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CLASSIFICATION OF WETLANDS
.AND DEEP-WATER HABITATS
OF THE UNITED STATES
(An Operational Draft)
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87.3
.C533
1977 October 1977
FISH AND WILDLIFE SERVICE
U.S.DEPARTMENT OF THE INTERIOR
.'
CLASslFICATIONOFWETLANDS AND DEEP-WATER
HABITATS OF THE UNITED STATES"-'-.........'-.-.....
,(An Operati.onaIDraft)
By Lewis M.Cowardin,U.S.Fish and Wildlife Service,Northern
Prairie Wildlife Research Center
Virginia Carter,U.S.'Geological Survey,Water Resources
Division
Francis C.Golet,Department of Forest and Wildlife
Management,University of Rhode Island
Edward T.LaRoe,U.S.National Oceanic and Atmospheric
Administration,Office of Coastal Zone Management ,
Edited by J.Henry Sather
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.C533
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October 1977
Fi~h and Wildlife Service
U.S.Department of the'lnterior ARLIS
Alaska Resources
Library &Information Servtces
Anchorage,Alaska
FOREWORD
•.~j
The NationalWetlimdInventoryProjectbf197S-'-/'9 has sought
(as did its predecessor of 1954,th,e first N<:lt'ional Wetland Inventory
,Proj ect),a wetland classification system that could be consistently
and equally applied to the varying'aspects of the wetland resources
of the United States.
The result of that search,"The Classification of Wetlands and
Deep-Water Habitats of the United States,"has been under development
for over two years,and has undergone peer review to an exteritwhich
is perhaps unprecedented in the history of documents published by the
Fish and Wildlif~Ser:vice.We want to acknowledg.e and thank those
individuals and organizations who gave us assistance and guidance during
the development of this system.'
We have termed this document "An Operational Draft,"and the
selection of that phrase was made for two reasons.'First,this classi-
fication system needs to be used in the field for a substantial period
of time so that inconsistencies can be isolated and viable alternatives
be examined.Secondly,similar to the Martin.,.et al<1953)system,this
system will most assuredly need revising in the~oriths and years ahead.
-1._.
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,-
TABLE OF ~ONTENTS
Page
I.INTRODUCT ION
I I.WETLANDS AND DEEP-WATER HABITATS
CONCEPTS .
DEFINITIONS
Wetland .,.
....
3
3
4
4
Systems and Subsystems
HI ERARCHI CAL STRUCTURE .
II I.THE CLASSIFICATION SYSTEM
Deep-water Habitats 5
5
7
7
7
.,
~.
.,
,. .
,.
.~
13
13
13
13
14
14
14
14
14
14
15
15
16
16
16
16
16
16.ARtis·16
Alaska Resources
Library &Information Semces
Anchorage,Alaska
",..
.,
..
,.
,.,
. ......
i i
.,
.,
..'
Definition
Limits ..•
Definition
Limits ..
Deseri pt ion
Subsystems •
Subtidal.
Intertidal
Classes
Definition
Limits ..
Des c rip t i on
Subsystems .
Subtidal.
Intertidal
Classes
RIVERINE.
MARl NE . . . .
ESTUARINE
LIMITS ...
...
..
TABLE OF CONTENTS (c,ontinued)
Classes,Subclasse~and Do~inahce Types
Defi nit i on
Oeser i pt ion
Subclasses and Dominance Types
Bedrock
Boulder
UNCONsoLIDATED BOTTOM
ROCK BOTTOM
If
,-
Page
17
17
18
18
18
19
19
19
19
20
20
20
20
20..21
21
21
21
21
22
22
23
26
26
26
27
27
27
27
27
27
29
.29
•29
30
30
Types
.'.
.'
Definition
Limi ts ..
Descri pt ion
Subsystems
Li'mnetic
Li ttoral
Classes
Defi nit i on
Limi ts ..
Description
Subsystems
Classes
Descri pti on
Subsystems .
Ti dal
Lower Perennial
Upper Perennial
Intermi ttent
Classes
Definition.
Description
Subclasses and Dominance
Cobble/Gravel
Sand . . .
Mud
Organic
LACUSTRINE
PALUSTRINE
i i j
TABLE OF CONTENTS (cot'!t inued)
Page
AQUATIC BED
ROCKY SHORE
Definition
Description
Oefi ni t ion
Descri pt ion
Subclasses and Dominance TYPeS
Cobble/Gravel
Sand ..
Mud
Organ ic
Defirrition .
Description .
Subc lasses and Domi nance Types
Submergent Algal ..
Submergent Vascular
Submergent Moss
Floating-leaved
Floating .
.30
30
30
30
31
32
32
33
33
33
33
34
34
34
34
35
35
35
36
36
36
36
37
37
37
37
37
38
38
38
39
39
39
39
39
,,
.,
..
,..
..
.,
"
..
".
".
Types "
Dominance Types •
Definition
Descri pti on ".
Subclasses and
Coral
Mollusc
Worm
Definition
- - -- -
Descri pt i on
Subclasses and Dominance
"Cobble/Gravel .•
Sand .•
Mud ."....
Organic
Vegetated Flats
REEF ....
FLATS
STREAMBED
..
•
·iv
TABLE OF CONTENTS (cont i nued)
FORESTED WETLAND
SCRUB/SHRUB WETLAND
EMERGENT WETLAND
Subclasses and Dominance Typei
Bedrock
Boulder
...
0,
.-
Page
39
40
40
..41
41
41
42
42
42
43
43
43
43
43
43
44
.44
44
44
44
45
46
46
46
46
46
46
47
..47
47
47
47
47
48
48
.'
Types
.'
Defi nit i on
Description
Subcl asses and Domi nance
Cobble/Gravel
Sand .
De fin I t i on . .
Description
Subclasses and Dominance Types
Moss ..'
Li chen .
De r i nit i on . .
Description
Subclasses and Dominance Types
Broad-leaved Deciduous
Needle-leaved Deciduous
Broad-leaved Evergreen.
Needle-leaved Everg~een
bead ....
De fin it i on . .
Description
Subclass~s and Dominance Types
Persistent.
Nonpersistent.
Def i nit ion . .
.Des cr i pt ion
Subclasses and Dominance Types
Broad~leaved Deciduous'
MOSS/LICHEN WETLAND
BEACH/BAR
v
TABLE OF CONTENTS (continued)
EXCAVATED
ART I FI CIAL
NONTIDAL .
!MPOUNDED
Page
48
48
49
49
49
49
50
50
50
51
51
51
52
52
.52
52
52
52
53
53
53
54
57
58
59
60
60
60
60
61
61
..
Need Ie-I eavedDec i duous
Broad-leaved Evergreen .
Needle-leaved Evergreen
Dead .
TIDAL
Subti da I
Irregularly Exposed
Regularly Flooded ~
Irregularly Flooded
Permanently Flooded.
Intermittently Exposed
Semi permanently Flooded
Seasonally Flooded.
Saturated.... . . . .
Temporarily Flooded ..
Intermittently Floo.ded
Artificially Flooded.
pH MO DI FIE RS
SALINITY MODIFIERS
DIKED
FARMED .•
PARTIAL~Y DRAINED
Water Regime Modifiers
Water Chemistry Modifiers
So i I Mod if ie rs
Special Modifiers
.MODIFIERS
..
•
vi
IV.REGIONALIZATION
TABLE OF CONTENTS (cant inued)
Page
62
V.USE OF THE CLASSIFICATION SYSTEM
HIERARCHICAL LEVELS AND MODIFIERS
RELATIONSHIP TO OTHER WETLAND CLASSIFICATIONS
LITERATURE CITED ~..
APPENDIX A -SCIENTIFIC AND COMMON NAMES OF PLANTS
APPENDIX B -SCIENTIFIC AND COMMON NAMES OF ANIMALS
APPENDIX C -CRITERIA FOR OIST1~GUISHING ORGANIC
SOILS FROM MINERAL SOILS .
vi i
67
69
71
79
90
94
97
Table
LI ST OF TABLES
Pilqe
pH Modifiers Used in This Classification System2
Salinity Modifiers Used in This
Classification System..",.......,56
58
3
4
5
Comparison of Wetland Types Described in
Circular 39 with Some of the MC;ljor Components
Of this Classification System .
Comparison of the Zones of Stewart and Kantrud's
(1971)Classification with the Water Regime
Modifiers used in this Classification System
Comparison of the Water Chemistry Subclasses of
Stewart and Kantrud (1972)with the Water
Chemistry Modifiers used in this Classification
System-. . . . . . • . . . . . . . . . . • .r
vi i i
72
77·
78
"L I ST OF FIGURES
.Figure Page
This figure is not included in this draft.
2 This fi gure is not included in th i.s draft.
3 DIAGRAM OF THE CLASSIFICATION HIERARCHY FOR
THE MARINE SYSTEM ····.~..· ······8
4 DIAGRAM OF THE CLASS IFICAT ION HIERARCHY FOR
THE ESTUARINE SYSTEM ·...·······.9
5 DIAGRAM OF THE CLASSIFiCATION HIERARCHY FOR
THE RIVERINE SYSTEM ·.· ·
.•.··· ··10
6 DIAGRAM OF THE CLASSIFICATiON HIERARCHY OF
THE LACUSTRINE SYSlEM ·.· ·
·....· · ·····11
7 DIAGRAM OF THE CLASSIFICATION HIERARCHY OF·
THE PALUSTRINE SYSTEM .
ix
e·• • • • •-.• •
12
".
I. I NTRODUCT ION
In 1954,the U.S.Fish andWildl ife Service conducted an
inventory of the wetlands of the United States (Shaw and Fredine 1956).
Since then,wetlands in this country have undergone considerable change,
both natural and man-related,and their characteristics and natural
values have become better defined and more widely known.During this
interval,state and federal legislation has been passed to protect wet-
lands,and some statewide wetland surveys have been conducted.
In 1974,the U.S.Fish and Wildlif~Service directed its Office
of Biological Services to design and conduct a new national inventory of
wetlands.Whereas the single purpose of the 1954 inventory was to assess
the amount and types of valuable waterfowl habitat,the scope of the new
project is considerably broader (Montanari and Townsend,..!..!!.press).It
will provide basrc data on the characteristics and extent of the nation's
wetlands and deep-water habitats and facilitate the management of these
areas on a sound,multiple-use basis.
Before the 1954 inventory was begun,Martin et al.(1953)had
devised a wetland classification system to serve as a framework for the
national inventory.The results of the inventory and an illustrated
description of the 20 wetland types were published as Circular 39
(Shaw and Fredine 1956).This document has been one of the most common
and most influential tools used in the continuous battle to preserve a
ra~idly vanishing and critically valuable national resource (Stegman
1976).However,the shortcomings of this work are well-known and have
been documented (e.g.,Leitch 1966,Stewart and Kantrud 1971).
2
In their attempt at simpl ification,Martin et ale ignored eco-
logically critical di~ferences,such as the distinction between fresh and
subsaline in~and wetlands;also,dissimilar habitats,such as boreal
black spruce forests and southern cypress-gum forests were often placed
in the same category,with no provision in the system for distinguishing
between them.Because of the central emphas is on waterfowlhab i tat,far
greater attention was paid to vegetated areas than tononvegetated areas.
Probably the greatest single disadvantage of the Martin et al.system was
the inadequate definition of types,which led to inconsistencies in
application.
