Sheviak, C. J. 1974. An Introduction to the Ecology of the Illi- 334-347.
nois Orchidaceae. Illinois State Mus. Sci. Pap. 45. Stuckey, R. 1. 1981. Distributional history of Juncus compressus
Sheviak. C. J. 1982. Biosystematic Study ofthe Spiranthes cernua (Juncaceae) in North America. Canad. Field-Nat. 95: 167-
Complex. New York State Mus. Bull. No. 448. 171.
Smith. J. G. 1895. Revision of the North American species of Stuckey. R.1. 1985. Distributional history of Najas marina
Sagitta ria and Lophotocarpus. Annu. Rep. Missouri Bot. Gard. (Spiny Naiad) in North America. Bartonia 51: 2-16.
6: 27-64. Svenson. H. K. 1932. Monographic studies in the genus Eleo-
Smith. S. G. 1967. Experimental and natural hybrids in North charis-II. Rhodora 34: 192-203. 215-227.
American Typha (Typhaceae). Amer. MidI. Nat. 78: 257-287. Svenson. H. K. 1957. Cyperaceae. Tribe 2. Scirpeae. North
Smith. S. G. 1969. Natural hybridization in the Scirpus lacustris American Flora 18: 505-556.
complex in north central United States. Pp. 175-200 in J. E. Terrell. E. E.. P. M. Peterson. J. 1. Reveal, and M. R. Duvall.
References 371
1997. Taxonomy of North American species of Ziziana Wentz, W. A., and R. L. Stuckey. 1971. The changing distribu-
(Poaceae). Sida 17: 533-549. tion of the genus Najas (Najadaceae) in Ohio. Ohio J. Sci. 71:
Thieret, J. W. 1971. Observations on some aquatic plants in 292-302.
northern Minnesota. Michigan Bot. 10: 117-124. Whiting, R. E., andP. M. Catling. 1986. Orchids of Ontario. Cana-
Thomas, W. W. 1984. The systematics of Rhynchospora section ColI Foundation, Ottawa.
Dichromena. Mem. New York Bot. Gard. 37: 1-116. Wiegand, K. M. 1921. The genus Echinochloa in North America.
Tucker, G. C. 1983. The taxonomy of Cyperus (Cyperaceae) in Rhodora 23: 49-65.
Costa Rica and Panama. Syst. Bot. Monogr. 2: 1-85. Wilson, K. A. 1960. Genera of Arales in the southeast United
Tucker, G. C. 1984. A revision of the genus Kyllinga Rottb. (Cyp- States. J. Arnold Arbor. 41: 47-72.
eraceae) in Mexico and Central America. Rhodora 86: 507- Wohler, J. R., 1. M. Wohler, and R. T. Hartman. 1965. The oc-
538. currence of Spirodela oligorrhiza in western Pennsylvania. Cas-
Tucker, G. C. 1985. Cyperus j1avicomus, the correct name for tanea 30: 230-23l.
Cyperus albomarginatus. Rhodora 87: 539-541. Wooten, J. W. 1970. Experimental investigations of the Sagitta-
Tucker, G. C. 1987. The genera of Cyperaceae in the southeast- ria graminea complex: Transplant studies and genecology.
ern United States. J. Arnold Arbor. 68: 361-445. J. Ecol. 58: 233-242.
Tucker, G. C. 1992. Scirpus (Cyperaceae) in Connecticut. Wooten, J. W. 1971. The monoecious and dioecious conditions
Newslett. Connecticut Bot. Soc. 20: 3-10. in Sagitta ria latifolia L. (Alismataceae). Evolution 25: 549-
Tucker, G. C.. and N. G. Miller. 1990. Achene microstructure in 553.
Eriophorum (Cyperaceae); taxonomic implications and paleo- Wooten, J. W. 1973a. Edaphic factors in species and ecotype dif-
botanical applications. Bull. Torrey Bot. Club 117: 266-283. ferentiation of Sagitta ria. J. Ecol. 61: 151-156.
Urbanska-Worythiewicz, K. 1975. Cytological variation within Wooten, J. W. 1973b. Taxonomy of seven species of Sagittaria
Lemna L. Aquatic Bot. 1: 377-394. from eastern North America. Brittonia 25: 64-74.
Voss, E. G. 1966. Nomenclatural notes on Monocots. Rhodora Wylie, R. B. 1917. The pollination of Vallisneria spiralis. Bot.
68: 435-463. Gaz. (Crawfordsville) 63: 135-145.