Numerous other classifications of wetlands.and deep-water habitats
have been developed,but most of these are regional systems and none
would fully satisfy natibnal need~(Stewart and Kantrud 1971,Golet and
Larson 1974;Jeglum et a1.1974,Odum et aL 1974,Zoltai et a1.1975,
Millar 1976).Because of the weaknesses inherent in Circular 39,and
because our understa·nding of wetland ecology has grown significantly since
1954,the U.S.Fish and Wildl ife Service elected to construct a new
national classification system as the first step toward a new national
inventory.The new classification has been designed to meet three long-
range objectives:J)to group ecologically similar habitats,so that
value judgments can be made;2)to furnish units·for inventory and
mapping; and 3)to provide uniformity in concepts and terminology through-
out the United States ..
"
....
3
I I.WETLANDS AND DEEP-WATER HABITATS
CONCEPTS
For centuries we have spoken of marshes,swamps and bogs,but
only relatively recently have we attempted to group these landscape units
under the single term,"we tland.11 The need to do this has grown .out of
our desire to understand and describe the characteristics and values of
all types of land,and to wisely and effectively manage wetland eco-
systems.There is no single,correct,indisputable,ecologically sound
definition for wetland,primarily because of the diversity of wetlands
and~because the gradation between dry and wet environments is continuous.
The reasons or needs for defining wetland vary;as a result,a great
prol iferation of definitions has arisen.Our primary task here is to
impose boundaries on natural ecosystems for the purposes of inventory,
evaluation and management.
In general terms,wetland is land where water is the dominant
factor determining the nature of soil development and the types of plant
and animal communities living in the soil and on its surface.It spans
a continuum of environments where terrestrial and aquatic systems
intergrade.The concept of wetland embraces a number of characteristics
including:1)the elevation of the water table with respect to the
ground surface;2)the duration of surface water;3)the soil types that
form under permanently or temporarily saturated conditions;and 4)the
various kinds of plants and animals that have become adapted to I ife in
a "
we t"env.ironment.The single feature that most wetlands share is soil
4
that,at least periodically,is saturated with water.This creates
severe physiological problems for all plants except hydrophytes,which
are adapted for life in water or in soil that is at least periodically
saturated.
Deep-water habitats include environments where surface water is
permanent and often quite deep so that water,rather than air,is the
principal medium within which the dominant organisms live,whether they
are attached to the substrate or not.
We define five major ecological systems:Marine,Estuarine,
Riverine,Lacustrine ahd Palustrine.The first four of these include
both wetland and deep-water habitats while the Palustrine includes only
wet landhab itat.
DEFINITIONS
Wet land
Instead of placing arbitrary limits on the position and
fluctuation of the water table for the purpose of defining wetland,we
have attempted to define wetland broadly and simply,and then to place
1 imits on the concept.For thepurpose of this classification system,
WETLAND is defined as land where the water table is at,near or above
the land surface long enough to promote the formation of hydric soils l
or to support the growth of hydrophytes.2 In certain types of wetlands,
IThe U~S.Soil Conservation Service is currently preparing a
preliminary list of hydric soils for use in this classification system.
2 .
The U.S.Fish and Wildife Service is currently preparing a
1 ist of the hydrophytes of the nited States.
5
vegetation is lacking and soils are poorly developed or ~bsent as a
result of freguent and drastic fluctuations of surface-water levels t
wave action t water flow t turbidity or high concentrations of salts or
other substances in the water or substrate.Such wetlands can be
recognized by the presence of surface water or saturated substrate at
some time during each year and their location within,or adjacent to,
vegetated wetlands or deep-water habitats.
Wetland as defined here includes land that is identified under
other categories in some land-use classifications.For example t wetland
and farmland are not necessarily exclusive.Many areas that we define
as wetland are farmed during dry periods,but if they are not tilled or
planted to crops,they will supporthydrophytes.
DeeE-water Habitats
Permanently flooded lands lying below the deep-water boundary of
wetland are defined as DEEP-WATER HABITATS in this classification.As
in wetlands,the dominant plants arehydrophytes;however t the sub-
strates are considered "not-soil 'l because the water is too deep to
support emergent vegetation (U.S.Soil Conservation Service 1975).
LIMITS
The upland 1 imit of wetland is designated as:1)the boundary
between land with predominantly hydrophytic cover and land with pre-
dominantly mesophytic or xerophytic cover;2)the boundary between soil
that is predominantly hydric and soil that is predominantly nonhydric;
or,in the case of wetlands without vegetation or soils;3)the boundary
6
between land that is flooded or saturated at some time during years of
normal precipitation and land that is not.Areas with drained hydric
soils that are no longer capable of supporting hydrophytes are not
considered wetlands.
The boundary between wetland and deep-water habitat in the Marine
and Estuarine Systems coincides with the elevation of the extreme low
water of spring tide (ELWS);permanently flooded areas are considered
deep-water habitats in these systems.The boundary between wetland and
deep-water habitat 1n the Riverine,Lacustrine and Palustrine Systems
I ies at a depth of 2 m (6.7 ft)below low water;however,if emergents,
shrubs or trees grow beyond this depth at any time,their deep-water edge
is the boundary.Figures I and 2 illustrate the identifying features
and limits of wetlands and deep-water habitats where tidal and nontidal
forces predominate.
The 2 m lower limit for inland wetlands was selected because it
represents the maximum depth to which emergent plants normally grow
(Welch 1952,Zhadin and Gerd 1963,Sculthorpe 1967).As Daubenmire
(1968:138)stated,emergents are not true aquatic plants,but are
"amphibious,"growing in both permanently flooded and wet,nonflooded
soils.In their wetland classification for Canada,Zoltai et al.(1975)
include only areas with water less than 2 m deep.
7
II I~THE CLASSIFICATiON SYSTEM
The structure of this classification is hierarchical,progress-
ing from systems and subsystems,at the most general levels,to classes,
subclasses and dominance types.Modifiers for water regime,water
chemistry and soi Is are applied to classes,subclasses and dominance
types.Figures 3-7 illustrate the classification structure within each
of the five ecological systems.Special modifiers are also included to
describe wetlands and deep-water habitats either created or highly modi-
fied by man or beavers.
HIERARCHICAL STRUCTURE
Systems and Subsystems
The term SYSTEM refers here to a complex of wetland and deep-
water habitats that share the influence of one or more dominant hydro-
logic,geomorphologic,chemical,or biological factors.We have chosen
to subdivide systems into more specific cate~ories called SUBSYSTEMS.
The characteristics of the five major systems have been discussed
at length in the scientific I iterature and the concepts are well-
recognized,but there is frequent disagreement as to which attributes
should be used to bound the systems in space.For example,both the
limit of tidal influence and the limit of ocean-derived salinity have
been proposed for bounding the upstream end of the Estuarine System
(Caspers 1967).As Bormann and Likens (1969)pointed out,boundaries of
ecosystems are defi ned to meet pragmat ic needs.
8
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Submergent Algal ---------------~
--1 Floating -----------------------Spirodela polyrhiza
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.
Submergent Moss ~---------------Orepanocladusfluitans
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.•~~·b~l:/;;~~~~l--------------
Pice a ma ria na
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Taxod iuni d ist i ch i um
Acer rubrum
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13
1.MARl NE
Definition.--The Marine System (Figure 3)consists of the open
ocean overlying the continental shelf and its associated high-energy
coastline.Marine habitats are exposed to the waves and currents of the
open ocean and the water regimes are determined primarily by the ebb and
flow of oceanic tides.SaHnites exceed 30%0 (parts per thousand),
with little or no dilution except opposite mouths of estuaries.Shallow
coastal indentations or bays without appreciable fresh-water inflow,and
coasts with exposed rocky islands that provide the mainland with 1 ittle
or no shelter from wind and waves,are also considered part of the
Marine System because they generally support typical marine biota.
Limits.--The Marine System extends from the outer edge of the
continental shelf to:1)the landward limit of tidal inundation (extreme
high water of spring"tides:EHWS)including the splash zone from break-
ing waves;2)the seaward limit of wetland emergents,trees or shrubs
where they extend into open ocean waters;or 3)the seaward limit of the
Estuarine System where this limit is determined by factors other than
vegetation.Deep-water habitats lying beyond the seaward limit of the
Marine System are outside of the scope of this classification system.
14
Subsystems
(1)Subtidal.This includes that part of the Marine System
in which the substrate is continuously submerged.
(2)Intertidal.This includes that part of the Marine
System in which the substrate is exposed and flooded by tides.It also
includes the associated splash zone.
Classes.--Rock Bottom,Unconsolidated Bottom,Aquatic Bed,Reef,
Flat,Rocky Shore and Beach/Bar.
2.ESTUARINE
Definition.--The Estuarine System (Figure 4)consists of deep-
water tidal habitats and adjacent tidal wetlands which are usually semi-
enclosed by land,but have open,partially obstructed,or sporadic access
to the open ocean and in which ocean water is at least occasionally
diluted by fresh water runoff from the land.The salinity may be
periodically increased above that of the open ocean by evaporation.
Along some low energy coastlines there is appreciable dilution of sea
water.Those offshore areas with typical estuarine plants and animals,
such as mangroves (Rhizophora mangle)and oysters (Crassostrea virginica),
are also included in the Estuarine System even though they are not semi-
enclosed by land.1
IThe Coastal Zone Management Act of 1972 defines an estuary as,
Iithat part of a river or stream or other body of water having unimpaired
connection with the open sea,where the sea water is measurably diluted
with fresh water derived from land drainage.11 The Act further states
that,lithe term includes estuary-type areas of the Great Lakes.11 However,
in this system we will not classify areas of the Great Lakes as estuarine.
15
Limits.--Estuaries extend upstream and landward to the place
where ocean-derived salts measure less than 0.5%0 during the period of
average annual low flow.The seaward limit of the Estuarine System is:
1)aline clos ing the mouth of a river,bay or sound;2)aline enclos-
in~an offshore area of diluted sea-water with typical estuarine flora
and fauna;or 3)the seaward 1 imit of wetland emergents,shrubs or trees
where these plants grow seaward of the line closing the mouth of a
river,bay,or sound.
Description.--The Estuarine System includes both estuaries and
lagoons.It is more strongly influenced by its association with land
than the Marine System.In terms of wave action,estuaries are generally
considered to be low energy systems.
Estuarine water regimes and water chemistry are affected by one
or more of the following forces:oceanic tides,precipitation,fresh-
water runoff from land areas,evaporation and wind.Estuarine salinities
range from hyperhaline to oligohaline (Table 1).The salinity may be
variable (poikilohaline),as in the case of hyperhaline lagoons (e.g.,
Laguna Madre,Texas)and most brackish estuaries (e.g.,Chesapeake Bay,
Virginia-Maryland);or it may be relatively stable (homoiohaline),as in
the case of sheltered euhaline embayments (e.g.,Chincoteague Bay,
Maryland)or brackish embayments with partially obstructed access or
small tidal range (e.g.,Pamlico Sound,North Carolina).(For an
extended discussion of estuaries and l~goons ~ee Lauff [1967J).
16
Subsystems
(1)Subtidal.This includes that part of the Estuarine
System in which the substrate is cOhtinuously submerged.
(2)Intertidal.This includes that part of the Estuarine
System in which the substrate is exposed and flooded by tides.It also
includes the assoii~t~d spla~h zone.
Classes.--Rock Bottom,Unconsolidated Bottom,Aquatic Bed,Reef,
Flat,Streambed,Rocky Shore,Beach/Bar,Emergent Wetland,Scrub/Shrub
Wetland and Forested Wetland.
3.RIVERINE
Definition.--The Riverine System (Figure 5)includes all wetlands
and deep-water habitats contained within a channel,except:1)wetlands
dominated by trees,shrubs,persistent ernergents,nonaquatic mosses or
lichens,aild 2)habitats with waters containing ocean-derived salts in
excess of 0.5 %°'A channel is,"an open conduit either naturally or
artificially created which periodically or continuously contains moving
water~or which forms a connecting 1 ink between two bodies of standing
water"(Langbein and Ised '1960:5).
Limits.--The Riverine System is bounded on the landward side by
upland~by the channel bank (including natural o~man-made levees),or by
wetland dominated by trees,shrubs,persistent emergents,honaquatic
mosses or lichens.In braided streams,the system is bounded by the
banks forming the outer limits of the depression within which the braid-
ing occurs.
17
The Riverine System terminates at the downstream end where the
concentration of ocean-derived salts in the water exceeds 0.5%0 during
the period of annual average low flow,or where the channel enters a
lake.It terminates at the upstream end where tributary streams originate,
whether their flow is perennial or intermittent,or where the channel
leaves a lake.Springs discharging into a channel are considered part
of the Riverine System.
Description.--Water is usually,but not always,flowing (lotic)
in the Riverine System.Upland islands or Palustrine wetlands may occur
ln the channel but they are not included in the Riveri~e System.
Palustrine Forested Wetlands,Emergent Wetlands,Scrub/Shrub Wetlands,
and Moss/Lichen Wetlands may occur adjacent to the Riverine System,often
on a floodplain.Many biologists have suggested that all the wetlands
occ~rring on the river flood plain should be a part of the Riverine
System because they consider their presence to be the result of river
flooding.However,we concur with Reid and Wood (1976:72,84)who state,
liThe floodplain is a flat expanse of land bordering an old river.
Often the floodplain takes the form of a very level plain occupied by
the present stream channel,and it may never,or only occasionally,be
flooded.It is this subsurface water [the ground water]that
controls to a great extent the level of lake surfaces,the flow of
streams,and the extent of swamps and marshes.11
Subsystems.--The Riverine System is divided into four subsystems:
the Tidal,the Lower Perennial,the Upper Perennial,and the
18
Intermittent.Each is defined in terms of water permanence,gradient,
water velocity,streambed compositlon and the extend of flood plain
development.The subsystems have characteristic water temperatures,
flora,and fauna (see Reid 1961,lIlies and Botosaneau 1963,Hynes 1970).
All four subsystems are not necessarily present in all rivers,and the
order of occurrence may be other than that given below.
(1)Tidal.In this subsystem,the gradient is low and
water velocity fluctuates under tidal influence.The streambed is
mainly mud with occasional patches of sand.Oxygen deficits may occur
at times and the fauna is similar to that in the Lower Perennial Sub-
system.The flood plain is typically well-developed and water
temperatures approximate those of the Lower Perennial Subsystem.
(2)Lower Perennial.This includes those channels that
contain nontidal flowing water throughout the year.The flow is slow
and the substrate consists mainly of sand and mud.Oxygen deficits may
occur at times,the fauna is composed mostly of species that reach their
maximum abundance in still water,and true planktonic organisms are
common.The gradient is low compared to that of the Upper Perennial
Subsystem and the flood plain is well-developed.Generally,the average
of mean monthly water temperatures is more than 20°C,and in tropical
latitudes-,the average of the monthly means during the summerlTlaV reach
2SoC (Illies and Botosaneau 1963).
(3)Upper Perennial.This includes channels that contain
flowing water throughout the year.The flow is fast and the substrate
consists of rock,cobbles,or gravel with occasional patches of sand.
19
The natural dissolved oxygen concentration is normally near saturation;
the fauna is characteristic of running water,and there are few or
no planktonic forms.The gradient is high compared to the Lower
Perr~nial Subsystem,and there is very little flood plain development.
Generally,the average of mean monthly water temperatures is about 20 C
(Illes and Botosaneau 1963).
(4)Intermittent.This includes those channels that
contain flowing water only part of the time.During those periods
when the water is not flowing,it may remain in isolated pools or
surface water may be absent.
Classes.--Rock Bottom,Unconsolidated Bottom,Aquatic Bed,
Flat,Streambed,Rocky Shore,Beach/BOar,and Emergent Wetland (non-
pers i stent).
4.LACUSTRINE
Definition.--The Lacustrine System (Figure 6)includes wetlands
and deep-water habitats with all of the following characteristics:
1)situated in a topographic depression or a dammed river channel;
2)lacking trees,shrubs,persistent emergents,nonaquatic mosses or
lichens with greater than 30 percent areal coverage;and 3)greater
than 8 hectares (20 acres)in size.Similar wetlands and deep-water
habitats smaller than 8 ha are also included in the Lacustrine System
if an active wave-formed or bedrock shoreline feature forms all or
part of the boundary,or if the water depth in the deepest part of the
basin is greater than 2 m at low water.Lacustrine waters may be
20
tidal or nontidal,but ocean-derived salinity is always less than
0.5%0'
Limits.--The Lacustrine System is bounded by upland or by wet-
land dominated by trees,shrubs,persistent emergents,nonaquatic
mosses or lichens.Lacustrine systems formed by damming a river
channel are bounded by the contour approximating the normal spillway
elevation or normal pool elevation except where Palustrine wetlands
extend lakeward of that boundary.Where a river enters a Jake,the
extension of the lacustrine shoreline forms the Riverine/Lacustrine
boundary.
Description.--The Lacustrine System includes permanently
flooded lakes and reservoirs (e.g.,Lake Superior),intermittent
lakes (e.g.,playa lakes)and tidal lakes with ocean-derived sal inities
below b.S %
o (e.g.,Lake Maurapas,louisiana).Typically,this
system contains extensive areas of deep water and exhibits considerable
wave action.Islands of Palustrine wetland may lie within the
boundaries of the Lacustrine System.
Subsystems
(1)Limnetic.·This subsystem includes all dee~-w~ter
habitats within the Lacustrine System.Many small Lacustrine Systems
have no Limnetic Subsystem.
(2)Littoral.This subsystem includes all wetland
habitats that fall within the Lacustrine System.It extends'from the
21
shoreward boundary of the system to a depth of 2 m below low water or
to the maximum extent of nonpersistent emergents if these grow beyond
the 2 m depth.
Classes.--Rock Bottom,Unconsolidated Bottom,Aquatic Bed,
Flat,Rocky Shore,Beach/Bar,and Emergent Wetland (nonpersistent).
5.PALUSTRINE
Definition.--The Palustrine System (Figure 7)includes all
nontidal wetlands dominated by trees,shrubs,persistent emergents,
nonaquatic mosses or lichens,and all such wetlands that occur in tidal
areas where salinity due to ocean-derived salts is below 0.5%0'It
also includes wetlands lacking such vegetation,but with all the
following characteristics:1)siz"e less than 8 hectares;2)absence
of an active wave-formed or bedrock shoreline feature;3)water depth
in the deepest part of basin less than 2 m at low water;and 4)
salinity due to ocean-derived salts less than 0.5%0'
Limits.--The Palustrine System is bounded by upland or by any
of the other four systems.
Description.--The Palustrine System was developed to group the
extensive vegetated wetlands traditionally called by such names as
marsh,swamp,bog,fen,and prairie which are found throughout the
country.It also includes small,shallow permanent or intermittent
water bodies,often called ponds.Palustrine wetlands may be situat~d
22
shoreward of lakes,river channels or estuaries;on river flood
plains;in isolated catchments;or on slopes,They may also occur as
islands in lakes or rivers.The erosive forces of wind and water
are of minor importance except in times of severe flood.
The emergent vegetation adjacent to rivers and lakes is often
referred to as lithe shore zone ll or the "zone of emergent vegetat ion l'
(Reid and Wood 1976),and is generally considered a separate community
from that of the river itself.As an example,Hynes (1970:85)says
in reference to riverine habitats,"We wi 11 not here consider the long
list cf emergent plants which may occur along the banks out of the
current,as they do not belong,strictly speaking,to the running
water habitat'"There are often great similarities between wetlands
lying adjacent to lakes or rivers and isolated wetlands of the same
class in basins without open water.
Subsystems.--No subsystems are recognized for the Palustrine
System.
C1asses.--Rock Bottom,Unconsolidated Bottom,Aquatic Bed,
Flat,Moss/Lichen Wetland,Emergent Wetland,Scrub/Shrub Wetland and
Fo res ted Wetland .
23
Classes,.Subclasses and Dominance Types
The CLASS is the highest taxonomic unit below the subsystem
level.It describes the general appearance of the habitat in terms of
either plant life form or physiography and composition of the_sub$~rate,
features which can be recognized without the aid of detailed environ-.
1mentalmeasurements.
Use of life forms at the class level has two major advantages:
I)it does not require a high level of biological expertise to distin-
guish between various life forms,and 2)it has been established that
various 1 ife forms are easi ly recognizable on a great variety of remote
sensing products (e.g.,R~dforth 1962,Anderson et al.1976).If plants
cover more than 30 percent of the substrate,we distinguish classes ()~
the basis of the life form of the plants which constitute the uppermost
layer of vegetation and possess an areal coverage greater than 30 p~rcent.
For example,an area with 50 percent areal coverage of trees over a
shrub layer with a 60 percent areal coverage would be classified as a
Forested Wetland;an area with 20 percent areal coverage of trees over
the same (60 percent)shrub layer would be classified a Scrub/Shrub Wet-
land.Finer differences in life forms are recognized at the SUBCLASS
level.For example,Forested Wetland is divided into Broad-leaved
Deciduous,Needle-leaved Deciduous,Broad-leaved Evergreen,Needle-leaved
lOur attempts to use familiar terms such as marsh,swamp,bog,
and meadow at the class level were unsuccessful primari ly because of
wide discrepancies in t~e use of these terms in various regi~ns of the
United States.In an effort to resolve that difficulty,we decided to
base the classes upon the fundamental componehts (1 ifeform,water
regime,substrate type,water chemistry)which give rise to such terms.
We bel ieve that this approach wi 11 greatly reduce the inisUilderstandirigs
and confusion that result from the use of the common terms.
24
Evergreen,and Dead Subclasses.Subclasses are named on the basis of
the predominant life form.
If plants cover less than 30 percent of the substrate,the
physiography and composition of the substrate are the principal
characteristics used to di~tinguish classes.The nature of the substrate
reflects regional and local variations in geology and the influence of
wind,waves,and currents upon erosion and deposition of substrate
materials.Rocky Shore~have been recognized a~a separate class,also
based on suostrate,even though these habitats may support more than
30 percent cover of macrophytic al~ae.Similarly,we decided to
characterize beaches and flats on the basis ofsLibstrate,although~in
some cases,macrophytic algae or I'pioneer"vegetation may cover more than
30 percent of the substrate.Reefs are a unique class in which the
subst rate i tse If is composed p rima r i I y'of 1 i"i ng and dead an i rna Is.
"Most classes based on substrate have been divided into subclasses
a\Ccon;Hng to the texture or composition of the substrate;for example,
four subcl asses of unconso 1i dated bottoms are recogn i zed:Cobb Ie/Grave 1,
Sand,Mudai1d Organic.In the special case of coral reefs,subclasses
are designated on the basis of the type of or~anlsm that has formed the
reef.