Voss, E. G. 1972. Michigan Flora. Part 1. Gymnosperms andMo- Yeo, R. R. 1964. Life history of Common Cattail. Weeds 12:
nocots. Cranbrook lnst. Sci. Bull. 55, Bloomfield Hills, Mich. 284-288.
Warwick, S. 1., and S. G. Aiken. 1986. Electrophoretic evidence Yeo, R. R. 1965. Life history of Sago Pondweed. Weeds 13:
for the recognition of two species in annual Wild Rice (Zi- 314-32l.
zania, Poaceae). Syst. Bot. 11: 464-473. Yeo, R. R., R. H. Falk, and J. R. Thurston. 1984. The morphol-
Webster, R. D. 1988. Genera of the North American Paniceae ogy of hydrilla (Hydrilla verticillata (L.f.) Royle). J. Aquatic PI.
(Poaceae: Panicoideae). Syst. Bot. 13: 576-609. Managem. 22: 1-16.
Wehrmeister, J. R. 1978. An ecological life history of the pond- Zenkert, C. A. 1960. The Old-World flowering rush: An attrac-
weed Potamogeton crispus 1. in North America. M.S. thesis, tive and aggressive immigrant. Science on the March, Mag.
Ohio State Univ., Columbus, Ohio. (Reprinted, CLEAR Tech. Buffalo Mus. Sci. 40: 71-75.
Rep. No. 99.)
Wehrmeister, J. R., and R. L. Stuckey. 1992. Life history of Pota-
mogeton crispus. Michigan Bot. 31: 3-16.
Index
Indexed terms set in italics are in synonymy; those in capital letters are family names. Page numbers set in boldface refer to treat-
ments of taxonomical categories. Boldface italic numerals indicate the major illustration of a plant species. Numerals set in italics
refer to minor illustrative appearances. and those in roman to incidental textual references to a species.
373
374 Index
ssp. angustifolia, 1:153 var. australis, 2: 151 pseudocyperus, 2:126. 151, 152
ssp. palustris, 1: 15 3 var. folliculata, 2:151 rariflora. 2:127,151,161
var. palustris, 1:153 garberi, 2:123, 130, 112 recta, 2:127
rotundifolia, 1:148, 150 gigantea, 2:168 retrorsa. 2:168
Cardinal Flower, 1: 3 6 6 glaucescens, 2:168 rhynchophysa, 2:151
Carduus muticus, 1:414 grayana, 2: 151 richii.2:141
Carex, 2:121 gynandra, 2:123, 130, 112 riparia.
alata, 2:125, 141, 141 gynocrates, 2:168 var. impressa, 2:151
albolutescens, 2:168 heleonastes. 2:168 var. lacustris. 2:141
alopecoidea, 2:124, 130, 116 hormathodes, 2:125, 141, 141 rostrata, 2:126, 151
angustior, 2:141 howei.2:141 var. utricu/ata, 2:151
var. gracilenta, 2:141 hyalinolepis, 2:126, 151, 152 ruthii,2:168
aquatilis, 2:130 hystericina. 2:126,151,155 salina, 2:123,127.128
var. altior, 2:123, 130, 111 impressa. 2:151 var. kattegatensis. 2:127
var. aquatilis, 2:123, 130, 111 incomperta.2:141 var. pseudofilipendula. 2: 12 7
var. substricta, 2:130 interior. 2:125, 141, 147 sartwellii, 2:124, 130, 116
atherodes, 2:126, 141, 149 ssp. charlestonensis. 2: 141 scabrata.2:168
atlantica, 2:141 var. capillacea. 2: 141 schweinitzii, 2:126, 151, 155
ssp. atlantica, 2:125, 141, 146 var. keweenawensis, 2: 141 scoparia, 2:124, 130, 140
ssp. capillacea, 2:125, 141, joorii,2:168 seorsa.2:168