The DOMINANCE TYPE forms the taxonomic category subordinate to
subclass.Dominance types are determined on the basis of dominant plant
species (e.g.,Jeglum et al.1974),dominant sedentary or sessile animal
species (e.g.,Thorson 1957)or dominant plant and animal species (e.g.,
Stephenson and Stephenson 1972)".A dominant plant species has
traditionally meant one that has control over the community (Weaver and
25
Clements 1938:91),and this plant is also usually the predominant
species (Cain and Castro 1959:29).When the subclass is based on life
form we name the dominance type for the dominant species or combination
of species (codominants)in the same layer of vegetation used to deter-
mine the subclass.1 For example,a Needle-leaved Evergreen Forested
Wetland with 70 percent areal coverage of Picea mariana and 30 percent
areal coverage of Larix laricina would be designated as a Picea mariana
Dominance Type.When the relative abundance of codominant species is
approximately equal,the Dominance Type consists of a combination of
species names.For example,an Emergent Wetland with approximately~qual
areal coverage of broad-leaved cattail (Typha latifol ia)and hardstem
bu I rush (Sci rpus acutus)wou Id be des i gnated as Typha 1at i fo 1i a/Sci rpus
acutus Dominance Type.
When the subclass is based on substrate material,the Dominance
Type is named for the predominant plant or sedentary or sessile macro-
invertebrate species without regard for 1 ife form.In the Marlne and
Estuarine Systems,sponges,alcyonarians,molluscs,crustaceans,worms,
ascidians and echinoderms may all be part of the community represented
by the Macoma Dominance Type.Sometimes it is necessary to designate two
or more codomihant species as a Dominance Type.Thorson (1957)has
recommended guidelines and suggested definitions for establishing
community types.and dominants on level bottoms.
Ipercent areal cover will seldom be measured in the application
of this system,but the term must be related to a frame of reference.
We suggest 2m 2 for herbaceous and moss layers,16m 2 for shrub layers and
100m 2 for tree layers (Meuller-Dombois and Ellenberg 1974:74).When per-
cent areal cover is the key for establishing boundaries between units of
the classification,it may be necessary to make cover measurements
occasionally on plots in order to maintain uniformity of ocular estimates
made in the field,or interpretations made from aerial photographs.
26
I.ROCK BOTTOM
Definition.--In the Marine and Estuarine Systems the class Rock
Bottom includes all deep-water (subtidal)habitats with rock substrates.
In the Lacustrine,Palustrine and Riverine Systems,Rock Bottom includes
all wetlands and deep-water habitats with rock substrates and
permanently flooded,intermittently exposed,and semipermanently flooded
W<:lter regimes.This class does not include those habitats classified
as Aquatic Beds.
Description.--The solid rock substrate of the rQcky benthi~or
bottom lone is one of the most important factors in determining the
abundance,variety and distribution of organisms.The stability of the
bottom allows a rich assemblage of plants and animals to develop.Rock
bottoms are usually high energy habitats with well-aerat~d waters.
Temperature,salinity,current and light penetration are also important
factors in determining the composition of the benthic community.Animals
that live on the rocky surface are generally firmly attached by hooking
or sucking devices although they may mOve about over the substrate in
search of food.Some may be permanently attached by cement.A few
<:lnimals hide in rocky crevices and under rocks,some ~ove rapidly enough
to avoid being swept away,and others burrow into finer substrates
between boulders.Plants are also firmly attached (e.g.,by holdfasts)
and,in the Riverine System,they are commonly streaml ined or flattened
in response to high water velocities.
27
Subclasses andDominan~e Types.--Rock Bottom has been divided
into two subclasses,Bedrock and Boulder.The Dominance Types for both
subclasses are similar.
(I)Bedrock.These bottoms consist of stable bedrock
surfaces.Grooves and crevices,when present,provide shelter and
microhabitats.
(2)Boulder.These bottoms consist predominantly of rela-
tivelystable,rock fragments larger than 256 mm (10 in)in diameter
(Wentworth 1922).Often,finer material is mixed with these boulders.
Examples of Dominance Types for the Marine and Estuarine Systems
are Hippospongia encr~sting sponges and Cnemidocarpa~Strongylocentrotus,
Pisaster,Muricea,and Laminaria.Examples of Lacustrine and Riverine
Dominance Types are Spongilla,L¥l1lnaea,Caenis,ChironQmidae,and
Hydrosyche.
2.UNCONSOLIDATED BOTTOM
Definition.--In the Marlneand Estuarine Systems,Unconsolidated
Bottom includes all deep-water (subtidal)habitats with unconsolidated
substrates.In Lacustrine,Palu~trine andR1verine Systems,the class
includes all unconsolidated substrates with permanently flooded,inter-
mittently exposed and semi permanently flooded water regimes.This class
does not include habitats classified as Aquatic Beds.
Description.--Unconsolidated Bottoms are characterized by the lack
of large stable surfaces for plant and animal attachment.They are
usually found in lower energy areas than Rock Bottoms,and may be very
I .,
28
unstable.Exposure to wave and current action,temperature,sa 1 in i ty
and 1 ight penetration .determine the composi.tion and distribution of
organisms.
Most macroalgae attach to the substrate by means of basal hold-
fast cells or discs;however~in sand and mud,algae penetrate the
substrate and higher plants can successfully root if wave action and
currents.are not too strong.The majority of animals in unconsolidated
sed·iments 1 ive wi thin the substrate,e.g.,Macoma and Mellita.Some,
such a~Chaetopterus,maintain permanent burrows,and others may live on
the surface,especially in coarse-grained sediments.
In the Marine and Estuarine Systems,Unconsolidated Bottom com-
i munities are relatively stable.They vary from the Arctic to the tropics,
depending largely on temperature,and from the open ocean to the upper
end of the estuary depending upon sal inity.Thorson (1957)has
summarized and described characteristic types of level bottom communities
in detail.
In the Riverine System,the substrate type is,to a great extent,
determined by current velocity,and plarit~alid animals exhibit a high
degree of morphologic and behavioral adaptation to flowing water.Some
species are confined to specific substrates and others are at least more
abundant in one type of substrate thanth-ey are in others ..AccO ....dirigto
Hynes (1970:208),liThe larger the stones,and hence the more complex the
substratum,the more diverse is the invertebrate fauna.1I In Lacustrine
and Palustrine Systems,there is usually a high correlation,within a
given watl body,between the nature of the substrate and the number of
species and individuals.For example,in the profundal bottom of
"29
eutrophic lakes where 1i ght ·is absent,oxygen contenti s low and carbon
dioxide concentration is high,the sediments are ooze-like organic
materials and species diversity is low."Each substrate type typically
supports a relatively distinct community of organisms (Reid 1961:307).
Subclasses and Dominance Types.--The class Unconsol idated Bottom
has been divided into four subclasses:Cobble/Gravel,Sand,Mud,and
Organi~.Differences in grain size and interstitial spa~e in unconsoli-
d<:lted substrates greatly affect the species composition of the benthic
flora and fauna.
(1)Cobble/Gravel.The substrate is predomina~tly cobble
and gravel although finer sediments may be intermixed.An example of a
Dominance Type fbr theMa~ine System is M6diolus and fot the Estuarine
System,Dendraster.Examples for the Lacustrine,Palustrine and Riverine
Systems are Diamesa~Nemoura/Eukiefferiella (Slack et al.1977).
Chironomus/Hydrosyche/Physa(Krecker and Lancaster 1933),Limnea,Baetis,
Spongilla,Lumbriculus,and Gammarus.
(2)Sand.The substrate is predominantly sand,although
finer or coarser sediments may be intermixed.The Sand Bottom has a
more limited fauna and flora th~n either Mud or Cobble/Gravel Bottom.
Examples of Dominance Types in the Marine System are Pecten,Tellina,
Penaeus,and Spatangus and for the Estuarine System,Tellina,Ar~nicola,
Dendraster,and Renilla.Examples for the Lacustrine,Palustrine and
Riverine Systems are Physa,Gammarus,Chironomidae,Limnodrilus,and
Ephemera.
30
(3)Mud.The substrate is predominantly silt and clay
although coarser sediments or organic material may be intermixed.
Organisms 1 iving in mud must often be able to adapt to rather low
oxygen concentrations.Exa~ples of Dominance Types for the Marine and
Estuarine Systems include Amphiura,Macoma,Echinocardium,and Urechis.
Examples of Dominance Types for the Lacustrine,Palustrine and Riverine
Systems are'Tubifex~Anodonta,Pisiduim,Chaoborus,~ndC~ironomus.
(4)Organic.The substrate is predominantly composed of
organic material.These habitats have a limited number of species and
are very low in fa.una 1 product iv I ty (We 1ch 1952).
3.AQUATIC BED
Defiriition.--The cl~ss Aquatic Bed represents wetlands and deep-
water habitats that the major i tyof the time are dominated by submergent
plants,floating-leaved plants or floating plants for the majority of
the growing season in most years.Water regimes are restricted to sub-
tidal,i rregu 1ar 1y exposed,permanent ly flooded,i nterm i ttent 1y exposed,
and semi permanently flboded.
Description.--Aquatic Beds represent a diverse group of plant
communities that require surface water for optimu~growth'and repro-
duction.They are best developed in relatively permanent water.
Subclasses and Dominance Types
(1)Submergent Algal.These habitats occur in both tidal
and nontidal locations,but they are far more diverse and widespread in
the Marine and Estuarine Systems.In these coastal areas,algal beds
31
oGcuPY rock substrates and unconsol idated substeates characterized by a
wide range of sediment depths and textures.They may extend to ~ater
depths of 30m (98 ft).Coastal algal beds are most luxuriant along the
rocky shores of the northeast and the west.Macrocysti s beds are
especially well-developed on the Pacific Coast.Along both coasts,
Fucus and Laminaria may dominate dense SubrTlergent Algal Beds.In
~ropical regions,this subclass is characterized by green algae,includ~
in~forms containing calcareous particles;Halimeda and Penicillus are
common examples.Caulerpa and Laurencia also may form large Submergent
Algal Beds.Other plants,such as Enteromorpha and Ulva,are tolerant
of fresh water and flourish in some estuaries.
Inland SObmergent Algal Beds are represented by plants such as
the Chara and Nitella which look much 1 ike vascular plants and may grow
in similar situations.However,Chara meadows may be found in Lacustrine
waters as deep as 40 m (131 ft)(Zhadin and Gerd 1963),where hydrostatic
pressur~limits the survival of vascular submergents (Welch 1952).
(2)Submergent Vascular.In the Marine and Estuarine
Systems this subclass has been referred to by others as temperate grass
flats (Phillips 1974);tropical marine meadows (Odum 1974);eelgrass beds,
tu rt legrassbeds and seagrass beds (Aki ns and Jefferson 1973,EI euter ius
1973,Phillips 1974).Submergent Vascular Beds extend to depths greater
Lhan 10 m (33ft)In clear marine waters.The greatest numbers of
plant species occur in shallow,clear,tropical or subtropical waters of
moderate current strength in the Caribbean and along the Florida and
Gulf Coasts.Principal Dominance Types in these areas include Thalassia
testudlnum~Halodule beaudettei,Syrin~odJumfiliformis,R~p~ia maritima,
32
~aJophila and Vall isneria americana.
Five major species dominate the Submergent Vascular Beds along
the temperate coasts of North America:Halodule beaudettei,Phyllospadix
scouleri,~.torreyi,Ruppia maritima and Zostera marina.Zostera beds
have the most extensive distribution,but they are limited primarily to
the more sheltered estuarine environment.In the lower sal inity zones
of estuaries,stands of Ruppia,Potamogeton and Vallisneria often occur,
along with Najas and
Submergent Vascular Beds in the Riverine,Lacustrine and
Palustrine Systems occur at all depths within the photic zone.Typical
inland genera include Potamogeton,Ceratophyllum,Myriophyllum,Najas,
Ruppia,Utricularia and Vallisneria.
(3)Submergent Moss.These Aquatic Beds are far less
abundant than Algal or Vascular Beds.They occur primarily in the
Riverine System and in permanently flooded and intermittently exposed
parts of some Lacustrine Systems.The most important Dominance Types
include genera such as Fissidens,Drepanocladus and Fontinalis.The
latter may grow to depths as great as 120 m (392 ft)(Hutchinson 1975).
For simpl icity,aquatic liverworts of the genus Marsupella are included
inth1s subclass.
(4)Floating-leaved.These Aquatic Beds are characterized
by floating-leaved plants.They are found in all systems except the
Marine.These beds typically occur in sheltered areas where there is
little water movement (Wetzel 1975).Typical dominants include Nymphaea,
Nuphar,Potamogeton natans and Brasenia schreberi.Plants such as
33
Nuphar advena andPolygorium amphibium,which may stand er~ctabove the
water surface or substrate.,may be considered emergentsorfloating-Ieaved
plants,depending upon·the life form adopted at a particular site.
(5)Floating.Thi~subclass is chsracterized by ~~nera
which float freely on the water surface,such as:Lemna,Spirodela,
Pistia,Eichhornia,Trapa,Salvinia and Azolla.These plants are
found primarily in protected portions of slow-flowing rivers and in
the Lacustrine and Palustrine Systems."Floating Beds Sre dynamic
habitats;they are easily moved about by wind or water currents.They
COver a large area of water in some parts of the country,particularly
the southeast.
4.REEF
Definition.--The class Reef"includes ridge-or mound-like
structures formed by the colonization and growth of sedentary
invertebrates.
Description.--Reefs are characterized by their elevation above
the surrounding substrate and their interference with norm~l wave flow;
they are primarily subtidal,but parts of some Reefs ~ay be intertidal
as well.Although corals,oysters and tubeworms are the most visible
organfs~s arid are mainly resporisible for Reef formation,other molluscs,
foraminifera,corral I ine algae and other forms of life also contribute
substantially to Reef growth.Frequently,Reefs contain an abundance
of dead skeletal material and shell fragments,in comparison to the
amount of living matter.
Subcla$ses andDo~1nance Types
(I)Coral.·Coral Reefs are widely distributed in shallow
waters of warm sea.s.They are found in Hawaii,Puerto Rico,the Virgin
Islands,and southern Florida.They ar~characterlzed by Odum(1971)
as stablE~,well-adapted,highly diverse and highly productive ecosystems
with a great degree of internal symbiosis.Coral Reefs lie almost
entirely within.the Marine Subtidal Subsystem;although the upper part
of some reefs is sometimes exposed.Examples of Dominance Types are
Porites,Acropora and Montipora;the distribution of these types
primarily reflects elevation,wave exposure,and the age of the Reef.
(2)Mollusc.This subclass occurs in both the Estuarin~
Intertidal and Subtidal Subsystems.These Reefs are found on the
Pacific,Atlantic,and Gulf Coastsand.in Hawaii and the Carribbean.
Mollusc Reefs may become extens ive ,affo·rd i ng.a substrate for sedentary
and boring organisms.arid a shelter for many others.Reef molluscs are
adapted to great variations in water level,salinity and temperature,
and these same factors control their distrib~tion.~xamples of
Dominance Types fbr this subclass are the oysters (Ostrea,trassostrea).
(3)Worm.Worm·Reefs are constructed by large c610nies of
sabellari id worms living in individual tubes constructed from cemented.
sand 9rains.Although they do not support as diverse a biota as do
Coral and Mollusc Reefs,they provide a distinct habitat which may
cover large areas.Worm Reefs are generally confined to tropical waters,
and are most cpmmon along the coasts of Florida,Puerto Rico,and the
Virgin Islands.They occur in both the Marine and Estuarine Systems
35
where the salinity approximates that .of sea water.The Dominance Type·
for this subclass isSabellarta.
5.FLATS
Definition.--The class Flat refers to level landforms composed of
unconsol idated sediments.Normally,Flats occur only in areaS sheltered
from strong currents "and wave action.They may be irregularly shaped or
elongate and continuous with the shore,whereas Bars generally are
elongate,parallel to the shorel ine,and separated from the shore by
water.Water regimes are restricted to irregularly ~xposed,regularly
flooded,irregularly flooded,seasonally flooded,tempor-arily flooded
and intermittently flooded •
.Description •...;-Estuarineand Marine Flats occur in the intertidal
zone.The distrib~tion of fauna is dependent on substrate texture,
current and wave action,and salinity;temperature and salLnity may be
extremely variable.Regularly flooded Flats support diverse populations
of tube-dwell ing and burrowing invertebrates including worms,clams
and crustacearis (Gray 1974).These invertebrates are mostly detritus
feeders.Irregularly flooded Flats have been called salt flats,pans or
pannes.They are typ i ca 11y highi n sal i h"ity and are usually surrounded
by,or lie on the landward side of,Emergent Wetland (Martin et al.1953,
,
Type 15).Flats are also commonly colonized by algae and diatoms and
there may be an algal crust or mat.
The distribution of organisms in Riverine and Lacustrine Flats is
.dependent upon substrate material,currelit and wave action,and the
36
frequency of inundation.Lacustrine Flats may include the entire basin
of a lake.Palustrine Flats are generally the result of high salinity,
removal of vegetation by man,animals,or fire,or the discharge of
thermal waters or pollutants.In manOy arid areas,Palustrine and
LacuStrine Flats are crusted or saturat~d with s~lt.Martin et al.
(1953)called these habitats iniand saline flats (Type 9);they are also
called alkali flats,salt flats,and salt pans.°Faunal diversity and
abundance caries with salinity,duration of inundation and temperature.
Subclasses and Dominance Types
(1)Cobble/Gravel.The Flats are-composed predominantly of °
cobbles or gravel,often with shell fragments or finer sediments inter-
mixed.Unlike Rocky Shores,Cobble/Gr~vel Flats are n6t ~table and
communities aroe more transitory.Examples of Dominance Types in the
Mar i ne and Estuari ne Systems are:Sa lanus,Patella ,Li nor i na,Thai s,
and Myt il us •
(2)Sand.Sand Flats are composed predominantly ~f ~and,
°often with particles of other sizes intermixed.Although population
density may be very high,species diversity is usually comparatively low.
In the Marine and Estuarine SyStems,some examples of Dominance Types are
~,Macoma,Tellina,Arenicola,Uca,Leptosynapta,and Paractis.
(3)Mud.Mud Fl ats are composed predomi nant 1y of s i It and
clay,and tend to be anaerobic below the surface.They usually have a
higher organic content than Cobble/Gravel and Sand Flats.In the Marine
and Estuarine Sy~tems some examples of Dominance Types are:Uca,
Callianassa,Nassari~s~Macoma,N~reis,Amphitrite,Cerianthiopsis and
Thyone.
..
37
(4)Organic.These Flats consist of exposed organfc soils
of former.y vegetated wetlands;In the Estuarine System,Organic Flats
are often dominated by rriicroinvertebrates such as foraminifera or smaller
snails (Cerit~ium).
(5)Vegetated Flats.Some Flats are exposed for a
sufficient period to be colonized by herbaceous annual~or seedling
herbaceous perennials (pioneer plants).When these plants cover more
than 30 percent of the substrate,the area is classified as a Vegetated
Flat.This vegetation is usually ki lIed by rising water levels and may be
removed before the begfnning of the next growing season.Examples of
Domfnance Types are Xanthium italicum,Cheno~od~um rubrum,Ec~inochlo~
crusgalli,and Eleocharis acicularis.
6.STREAMBED
Definition.--The class Streambed is restricted to the Riverine.
and Estuarine Systems.I·t includes all p~rts of channels that are not
included in any of the oth~r classes.Streambed may have the classes
Beach/Bar and Flat included within it,but in the Intermittent Subsystem
of the Riverine System the entire channel frequently contains only the
c I ass S tr'eamb~d.
Description.--Streambedsvary greatly in substrate and form
depending on the gradient of the channel and the velocity of the water.
In the Rfveririe Sys~em,material on the bed is continually being moved
downstream;in Estuarine streambeds,material may be moved upstream or
downstream according to the direction of tidal flow.Frequently Bars
occur on the convex side of single channels or they may be included as
38
islands within the bed of braided streams (Crickmay 1974}.Flats and
Beaches are particularly common adjacent to Streambed,particularly in
the Lower Perennial and Tidal Subsystems of larger rivers and in estuarine
streams where wave action is especially strong.In estuarine areas,
Streambed is the proper class for the entire channel of tidal creeks that
are dewatered at low tide.In most cases,Streambeds are not vegetated
because of the scouring effect of the moving water,but,I ike Flats,they
may be colonized by "p ioneering"annuals during periods of low flow or
they may have perennial emergents and shrubs that are too scattered to
classify as Emergent Wetland or Scrub/Shrub Wetland.
39
13)Mud.This subclass is characteristic of the Lower
Perennial and Tidal Subsystems of the Riverine.Mud Streambeds are
frequently associated~ith l~rge Mud Flats~
(4)Organic.This is not a common sub~lass in the Riverine
System but it occurs along the edge of streams flowing over deep peat
deposits,usually in the Lower Perennial Subsystem.Organic Streambeds
are common in creeks draining Estuarine Einergent Wetlands with orga"nic
soi Is.
7.ROCKY SHORE
Definition.--The class Rocky Shore includes wetland environments
characterized by stable bedrock surfaces or relatively stable,large rock
fragments.Water regimes ate ~e~tticted to irregularly exposed,
regul ar Iy flooded,i rregu 1ar 1y flooded,seasona I ly flooded,temporar i ly
flooded and intermittently flooded.
Description.--I~Marine and Estuarine Systems,Rocky Shores are
generally high energy habitats which lie exposed as a result of
continuous erosion by wind-driven waves or strong currents.The substrate
is stable enough to permit the attachment and growth of sessile or
sedentary invertebrates and attached algae or lichens.Rocky Shores
usually display a vertical zonation which is a function of tidal range,
wave action and degree of exposure to the sun.In the Lacustrine ahd
Riverine Systems,Rocky Shores support sparse plant and animal communities.
SUbclasses and Dominance Types.--The class Rocky Shore is divided
into two subclasses,Bedrock and Boulder.The Dominance Types are
40
similar for both subclasses.
(1)Bedrock.These wetlands consist of stable bedrock
surfaces ..
(2)Boulder.These wetlands consist of relatively stable
rock fragments larger than 256 mm in diameter.
Communities or zones of Marin~ana Estuarine Rocky Shores have
been studied in detail (Lewis 1964,Ricketts and Calvin 1968,Stephenson
and Stephenson 1972).Each zone supports a rich assemblage of
invertebrates and algae.Dominance Types of the Rocky Shore often can be
characterized by one or two dominant genera from these zones.