146 josselynii, 2:141 stellata var. conferta. 2: 141
var. incomperta, 2:141 lacustris. 2:126, 141, 150 stellulata. 2: 141
aurea, 2:123, 130 var. laxiflora. 2: 151 var. scirpoides, 2:141
barrattii, 2:127, 151, 161 lanuginosa. 2:126 sterilis. 2:168
bebbii, 2:124, 141, 142 laricina.2:141 stipata. 2:124. 130, 117
brunnescens, 2:125, 141 lasiocarpa. 2:126. 141, 148 var. maxima. 2:130
bullata, 2:127, 151, 158 var. americana, 2: 141 straminea. 2:125. 141, 141
var. greenii, 2: 151 leersii. 2:141 striata. 2:126. 127. 141, 150
buxbaumii, 2:127,151,161 lenticularis, 2:123, 130, 114 var. brevis. 2:141
canescens, 2:125. 141, 144 var. lenticularis, 2:130 stricta. 2:130
var. disjuncta, 2:141 lepidocarpa. 2:168 var. stricta. 2:124. 130, 115
var. subloliacea. 2:141 leptalea, 2:123, 127 var. strictior. 2:124. 130, 115
capitata.2:168 ssp. harperi. 2:127 strictior, 2:130
cephalantha.2:141 limosa. 2:127, 151, 162 suberecta. 2:124. 130, 118
chordorrhiza, 2:124, 130, 116 livida. 2:127, 151, 164 substricta.2:130
comosa, 2:126, 151, 154 var. grayana, 2:151 tenuiflora, 2:125. 141
crawei.2:168 var. radicaulis, 2:151 tetanica. 2:127. 151, 167
crawfordii, 2:124, 140, 141 longii.2:168 torta. 2:123. 127, 128
crinita, 2:130 louisianica, 2: 168 tribuloides, 2:168
var. brevicrinis, 2:123, 130 lupuliformis, 2:127,151,160 trichocarpa. 2:126. 141, 148
var. crinita, 2:123, 130, 111 lupulina. 2:127, 151, 162 trisperma. 2:125. 141
var. gynandra. 2:130 lurida. 2:126,151,156 tuckermanii. 2:126. 151, 157
var. mitchelliana, 2:127 lyngbyei, 2:123. 130, 111 uberior; 2:130
cristatella, 2:124, 141, 142 mackenziei. 2:124, 125. 141, 144 utriculata, 2:126. 151, 157
crus-corvi, 2:124. 130, 117 michauxiana, 2:126, 151, 151 vesicaria. 2:127. 151, 159
cryptolepis, 2:127, 167,168 mitchelliana, 2:123. 127. 129 viridula, 2:151
davisii,2:168 muricata var. cephalantha. 2:141 ssp. brachyrrhyncha, 2:127. 151,
decomposita, 2:124, 130, 117 muskingumensis, 2:168 167
diandra, 2:124,130,118 nigra, 2:123, 130, 114 ssp. viridula. 2:127. 151,165
disperma, 2:124, 130 var. strictiformis. 2:130 vulpinoidea. 2:124. 130, 119
echinata, 2:125, 141, 147 oederi var. viridula, 2: 151 walteriana. 2: 141
var. conferta, 2:141 oligosperma, 2:126. 151, 157 wiegandii. 2:125, 141, 145
emoryi, 2:124, 130, 115 paleacea, 2:123, 127, 129 Carolina Ash. 1:323
exilis, 2:123, 141, 145 pauciflora. 2:123. 127, 128 Carolina Quillwort. 1:25
festucacea, 2:168 paupercula. 2:127, 151, 161 Caryophyllidae. 1:79
flava, 2:127, 166, 168 praegracilis, 2:168 Cashew Family, 1:245
var. fertilis, 2: 168 prairea, 2:124, 130, 118 Cassandra calyculata. 1:161
var. viridula, 2:151 prasina, 2:168 Castalia
folliculata, 2:126,151,151 projecta, 2:168 leibergii. 1:41
378 Index
Honeysuckle Family. 1:372 ellipticum. 1:113. 115, 118 Iris Family. 2: 323
Hooded Ladies-tresses. 2:347 gymnanthum. 1:113. 117, 118 ISOETACEAE. xxvi. xxxviii. xxxix.
Horned Pondweed. 2:58 hypericoides. 1:110. 113, 114 1:25
Horned Pondweed Family. 2:58 var. hypericoides, 1: 113 Isoetes. xxvii. xxxviii. xxxix. xliv. 1:25
Hornpod. 1:267 var. multicaule. 1:113 acadiensis. 1:25
Hornwort. 1:50 kalmianum. 1:113, 114 braunii. 1:25
Hornwort Family. 1:50 majus. 1:113.116, 118 Xbrittonii. 1:27
Horsetail. 1: 17 mutilum. 1:113. 117,118 caroliniana. 1:25
Common. 1:17 var. latisepalum,l: 118 xdodgei. 1:27
Field. 1:17 var. parviflorum. 1: 118 Xeatonii. 1:27
Marsh. 1:17 petiolatum. 1: 11 0 echinospora. 1:25,26,27
Meadow. 1:17 punctatum. 1:113. 116, 118 var. braunii. 1:25
Variegated. 1: 17 pyramidatum. 1: 118 var. muricata. 1:25
Water. xvii. 1:17 stans. 1:113, 114 engelmannii. 1:25,26,27
Wood. 1:17 tubulosum. 1:110 var. canadensis. 1:25
Horsetail Family. 1: 17 var. walteri, 1: 11 0 var. caroliniana. 1:25
Hottonia. xxvi. xxxi. xxxii. 1: 168 virginicum. 1:110 var. valida. 1:25
infiata. 1:168, 170 var. fraseri. 1: 11 0 xfoveolata. 1:27
Huckleberry. 1: 164 Xharveyi. 1:27
Black. 1:168 Ibidium Xheterospora. 1:27
Dwarf. 1:168 cernuum.2:347 Xhickeyi. 1:27
Hybrid Cattail. 2:306 laciniatum. 2:347 hieroglyphica. 1:25
Hydrilla. xvi. xlvii. 2:28. 33 odoratum. 2:347 Xjeffreyi.1:27
verticillata. xvi. 2:11, 33 praecox. 2:347 lacustris. 1:25,26,27
Hydrocharis, xxiii. xxxv. xlvi. 2:11. romanzojJianum. 2:347 macrospora. 1:25
28 Ilex. xxi. 1:237 f. hieroglyphica. 1:25
morsus-ranae. xvi. 2:28, 29 bronxensis. 1:243 melanopoda. 1:25,26
HYDROCHARITACEAE. xxiii. xxvi. fastigiata. 1:243 muricata. 1:25
xxxv. xl. xlvi. 2:11 glabra. 1:237,242,243 prototypus. 1:25
Hydrocotyle. xxxiv, xxxv, 1:245. 250 laevigata. 1:237.242,243 riparia. 1:25,26,27
americana. 1:250,251 verticillata. 1:237.241,243 var. canadensis. 1:25
australis. 1:250 var. fastigiata. 1 :243 var. palmeri. 1:25
canbeyi. 1:250 var. padifolia. 1:243 var. reticulata. 1:25
ranunculoides, xxxiii. 1:250,251 var. tenuifolia, 1:243 saccharata, 1:25
umbellata. 1:250,251 Illinois Pondweed. 2:58 tuckermanii. 1:25,26,27
verticillata. 1:250,251 Impatiens. xlix. 1:245 valida. 1:25
var. featherstoniana. 1:250 biflora. 1 :245 Isolepis. 2:224
var. triradiata. 1:250 capensis. 1:245,246 koilolepis. 2:233
var. verticillata. 1:250 fulva. 1:245 Isotria. 2:328. 330
Hydrolea. I. 1:285 nortonii. 1:245 verticillata. 2:330, 112
ajJinis. 1:285 pallida. 1:245,246 Iva. xx, 1:379.414
ovata. 1:285,287 Indian Lotus, 1:44 frutescens. 1:414,417
quadrivalvis. 1:284, 285 Indian Poke. 2:320 ssp. frutescens. 1:414
uniflora. 1:285,286 Inkberry. 1:243 ssp. oraria. 1:414
HYDROPHYLLACEAE.I.1:285 IRIDACEAE. xxxviii, 2:323 var. oraria, 1:414
Hydrotida caroliniana. 1:345 Iris. xxxviii. 2:76. 323 Ivy. Poison. 1:245
Hypericum. xx. xlviii. 1:110 brevicaulis. 2:323, 127
adpressum. 1:113. 117,118 foliosa. 2:323 Jacob's-ladder. 1:285
ascyron. 1:113. 115, 118 fulva. 2:323, 124 Jewelweed. 1:245
boreale. 1:113, 116, 118 Lamance.2:323 Joe-Pye Weed. 1:418. 426
f. callitrichoides, 1: 118 prismatica, 2:323 Jumpseed. 1:88
canadense. 1:113. 117,118 var. austrina. 2:323 JUNCACEAE. xxv. xxvi. xxxix. xl. 2:97
var. galiforme. 1: 118 pseudacorus. 2:323, 125 JUNCAGINACEAE. xxxv. xxxix. xl.