The uppermost zone (termed the Littorine/Lichen Zone)is dominated
by periwinkles (Littorina and Nerita)and lichens.This zone frequently
takes on a dark,or even black appearance,although abundant lichens may
lend a colorful tone.These organisms are rarely submerged.but are kept
moist by sea spray.Frequently,this habitat is invaded from the
landward side by semi-marine genera such as Lygia.
The next lower zone (the BalanoidZone)is commonly dominated by
molluscs,green algae and barnacles of the bi:l1anoid group.From a
distance,the zone appears white~Dominance Types such as BalanUS,
Chthamalus and Tetrac1ita may form an almost pure sheet of barnacles,or
these animals may be interspersed with molluscs,tubeworms and algae
such as Pelvetia.
The transition between the littorine/lichen and balanoid zones is
frequently marked by the replacement of the periwinkles with both true and
false 1 impets such as Acmaea and Siphonaria.The limpet band approximates
the upper limit of the regularly flooded intertidal zone.
41
In lhc middle and lower intertidal areas,which are flooded and
exposed by tides at least once daily,lie a number of other communities
which can be characterized by dominant genera.Mussels (Mytilus)and
gooseneck barnacles (Mitella)form communities exposed to strong wave
action.The Fucus and Laminaria Dominance Types liesl ightly lower,
just above the coralline algae (Lithothamnion)Dom.inance Type.The
Laminaria Domin~nce Type approximates the lower end of the Intertidal
Subsystem;it is generally exposed at least once daily.The Lithothamnion'
Dominance Type forms the transition to the Subtidal Subsystem and is
exposed only i~regularly.
We have not identifled DomlnanceTypes foithe Lacustrine and
Riverine Systems.
8.BEACH/BAR
Definition.--The class Beach/Bar consists of sloping landforms
generated by waves and currents and composed predomi nant lyof unconso 1 i-.
dated sand,gravel or cobbles.'They have less than 30 percent vegetative
cover.Beaches are generally continuous with the shore,extending
landward to a distinct break in landform ~r substrate type (e.g.,a
foredune,cl iff or bank)or to the point where vegetation covers 30 per-
cent or more of the substrate.Bars are elongate ridges,banks or mounds
which are bordered by water on at least two sides.Water regimes are
restricted to irregularly exposed;regularly flooded,irregularly fiooded,
seasonally flooded,temporarily flooded and intermittently flooded.
Description.--Beaches and Bars are characterized by a shifting,
unstable substrate with high permeability~variable surface moisture,
42
and relatively low organic niatter content.The surface layer has a high
oxygen content and there is a deeper anaerobic layer (Hedgpeth 1957,
Ranwell 1972).They may be sparsely vegetated and populated by a
diversity of specialized burrowing invertebrates such as molluscs,
crustaceans and polychaetes.Faunal distribution is controlled by waves,
currents,interstitial moisture,sal inity and sediment grain size.In
Marine and Estuarine Beaches and Bars the fauna is usually more diverse.
than in Lacustrine Of Riverine Beaches and Bars (Hedgpeth 1957,Riedl
and McMahan 1974).
Subclasses and Dominance Types
(I)Cobbl~/Gravel.Cobble/Gravel Beaches and Bars
typically form where wave action is especially strong so that sand .and
silt particles are largely eroded and .transported from the Beach or Bar
and ~eposited in deeper waters.While cobbles and gravel predominate,
sand is usually mixed with these larger particles.Some of the larger
cobbles and occasional boulders found on these Beaches may support
sedentary organisms such as Balanus.
(2)Sand.Sand Beaches and Bars are composed predominantly
of calcareous or terrigenous sand.Examples of Dominance Types in the
Marine and Estuarine Systems are Donax,Mya,Tivela,Oliva,Thoracophelia,
Haustorius,Orchestia,Chiredotea,EmeritajOcypode (Hedgpeth 1957,Riedl
and McMahan 1974).Examples of Dominance Types in the Riverine,
Lacustrine,and PalOstrine Systems are Parastehocaris,Phyllognathus,and
Pristina (Neel 1948).
43
9.MOSS/LICHEN WETLAND
Definition.--The Moss/Lichen ·Wetland class includes areas where
mosses or I ichens cover the substrate and where other plants such as
emergents,shrubs or treestomprise less than·30 percent of the areal.
cover.
Description~-~Mosses and lichens are important components of the·
flora in many wetlands,espetially in the north,but these plants
usually form a ground cover under a dominant layer of trees,shrubs or
emergents.In some instances higher plants a re uncommon and mosses or
lichens dominate the flora.Such Moss/Lichen wetlands are not common,
even in the northern United States where they occur ~ost frequently.
Subclasses abd Dominance Types
(I)Moss.These wetlands are most abundant in the far
north.Areas.covered wi th Sphagnum spp.are usual 1y ca lIed bogs (Go let
and Larson 1974,Jeglum et al.1974,Zoltai et al.1975),whether
Sphagnum or higher plants are dominant.In Alaska,mosses such as
Drepanocladus and the liverwort Chiloscyphus fragilis may dominate shallow
pools with impermanent water;Sphagnum,Campyl ium stellatum,Desmatodon
heimi i,Aulacomnium palustre and Oncophorus wahlenbergii are typical of
wet soil in this region (Britton 1957,Drury 1962).
(2)Lichen.These wetlands are also a northern subclass.
Cladonia forms the most important Dominance Type.Pollett and Bridgewater
(1973)described areas with mosses and lichens as bogs or fens,the
distinction being based on the availabhity of nutrients and the
44
panicularp1ant species present.Sj8rs (1959)and Jeg1um et a1.(1974)
mentioned the presence of Lichen Wetlands in the Hudson Bay Lowlands and
in Ontario,respectively.
10.EMERGENT WETLAND
Definition.--The Emergent Wetland class is characterized by erect,
rooted~herbaceous hydrophytes,excluding mosses and lichens.This
vegetation is present for most of the growing season in most years.
These wetlands are usually dominated by perennial plants.
Description.--In areas with relatively stable climatic conditions,
Emergent Wetlands maintain the same appearance year after year.In other
areas,such as the prairies of the central United States,violent
cl imatic fluctuations cause Emergent Wetlands to revert to an open water
phase in some years (Stewart and Kantrud 1972).They are found throughout
the United States and occur in all systems except the Marine.Emergent
Wetlands are known by many names including marsh,meadow,fen,prairie
pot hote,and slough.Areas that are dominated by pioneer plants that
become estab1 ished during periods of low water are not Emergent Wetlands
~nd should be classified as Vegetated Flats.
Subclasses and Dominance Types
(1)Persistent.Wetlands in this subclass are dominated by
species that normally remain standing at least until the beginning of the
next growing season.This subclass is found only in the Estuarine and
Palustrine Systems.
45
Persistent Emergent Wetlands dominated bySpartina alterniflora J
S.patens,.?:cynosuroides J Juncus roemerianusJTypha angustifolia,and
Zizaniopsis mili.:kea are major components of the Estuarine Systems of
the Atlanti~and Gulf Coasts of the United States.On the Pacific Coast,
Sal icornia pa~ifica,Suaeda cal ifornicaJ Triglochin maritima and Spartina
foliosa are common dominants.
Palustrine Persistent Emergent Wetlands contain a vast array of
grass-like plants such as Typha spp.,Scirpusspp.,Cladium jamaicens
Carex spp.,and true grasses such as Phragmites communis,Glyceria spp.,
Beckmania syzigachne and Scolochloa festucacea.There is also a variety
of broad-leaved persistent emergentssuch as Lythrum salicaria,Rumex
mexicanus,Decodonvertfcillatus,and many species of Polygonum.
(2)Nonpersistent.Wetlands in this subclass are dominated
by plants which fall to the surface of the substrate or below the surface
of the water at the end of the growing season so that J at certain seasons
of the year,there is no obvious trace of emergent vegetation.For
example,Zizania aquatica in the north central states does not become
apparent until mid-summer and fail when it forms dense emergent stands.
Nonpersistent emergents also include species such as Peltandra virginica,
Pontederia cordata and Sagittaria spp.Movement of ice in Riverine and
Lacustrine Systems often removes all traces of emergent vegetation during
the winter.Where this occurs,the area should be classified as Non-
persistent EmeigentWetland.
46
11.SCRUB/SHRUB WETLAND
befinition.--The class Scrub/Shrub Wetland includes areas domi-
nated by woody vegetation less than 6 mm (20 ft)in height.The species
include true shrubs,young trees,and trees or shrubs that are small or
stunted betauseof environmental conditions.
Descripti6n.--Scr~b/$hrubWetlands may represent a successional
stage leading to Forested Wetland,or they may be relatively stable
communities.They occur only in the Estuarine and Palustrine Systems,
but are one of the most widesp"read classes in the country (Shaw and
Fredine 1956).Scrub/Shrub Wetlands are known by many names such as
shrub swamp (Shaw and Fredine 1956),shrub carr (Curtis 1959),bog
(Heinselman 1970),and pocosin (Kologiski 1977).For practical reasons
we have also included forests composedbf young trees lessthah 6 m tall
in this class.
Subclasses and Dominance Types
(1)Broad-leaved Declduous.These are wetlands where
deciduous trees or shrubs less than 6 m high represent more than 50 per-
cent of the total areal cover.In the Estuarine System they are
dominated by such plants as Baccharis halmifolia,and Iva frutescens.
Alnus spp.,Salix spp~,Cephal~nthus occidental is,Zenobia pulverulenta,
BcLulapulilila and young trees such as Acerrubrum are typical Dominance
Types in the Palustrine System.
(2)Needle-leaved Deciduous.This subclass is found only in
the Palustrine System where it is represented by young or stunted trees
47
such as Larix laricina or Taxbdium distichium.
(3).Broad-leaved Evergreen.These wetlands are found in
both the Estuarine and Palustrine Systems.In the Estuarine System,vast
acreages are dominated by mangroves (R~izophora mangle,~anguncularia
racemose,Conocarpus erecta,Avicennia germinans)~h~t are less than 6 m
in height.In the Palustrine System,the broad-leaved evergreen species
are typically found on organic soils.In the north,typical representa-
·tives are Ledum groenlandicum,Andromeda glaucophylla,Kalmia polifolia,
~nd the semi-evergreen Chamaedaphnecalyculata.In the south,Lyon1a
lucida and Leucothoe axillaris are characteristic broad-leaved evergteen
species.
(4)Needle-leaved Evergreen.These wetlands are found only
in the PalUstrine System.The dominant species are young or stunted
trees such as Pice~mariana or Pinus setotina.
(5)Dead.These wetlands ~redominated by dead woody
vegetation less than 6 m in height.They are usually produced by a
prolonged rise in the water table,which often results from the activities
of man or beaveis,but they may result from other natural causes such as
fire,salt spray which often accompanies severe coastal storms,and
insect infestations;and from air pollution or man's use of herbicides.
12.FOR~ST~D WETLAND
Definition.--The class Forested Wetland is characterized by woody
vegetation that is 6 m or more in height.
Descriptibn.--Forested Wetlands are most common in the eastern
United States and in those sections of the west where moisture is
----------~
48
relatively abundant,particularly along rivers and in the mountains.
They occur only in the Palustrine and Estuarine Systems.Palustrine
Forested Wetlands normally possess an overstory of trees,an understory of
young trees or shrubs and an herbaceous layer.Forested Wetlands in the
Estuarine System are restricted to the mangrove forests of Florida,
Puerto Rico and the Virgin Islands.