densiflorum. 1:113, 115 Red. 2:323 2:33
denticulatum. 1:113. 118 shrevei. 2:328 Juncus. xxxix. xl. xlv. 2:97
var. acutifolium. 1: 118 versicolor. 2:323. 126, 127, 328 abortivus. xxv. 2:100.109,117
var. denticulatum. 1:118 virginica. 2:323. 327, 328 acuminatus. 2:99. 109, 118
var. ovalifolium. 1: 118 var. shrevei. 2:328 alpinoarticulatus, 2: 1 09
var. recognitum. 1: 118 var. virginica. 2:328. 126 alpinus. 2:100. 109, 118
dissimulatum. 1:113. 118 Yellow. 2:323 ambiguus. 2:98. 100
Index 385
Puccinellia. 2:254. 267 flammula. 1:56. 57, 62 Reed Canary Grass, 2:283
fernaldii. 2:267,269 var. fili/ormis. 1: 57 Reed Grass. 2:271
langeana.2:267 var. laxicaulis. 1: 5 7 Salt,2:273
ssp. alaskana. 2: 267 var. ovalis. 1: 5 7 Reed Manna Grass, 2:271
maritima. 2:267, 269 gmelinii. 1:56. 57, 61 Reed Meadow Grass. 2:271
pallida. 2:267, 270 var. hookeri. 1:57 Reedgrass.2:283
paupercula var. alaskana. 2:267 hederaceus. 1:56. 57,65 RHAMNACEAE, xxi. 1:243
Pumpkin Ash. 1 :323 hispidus. 1:57 Rhamnus, xxi. 1 :243
Purple Avens, 1:188 var. caricetorum. 1:56. 58, 66 alnifolia, 1:242, 243
Purple Chokeberry. 1: 182 var. nitidus. 1:56. 57, 66 cathartica, 1:243
Purple Gerardia. 1:338 hyperboreus. 1:56. 57, 65 frangula, 1:243
Purple Loosestrife. xvii, 1:203 intermedius. 1:58 Rhexia, xlvii. 1:214
Purple Meadow-rue. 1:52 laxicaulis. 1:56. 57, 63 aristosa, 1:230,231
Purple-fringed Orchid longirostris. xxvii. 1:56. 57, 59 ciliosa, 1: 2 30
Large. 2:334 mississippiensis. 1: 5 7 delicatula, 1:230
Small. 2:334 nitidus. 1: 5 7 interior, 1:230
Purple-honeysuckle. 1:164 obtusiusculus. 1: 5 7 lanceolata, 1:230
Purple-leaved Willow-herb. 1:210 pensylvanicus. 1:57. 58,66 mariana, 1:230
Purple-stemmed Aster. 1:406 purshii. 1:57 var. interior, 1:230, 232
Pycreus albomarginatus. 2:212 pusillus. 1:56. 57, 63 var.leiosperma, 1:230
Pyrus var. angustifolius. 1:57 var. mariana, 1:230, 233
arbutifolia. 1:182 repens. 1:57. 58, 65 var. purpurea, 1:230
var. atropurpurea. 1:182 var. nitidus. 1:57 var. ventricosa. 1:230,234
floribunda. 1:182 reptans. xxxix. 1:57 nashii, 1:230,235
melanocarpa. 1: 18 2 var. ovalis. xliv, xlvi, lv, 1:56. 57, petiolata. 1:214.230,231
64 stricta, 1: 230
Quillwort. 1:25 var. reptans, 1:56. 57, 64 ventricosa, 1:230
Acadian. 1:25 sceleratus. xxxiv. 1:56, 57, 64 virginica. 1:230,232
Big. 1:25 ssp. multifidus, 1:56, 57 var. purshii, 1:230
Black-footed. 1:25 ssp. sceleratus, 1:56, 57 var. septemnervia. 1:230
Carolina. 1:25 septentrionalis, 1: 5 7 Rhododendron, 1:158. 164
Engelmanns. 1:25 var. caricetorum, 1:58 atlanticum, 1:164. 165
Lake. 1:25 sicae/ormis. 1: 5 8 canadense, 1:164, 165
Riverbank, 1:25 subrigidus, 1:56, 57 canescens, 1:164, 166
Spiny-spored. 1:25 tener, 1:57 coryi, 1: 164
Quillwort Family. 1:25 texensis, 1: 5 7 maximum. 1:163, 164
trichophyllus, 1:56, 57,59 nudiflorum, 1: 164
Radicula aquatica. 1:148 ssp. eradicatus, 1: 5 7 oblongifolium, 1: 164
Ragged Fringed Orchid. 2:334 ssp. trichophyllus. 1: 5 7 periclymenoides, 1:163, 164
Ragwort. 1:382. 398 var. calvescens, 1:57 serrulatum, 1:164
Golden. 1:398 var. eradicatus, 1: 5 7 viscosum. 1:164. 165
Ram's-head Ladys-slipper. 2:330 Rattle-box. 1:214 Rhodora, 1:164
RANUNCULACEAE. xxvi. xxxi. xxxiii. Rattlesnake Manna Grass. 2:271 canadense. 1: 164
xxxv. xxxix. xl. xlvi. xlix. I. 1:50 Red Alder, 1: 74 Rhus vernix, 1:245
Ranunculus. xxviii. xxxiii. xxxiv. I. 1:50. Red Ash, 1:323 Rhynchospora, 2:121. 168, 248
52 Red Birch. 1:74 alba, 2:173,174,178
ambigens. 1:56. 57, 62 Red Chokeberry, 1:182 caduca, 2:173,174,178
aquatilis. 1:57 Red Iris. 2:323 capillacea, 2:173, 180, 182
var. calvescens. 1:57 Red Maple. 1:243 capitellata. 2:173, 179,182
var. capillaceus. 1: 5 7 Red-osier Dogwood. 1:237 cephalantha. 2:173.174,178
var. eradicatus. 1: 57 Red-root. 2: 317 var. microcephala, 2:182
var. subrigidus. 1:57 Red-rooted Cyperus. 2:217 var. pleiocephala, 2: 174
caricetorum. 1:58 Red-stemmed Dogwood. 1:237 chalarocephala, 2:173, 180, 182
carolinianus. 1: 5 8 Red-stemmed Gentian. 1:274 colorata, 2:168. 170,174
circinatus var. subrigidus. 1: 57 Red-veined Dock, 1:100 corniculata, 2:174
cymbalaria var. cymbalaria. 1:56. Red-willow. 1:237 var. corniculata. 2:172,173.
57,61 Redtop, 2:285 174
delphini/olius. 1: 5 7 Reed, xvii, 2:261 var. interior. 2:173,174
filiformis. 1: 5 7 Common, 2:261 cymosa, 2:174
flabellaris. 1:56. 57, 60 Reed Bentgrass. 2:283 debilis, 2:174, 181,182
Index 393
fascicularis, 2:181, 182 ssp. hispida, 1:144, 147 maritima, 2:58, 65, 66
var. distans, 2:174,182 var. hispida, 1: 148 var. occidentalis, 2:58
var. fascicularis, 2: 1 74, 182 ssp. palustris, 1:144, 147 occidentalis, 2: 5 8
filifolia, 2:174, 181, 182 var. palustris, 1:144 RUPPIACEAE, xxv, 2:58
fusca, 2:174,181,182 var. hispida, 1:148 Rush,2:97
globularis, 2:173,174,177 var. Xreptabunda, 1:148 Beak-, 2:168
var. recognita, 2: 174 sessiliflora, 1:144, 145 Common Scouring, 1:23
g1omerata, 2:173, 179,182 sinuata, 1:144, 145, 148 Flowering-, xvii, 2:3
gracilenta, 2:174, 181, 182 sylvestris, 1:144, 145, 148 JOinted Spike-, 2:193
harveyi, 2:173,174,175 Rosa, xx, 1:182, 184 Needle-, 2:100, 193
inexpansa, 2:173, 174, 176 dasistema, 1: 184 Nut-, 2:168
inundata, 2:172, 173, 174 floribunda, 1:184 Scouring, 1:17
latifolia, 2:174 lancifolia, 1:184 Slender Spike-, 2:193
macrostachya, 2:172,174 nitida, 1:184, 186 Smooth Scouring, 1:23
var. colpophila, 2:173, 174 palustris, 1:184, 187 Soft, 2:97, 100
var. macrostachya, 2:172,173,174 var. dasistema, 1: 184 Spike-, 1:191
microcephala, 2:173, 178, 182 virginiana, 1:184, 186 Square-stem Spike-, 2:193
miliacea, 2:173,174,179 ROSACEAE, xx, xxi, xxxi, 1: 179 Sweet-, 2:202
nitens, 2:168, 171,174 Rose, 1:184 Triangle Spike-
oligantha, 2:173,174,176 Northeastern, 1:184 Twig-, 2:182
pallida, 2:173, 180,182 Swamp, 1:184 Rush Aster, 1:406
perplexa, 2:173,174,177 Virginia, 1:184 Rush Family, 2: 9 7
var. virginiana, 2:174 Rose Family, 1: 179 Rush Family, Flowering 2:3
rariflora, 2:173,174,175 Rose Pogonia, 2:334 Rye
scirpoides, 2:168, 171,174 Rosebay, 1:164 Virginia Wild, 2:261
torreyana, 2:173,174,176 Rose-gentian, 1:270 Wild,2:261
Ribwort, 1:323 Rose-mallow, 1: 118,
Riccia, xxiii, xxiv Swamp, 1:120 Sabatia, 1:267,270
RICCIACEAE, xxiii Wooly, 1:120 calycina, 1:270
Ricciocarpus, xxiii, xxiv Rosidae, 1: 1 77 campanulata, 1:270,271
Rice Rotala, xlviii, 1:203,206 var. gracilis, 1:270
False Wild-, 2:261 ramosior, 1:206,208 difIormis, 1:270
Jungle, 2:292 Rough-leaved Goldenrod, 1:400 dodecandra, 1:270,272
Northern Wild-, 2:255 Round-leaved Sundew, 1:125 var. kennedyana, 1:270
Southern Wild-, 2:255, 261 RUBIACEAE, xx, xlviii, 1:370 gracilis, 1:270
Wild,2:255 Rubus, 1:182, 188 harperi, 1:270
Rice Cutgrass, 2:281 acaulis, 1:188 kennedyana, 1:270,271
Richweed, 1:68 chamaemorus, 1:188, 190 lanceolata, 1:270
River Birch, 1: 74 pubescens, 1:188 Large, 1:270