Subclasses and Dominance Types
(1)Broad-leaved Deciduous.These wetlands are found
throughout the country,but reach their greatest abundance in the south
and east.Dominant trees typical of this subclass include:Acer rubrum,
Ulmus americana,Fraxinus pennsylvanica,~.nigra,Nyssa sylvatica,~.
aquatica,Quercus bicolor,..9:..lyrata,and Q.michauxii.Wetlands in
this subclass generally occur on mineral soils or highly decomposed
organic soils.
(2)Needle-leaved Deciduous.These wetlands are represented
by a limited group of species.Larix laricina is characteristic of the
Boreal Forest Region where it occurs as a dominant on organic soils.The
southern representatives of this subclass include Taxodium distichum and
T.ascendens which are noted for their abil ity to tolerate long periods of
surface inundation.
(3)Broad-leaved Evergreen.These wetlands reach their
greatest development in the southeast where Persea borbonia,Gordonia
lasianthus and Magnolia virginiana are prevalent,especially on organic
soils.This subclass also includes Rhizophora mangle,Avicennia nitida
and Laguncularia racemosa which are adapted to varying levels of salinity.
49
(4)Needle~leaved Evergreen.These wetlands are widespread
in the north where Picea mariana,growing on organic soils,represents a
major dominant.Whereas~.mariana is common on nutrient-poor soils,
Thuja occidentalis dominates northern wetlands on more nutrient-rich
sites.Along the Atlantic Coast,Chamaecyparis thyoides is one of the
most common dominants on organic soils.Pinus serotina is a common
needle-leaved evergreen found in the southeast in association with dense
stands of broad-leaved evergreen and deciduous shrubs.
(5)Dead.These wetlands are dominated by dead woody
vegetation greater than 6 m in height.Like Dead Scrub/Shrub Wetlands,
they are most common In,or around the edges of,man-made impoundments
and in beaver ponds.The same factors that produce Dead Scrub/Shrub
Wetlands also produce Dead Forested Wetlands.
MODIFIERS
In order to fully describe wetlands and deep-water habitat~,
certain modifiers must be appl ied at the class level and at lower levels
in the classification hierarchy.The modifiers described below were
adapted from existing classifications or were developed specifically for
this system.Modifiers for water regime,water chemistry,soil,and
special modifiers for disturbed wetlands are defined in this section.
Waie~Regime Modifiers
Precise description of hydrologic parameters requires detailed
knowledge of the duration and timing of surface inundation on both a
yearly and long-term basis,as well as an understanding of ground-water
50
fluctuations.Because such information is seldom available,the water
regimes that,in part,determine characteristic wetland and deep-water
plant and animal communities are described here in only general terms.
Water regimes are defined under two major headings,Tidal and Nontidal.
~iheseterms are not intended as modifiers.
Tidal water regime modifiers should be used for wetlands and
deep-water habitats in the Estuarine and Marine Systems and nontidal
modifiers should be used for all nontidal parts of the Palustrine,
Lacustrine and Riverine Systems.The Tidal Subsystem of the Riverine
System and tidally-influenced parts of the Palustrine and Lacustrine
Ecological Systems require careful selection of water regime modifiers.
Subtidal and irregularly exposed wetlands and deep-water habitats.in the
Palustrine,Riverine,and Lacustrine Systems should be called
permanently flooded-tidal rather than subtidal.Palustrine,Riverine,
an9 Lacustrine wetlands regularly flooded by the tide should be
\
designated regularly flooded.Palustrine,Riverine and Lacustrine wet-
lands which are only irregularly flooded by tides should be designated
by the appropriate nontidal water regime modifier with the word tidal
added,as in seasonally flooded-tidal.
1.TI DAL:
The water regimes are largely determined by oceanic tides.
Subtidal.--The substrate is permanently flooded with tidal water.
Irregularly Exposed.--The land surface is exposed by tides less
often than daily~
..
~'.
-----~~~------~~---~-~-----------~
51
Regularly Flooded.--Tidal water alternately floods and exposes the
land surface at least once daily.
Irregularly Flooded.--Tidal water floods the land surface less
often than daily.
The periodicity and amplitude of tides vary in different parts of
the United States,mainly because of differences in latitude and
geomorphology.On the Atlantic Coast,two approximately equal high tides
are the rule (semidiurnal).On the Gulf Coast,there is frequently only
one high tide and one low tide each day (diurnal).The usual pattern on
the Pacific Coast is two unequal high tides and two unequal low tides
(mlxed semidiur~al).
Individual tides range in height from approximately 9.5 m (31 ft)
at St.John,New Brunswick (U.S.National Oceanic and Atmospheric
Administration 1973)to less than 1 m (3.3 ft)along the Louisiana coast
(Chabreck 1972).Tides of only 10 cm (25 in)are not uncommon in
Louisiana.Therefore,while there can be no hard and fast rules,the
division between regularly flooded and irregularly flooded water regimes
would probably occur approximately at:mean high water on the Atlantic
Coast,lowest higher high tide on the Pacific Coast,and just-above mean
tide level of the Gulf Coast.The width of the intertidal zone is
determined by the tidal range,the slope of the shoreline,and the degree
of exposure of the site to wind and wave.
2.NONT I DAL
These water regimes are not influenced by oceanic tides,although
they may be affected by wind or seiches in lakes.Water regimes are
--------------------------------------------
52
--------------------------------------------------
defined in terms of the growing season which we equate to t'he frost-free
period (see the U.S.Department of Interior National Atlas 1970:11 for
generalized regional delineation).The remainder of the year is defined
as the dormant season,a time when even extended periods of flooding may
have little influence on the development of plant communities.
Permanently Flooded.--Water covers the land surface at all times
of the year in all years.Vegetation is composed of obligate hydrophytes.
Intermittently Exposed.--Surface water is present throughout the
year except in years of extreme drought.
Semi permanently Flooded.--Surface water persists throughout the
growing season in most years.When surface water is absent,the water
table is usually at,or very near,the land surface except during severe
drought.
Seasonally Flooded.--Surface water is present for extended periods
early in the growing season,but is absent by the end of the season in
most years.When surface water is absent,the water table is usually
near the land surface.
Saturated.--The substrate is saturated to the surface for
extended periods during the growing season,but surface water is seldom
present.
Temporarily Flooded.--Surface water is present for brief periods
during the growing season,but the water table usually lies well below
the soil surface for most of the season.Plants that grow both in uplands
53
and wetlands are characteristic of the temporarily flooded regime.
f~ter~itteritly Flooded.--The substrate is usually exposed,but
surface water is present for variable periods without detectable seasonal
periodicity.Weeks,months,or even years may intervene between periods
of inundation.The dominant plant communities under this regime may
change as soil·moisture conditions change.Some areas exhibiting this
regime do not fall within our definition of wetland because they do not
have hydric soils or support hydrophytes.
Artificially Flooded.--The amount and duration of flooding is
under the direct and purposeful control of man;it does not depend just on
normal variatiori in precipitation or riverflow.The vegetation growing
in these areas cannot be considered a reliable predictor of water regimes.
Agricultural l~nds managed under a rice-soybean rotation and wildlife
.management areas where crops or forests may be flooded or dewatered to
attract wildlife are typical of this regime.
Water Chemistry Modifiers
The accurate characterization of water chemistry in wetlands and
deep-water habitats is a very difficult task,both because of problems in
meastirement and because values tend to vary wtth changes in the season,
weather,time of day and other factors.Yet,very subtle changes in
water chemistry,which occur over short distances,may have a dramatic
influence on the types of plants or animals that inhabit an area.We
feel that a description of water chemistry must be an essential part of
this classification system.
The two key parameters employed in this system are sal inity and
hydrogen ion concentration (pH).All habitats are classified according to
salinity,and fresh-water habitats are further subdivided by pH levels.
I.SALINITY MODIFIERS
Differences in sal inity are reflected in the species composition
of plants and animals.Many authors have suggested using biological
changes as the basis for subdividing the salinity range between sea water
and fresh water (Remane and Schlieper 1971).Others have suggested a
similar subdivision for the inland salinity spectrum (Moyle 1946,Bayly
1967,Stewart and Kantrud 1971).Since the gradation between fresh and
hypersaline or hyperhaline waters is continuous,any boundary will be
artificial,and few classification systems agree completely.In inland
environments,salinity is usually expressed in specific conductance,
milligrams per liter (mg/l),milliequivalents per liter (meq/l),percent
salt or total dissolved solids.
Estuarine and marine waters are a complex solution of salts;these
are dominated by sodium chloride (NaCl).The term haline is used to
indicate the domihance of ocean salt.The relative proportions of the
various major ions are usually similar to those found in sea water,even
if the water is diluted below sea strengtho Dilution of sea water with
fresh water and concentration of sea water by evaporation result in a wide
range of recorded salinities in both surface water and interstitial (soil)
water.
We have modified the Venice System,suggested at a IISymposium on
the Classification of Brackish Waters"in 1958,for use in the Marine and
55
Estuarine Systems (Table I).The system has been widely used during
recent years (Macan 1961,1963,Burbank 1967,Carriker 1967,Reid and
Wood 1976),although there has been some criticism of its applicability
(den Hartog 1960,Price and Gunter 1964).
The salinity of inland waters is dominated by four.major cations:
calcium (Ca),magriesium (Mg),sodium (Na),and potassium (K),and the
major anions:carbonate (C0 3),sulfate (S04)and chloride (Cl)(Wetzel
1975).Salinity is governed by the interactions between precipitation,
surface runoff,ground-water flow,evaporation,and,in some cases,
evapotranspiration by plants.The ionic ratios of inland waters usually
differ appreciably from those found in the sea,although there are some
exceptions (Bayly"196]).The great chemical diversity of these waters,
the wide variation in physical conditions such as temperature,and the
relative impermanence of surface water in many cases,make it extremely
difficult to subdivide the inland salinity range in a meaningful way.
Bayly (196])has attempted a subdivision on the basis of animal life;
Stewart and Kantrud (1971)and Moyle (1945)have suggested two very
different divisions on the basis of plant I ife.We have decided to employ
a subdivision which is identical t.o that used in the Estuarine and Marine
Systems (Table I).
The term saline is used to indicate that any of a number of ions
may be dominant or co-dominant.The term brackish has been applied to
inland waters of intermediate salinity (R~mane and Schlieper 1971,
Stewart and Kantrud 1971),but is not universally accepted (see Bayly
j
1967:84).;therefore,mixosaline will be used here.In some inland wet-
lands high soii salinities control the invasion or establ;ishment of many
Table J.Salinity Modifiers Used in This Classification System.
Coastal
Modifiers l
Hyperha line
Euha.1 ine
Mixohaline (Brackish)
Polyhaline
Hesohaline
01 igohaline
Fresh
Approximate Specific
Inland Sal inity Conductance
Hodifiers 2 (%0)(llHhos at 25°C)
Hypersal i ne >40 >60,000
Eusal ine 30 -40 45.000-60.000
Mixosal ine3 0.5-30 800-45,000
Polysal ine 18 -30 30,000-45.000
Hes.osa line 5 -18 8.000-30.000
01 i gosa I i.ne 0.5-5 800-8,000
Fresh <0.5 <800
ICoastal modi·f.iers are employed in the Marine and Estuarine Systems.
21nland,modifiers are employed in the Riverine •.lacustrine and Palustrine Sy stems.
3The term Brackish should not be used for inland wetlands or deep-water habitats.