River Bulrush, 2:233 setosus, 1:188 stellaris, 1:270, 271
River-beauty, 1:210 vigil, 1:188 Sacciolepis, 2:254, 288
Riverweed, 1: 191 Rumex, xlix, Ii, liv, 1:85, 100 striata, 2:288, 291
Riverweed Family, 1:191 a1tissimus, 1:100, 101, 106 Sacred Bean, 1:44
Robbins' Pondweed, 2:44 britannica, 1:101 Sacred Lotus, 1:44
Rorippa, 1: 144 crispus, 1:100, 103 Sagittaria, xxvi, xxix, xxxv, xxxix, xlii,
amphibia, 1:144, 145, 148 maritimus, 1: 1 00 2:3,5,78
aquatica, 1:148 var. fueginus, 1:100, 102 ambigua, 2:10, 11,22
hispida, 1:148 var. persicarioides, 1:100, arifolia, 2:11
islandica, 1: 144 102 australis, 2:10, 11,22
var. fernaldiana, 1: 144 mexicanus, 1:101 brevirostra, 2:10, 11,26
var. glabra, 1:144 obtusifolius, 1:100, 102 calycina, 2: 1 0
var. glabrata, 1: 144 orbiculatus, 1:100, 101, 104 var. spongiosa, 2:11
microphyIIa, 1:153 pallidus, 1:100, 101, 105 cristata, 2:10, 11
nasturtium-aquaticum, 1: 15 3 persicarioides, 1: 100 cuneata, 2:10, 11,25
var. longisiliqua, 1: 15 3 salicifolius ssp. triangulivalvis, eatonii, 2:11
palustris, xxxii, xxxiv, xlix, 1: 144, 1:101 engelmanniana, 2:10, 11,27
146 triangulivalvis, 1:100, 101, 105 ssp. brevirostra, 2: 11
ssp. glabra, 1:144, 147 verticillatus, 1:100. 101, 104 ssp. longirostra, 2: 11
var. fernaldiana, 1:144, 148 Ruppia, xxv, xxviii, 2:58 faIcata, 2: 11
var. glabrata, liv, 1: 144 cirrhosa, 2:58 filiformis, 2:10, 11, 15
394 Index
Sagittaria (continued) candida, 1:131. 132, 138, 139, 141, Salsola salsa var. americana, 1:82
graminea, 2:11,19 142, 143 Salt Grass, 2:255
ssp. graminea, 2:10,11,19 var. denudata, 1:143 Salt Marsh Aster
ssp. weatherbiana, 2:10,11,20 caroliniana, 1:131. 132, 133, 134, Annual, 1:406
var. cristata, 2, 11,21 142 Perennial, 1 :406
var. platyphylla, 2:11 coactilis, 1:131, 132, 133, 140, 143 Salt Marsh Bulrush, 2:233
var. weatherbiana, 2:11,20 cordata, 1:131, 132, 133, 140, 141 Salt Reed Grass, 2:273
heterophylla, 2:11 var. abrasa, 1:133 Saltmeadow Cord Grass, 2:273
var. rigida, 2: 11 var. missouriensis, 1: 13 3 Saltmeadow Grass, 2:273
lancifolia, 2:10, 11, 18 cryptodonta, 1:131. 132, 140, 143 Saltwater Cord Grass, 2:273
ssp. media, 2:11 discolor, 1:131, 132, 133, 136, 141, Saltwort, 1: 79
latifolia, xxxvii, 2:5, 10, 11,23,24 142 Dwarf,1:79
var. obtusa, 2:11 var. eriocephala, 1:133, 137 Perennial, 1:79
var. pubescens, 2: 11 var. latifolia, 1: 13 3 Salvinia, xxiii, 1: 14
longirostra var. australis, 2: 11 var. overi, 1:133 minima, 1:14, 16
mohrii, 2: 11 eriocephala, 1:131. 132, 133, 142 SALVINIACEAE, xxiii, 1:14
montevidensis, 2:10, 12 exigua, 1:129, 132, 133, 134, 141, Sambucus, xx, 1:372
ssp. calycina, 2:5, 10 142 canadensis, 1:372,373
ssp. spongiosa, 2: 1 0, 13 ssp. interior; 1: 13 3 var. submollis, 1:372
platyphylla, 2:10, 11, 14 j1uviatilis, 1: 13 3 Samolus, I, 1:168, 171
rigida, 2:10, 11,17 glaucophylloides, 1: 143 j1oribundus, 1: 171
spatulata, 2: 11 var. albovestita, 1:143 parviflorus, 1:170, 171
stagnorum, 2:11 var. glaucophylla, 1:143 valerandi ssp. parvijlorus, 1: 171
subulata, 2:10, 11, 15 gracilis, 1: 14 3 Samphire, 1:79
var. gracillima, 2: 11 var. angustifolia, 1:143 Sampson's Snakeroot, 1:274
var. natans, 2:11 var. rosmarinoides, 1:143 Sanguisorba, xxxi, 1:182, 188
teres, 2:10, 11, 16 interior; 1:133 canadensis, 1:188, 190
weatherbiana, 2:11 longifolia, 1: 13 3 Sarcocornia perennis, 1: 79
Sago Pondweed, 2:39 longipes, 1:133 Sarracenia, xxxix, xlvi, 1:120
st. Andrew's Cross, 1:113 lucida, 1. 129, 132, 133, 135, 141 flava, 1:120, 124
st. John's-wort, 1:110 ssp. lucida, 1:133 purpurea, 1:120, 124
Creeping, 1:118 var. angustifolia, 1:133 SARRACENIACEAE, xxxix, xlvi,
Great, 1:118 var. intonsa, 1:133 1:120
Kalm's, 1:113 var. serrisima, 1: 13 3 Satureja
Marsh, 1: 11 0 myricoides, 1:131, 132, 133,137, arkansana, 1:306
Narrow-leaved, 1:118 141, 143 glabella, 1:306
st. John's-Wort Family, 1:110 var. albovestita, 1: 143 glabra, 1:306
St. Peter's-wort, 1: 113 myrtillifolia, 1:131. 132, 133, 139 SAURURACEAE, xxxvi, 1:35
SALICACEAE, xxi, 1:129 nigra, 1:131, 132, 133, 135, 142 Saururus, xxxvi, 1:35
Salicornia, xxxviii, xlvi, 1:79 var. falcata, 1: 13 3 cernuus, xxxvii, 1:35,37
bigelovii, 1:79,80 pedicellaris, 1:131, 132, 133, 138, Saw-grass, 2:182
europaea, 1:79,81 141, 142, 143 Saxifraga, xlvi, 1: 17 7, 179
var. pachystachya, 1:79 var. hypoglauca, 1:143 pensylvanica, 1:178,179
var. prostrata, 1:79 var. tenuescens 1:143 SAXIFRAGACEAE, xlvi, xlvii, xlix,
var. simplex, 1:79 pellita, 1:131, 132, 133, 140, 141, 1:177
perennis, 1:79 143 Saxifrage, xlvi, 1: 179
rubra, 1:79, 80 petiolaris, 1:131. 132, 133, 137, Golden, 1: 177
virginica, 1:79,81 141,142,143 Swamp, 1:179
Salix, xxi, 1:129 var. gracilis, 1:143 Saxifrage Family, 1: 177
adenophylla, 1: 13 3 var. rosmarinoides, 1: 143 Scarlet Maple, 1:243
amygdaloides, 1:131. 132, 133, 135, rigida, 1:133 Scheuchzeria, xli, 2:33
141,142 var. angustata, 1:133 palustris, 2:33,35
angustata, 1:133 sericea, 1:131, 132, 133, 136, 138, ssp. americana, 2:33
bebbiana, 1:131. 132, 138, 141, 141,142,143 var. americana, 2:33
142, 143 serrisima, 1:129, 132, 133, 134, Scheuchzeria Family, 2: 3 3
var. capreifolia, 1: 143 141, 142 SCHEUCHZERIACEAE, xli, 2:33
var. luxurians, 1:143 synticola, 1: 13 3 Schizaea, xl, 1:3
var. perrostrata, 1:143 wardii, 1:133 pusilla, 1:3, 6
var. projecta, 1:143 wheeleri, 1: 13 3 SCHIZAEACEAE, xl, 1:3
Index 395