57
plants.These salinities are expressed in units of specific conductance
as well as percent salt (Ungar 1974),and they are also covered by the
salinity classes in Table 1.
2.pH MODIFIERS
Acid waters are,almost by definition,poor in calcium and often
generally low in other ions,but some very soft waters may have a
neutral pH (Hynes 1970).It is difficult to separate the effects of high
concentrations of hydrogen ions from low base content,and many studies
suggest that acidity may never be the major factor control 1 ing the
presence or absence of particular plants and animals.Nevertheless,some
researchers have demonstrated a good correlation between pH levels and
plant distribution (SjHrs 1950,Jeglum 1971).Jeglum (1971)showed that
plants can be used to predict the hydrogen ion concentration of moist
peat.
There seems to be little doubt that,where a peat layer isolates
plant roots from the underlying mineral substrate,the availability of
minerals in the root zone strongly influences the types of plants that
occupy the site.For this reason,many authors subdivide freshwater,
organic wetlands into mineral-rich and mineral-poor categories (SjHrs
1950,Heinselman 1970,Jeglum 1971,Moore and Bellamy 1974).We have
instituted pH modifiers for freshwater wetlands (Table 2)because pH has
been widely used to indicate the difference between mineral-rich and
mineral-poor sites,and because it is relatively easy to determine.The
ranges presented here are similar to those of Jeglum (1971),except that
the upper limit of the circumneutral level (Jeglum's mesotrophic)was
58
raised to bring it into line with usage of the term in the United States.
The ranges given apply to the pH of water.They were converted from
Jeglum's moist-peat equivalents by adding 0.5 pH units.
Table 2
pH Modifiers Used in This Classification System
Modifier
Acid
Circumneutral
Al ka 1 i ne
pH of Water
<5.5
5.5-7.4
>7.4
So i 1 Mod i fie rs
Soil is one of the most important physical components of wetlands.
Through its depth,mineral composition,organic matter content,moisture
regime,temperature regime and chemistry,it exercises a strong
influence over the types of plants that live on its surface and the kinds
of organisms that dwell within it.In addition,the nature of soil ina
wetland,particularly the thickness of organic soil,is of critical
importance to engineers planning construction of highways or buildings.
For these and other reasons,it is essential that soil be considered in
the classification of wetlands.
According to the U.S.Soil Conservation Service (U.S.Soil
Conservation Service,Soi 1 Survey Staff 1975:1-2),soil is limited to
terrestrial situations and shallow waters;however,"areas are not con-
sidered to have soil if the surface is permanently covered by water deep
59
enough [50]that only floating plants are present.II Since emergent
plants do not grow beyond a depth of approximately 2 m in inland waters,
thewaterward limit of soil is virtually equivalent to thewaterward
limit of wetland,according to our definition.In marine and estuarine
areas,subtidal lands do not have soil.Wetlands can then be regarded as
having soil in most cases,while deep-water habitats are never considered
to have soi 1.
The most basic distinction in soil classification in the United
States is between mineral soil and organic soil (U.S.Soil Conservation
Service,Soil Survey Staff 1975).The Soil Conservation Service
recognizes nine orders of mineral soils and one order of organic soi 15
(Histosols)in.their taxonomy.Their classification is hierarchical and
permi ts the description of soi Is at several levels of detai 1.For
example,.suborders of Histosols are recognized according to the degree of
decomposition of the organic matter.
We have chosen the modifiers mineral and organic for use in this
classification.Mineral soils and organic soils are differentiated on
the basis of very specific criteria which are enumer~ted in the Soil
Taxonomy (U.S.Soil Conservation Service,Soil Survey Staff 1975:13-14,
65).These criteria have been restated in Appendix C of this classifi-
cation for ready referen.ce.If more detail is desired,the U.S.Soil
Conservation Service classification system should be used.
Special MOdifiers
Many wetlands and deep-water habitats are man-made,and natural
ones have been modified to some degree by the activities of man or
60
beavers.Since the nature of these modifications often greatly influences
the character of such habitats,special modifying terms have been
included here to emphasize their importance.The following modifiers
should be used singly or in combination wherever they apply to wetlands
and deep-water habitats.
1.EXCAVATED
Lies within a basin or channel excavated by man.
2.IMPOUNDED
Created or modified by a barrier or dam designed to obstruct the
outflow of water.Both man-made and beaver dams are included.
3.DI KED
Created or modified by a man-made barrier or dike designed to
obstruct the inflow of water.
4.PARTIALLY DRAINED
Water level has been artificially lowered,but the area is sti 11
classified as wetland because soil moisture is sufficient to support
hydrophytes.Included under this heading also are lands where low water
levels are maintained by continuous pumping;although such areas may not
support hydrophytes presently,they could if pumping ceased.Permanently
drained areas are not considered wetland.
61
5.FARMED
The soil surface-has been mechanically or physically altered for
production of crops,but wetlandplahts will become reestabl ished if·
farming is discontinued.
6.ART!FICIAL
Refers to substrates classified as Bottom;Rocky Shore,Flat or
Beach/Bar that were emplaced by man using either natural materials such
as dredge spoil or synthetic materials such as discarded automobiles,
tires or concrete.JettIes and breakwaters are examples of Artificial
Rocky Shores.Man-made reefs are ah example of Artificial Rock Bottoms.
62
IV.REGIONALIZATION
In this classification system,a giveh taxon has no particular
regional alliance;its representatives may be found in one or many parts
of the United States.However,regional variations in climate,geology,
soils and vegetation are important in the development of different wet-
land habitats;and,often,different regions have bery different
management problems.For these reasons,we felt the need to recognize
regional differences.Regionalization is designed to facilitate:1)
planning at th~national level,where it is necessary to study management
problems and potential solutions on a regional basis;2)organization and
retrieval of data gathered in a resource inventory;and 3)interpretation
of inventory.data,including differences in indicator plants and~nimals
among the regions.
We have recommended the classification ahd map of Bailey (1976)
to fill the need for reglonallzation inland.Bailey's classification of
ecoregions is hierarchical.The upper four levels are:domain,defined
as including.subcontinental areas of related climates;division,defined
II ()as including regional climate at the level of Koppen's 1931 types;
province,defined as including broad vegetational types;arids~ction,
defined as including climax vegetation at the level of ~chler's (1964)
types.On the map,the boundaries between the different levels are
designated by various lines of various widths and the sections are
numbered with a four-digit code;digits 1 through 4 represent the first
four levels in the hierarchy.The reader is referred to Bailey's map for
a discussion and description of the units appearing on the map.
63
The Bailey system terminates at the ocean,whereas this wetla~d
classification includes .marine and estuarine habitats.Many workers have
divided marine and estuarine realms into series of biogeographic
provinces·(e.g.,U.S.Senate 1970,Ketchum 1972).These provinces differ
somewhat in detail,but the broader concepts are similar.We have
developed the following marine and estuarine provinces for North America:
ARCT i C PROV I NCE·
Arctic Province extends from the southern tip of Newfoundland
(Avalon Peninsula),~orthward around Canada to the west coasts of the
Arctic Ocean,Bering Sea,and Baffin and Laborador Basins.It is
characterized by the southern extension of floating ice,the 4°C summer
isotherm,and arctic bJbt~~
ACADIAN PROVINCE
Acadian Province extends along the northeast Atlantic coast from
the Avalon Peninsula to Cape Cod and is characterized by a well-developed
algal flora and boreal biota.The shoreline is very indented and
frequently rocky.It has a large tidal range and is ~trongly influenced
by the Laborador Current.
VIRGINIAN PROVINCE
Virginian Province extends along the middle Atlantic coast from
Cape Cod to Cape Hatteras.The province is transitional between Acadian
and Carolinian.The biota is primarily temperate with some boreal
representatives.The Laborador Current occasionally extends down to Cape
64
Hat teras and wi nter temperatures may approach 4°C •The t ida I range is
moderate.
CAROLINIAN PROVINCE
Carolinian Province is situated along the south Atlantic coast
from Cape Hatteras to Cape Kennedy •.It contains extensive marshes and
well-developed barrier islands.Waters are turbid and productive.The
biota is temperate with seasonal tropical elements.The Gulf Stream is
the primary influence and winter temperatures reach a minjmum of 10°C;
summer temperatures are tropical (in excess of 20°C).The tidal range is
small to moderat~~
WEST INDIAN PROVINCE
West Indian Province extends from Cape Kennedy to Cedar Key,
Florida and also includes the southern Guif of Mexico,the Yucatan
Peninsula,Central America and the C~rribbean Islands.The shdreland is
usually low lying 1 imestone with calcareous sands and marls except for
volcanic islands.The biota is tropical and includes reef c6rals and
.."" °mangroves.Minimum winter temperatures are about 20e and the tidal
range is small.
LOUISIANIAN PROVINCE
Louisianian Province extends along the northern coast of the Gulf
of Mexico from Cedar Key to Port Aransas,Texas.The characteristics of
the province are similar 'to those of the Carolinian,reflecting the past
submergence of the Florida Peninsula.The bi6ta is prImarily temperate
and the tidal range is small.
65
CAL1FORNIAN PROVINCE
Cal lforriian Provirice ext~nds al~ng the Pacifit coast from Mexico
northward to Cape Mendocino~The sho~eland is strongly influenced by
coastal mountains and the coasts are rocky with I imited freshwater runoff.
In the southern part volcanic sands are present;marshes and swamps are
scarce throughout the province.The climate is Mediterranean and
influenced by the California Current.The biota is temperate,and
includes well-developed offshore kelp beds.The tidal range is moderate.
COLUMBIAN PROVINCE
Columbian Province extends along the northern Pacific coast from
Cape Mendocino to Vancouver Island.Mountainous shore land with rocky
foreshores are prevalent.Estuaries are strongly influenced .by fresh-
water runoff.The biota is primarily temperate wit~some boreal
components,and there are extensive algal communities.The province is
influenced by both the Aleutian and ·CaliforniaCurrents.The tidal range
is moderate to large.
FJORD PROVINCE
Fjord Province extends along the Pacific coast from Vancouver
Island to the southern tip of the Aleutian Islands.Precipitous
mountains,deep estuaries (some with glaciers),and a heavily indented
shoreline subject to winter icing are typical of the coast.The biota is
boreal to subarctic.The province is influenced by the Aleutian and
Japanese Currents,and the tidal range is large.
66
PACIFIC INSULAR PROVINCE
Pacific Insular Province·surrounds all of the Hawaiian Islands.
The coasts h~ve pfecipit6us mount~ins and wave action is stronger than in
most of the other provinces.The biota is largely endemic and composed
of tropical and subtropical forms.The tidal range is small.
Use of Bailey's sections for the rive~ine,lacustrine and
palustrine systems and the provinces defined above for the marine and
estuarine systems provide a regional locator for any wetland in the
United States.
67
v.USE OF tHE CLAsSIFICATION SYSTEM
This system was desi~nedfor use Over an extremely wide geographic
area and for use by individuals and Qrganization~with varied interests
and objectives.The classification employs 5 system names,8 subsystem
names,12 class names,24 subclass names and an uhspecified number of
dominance types.It is,of necessity,a complex system when viewed in its
entirety,but use of the system·for a speci~lc purpose at a local site
should be simple and straightforward.The purpo.se of this section is to
illustrate how the system should be used and some of the potential pit-
falls that courd lead to its misuse.
Before attempting to apply the system,·the user should consider
certain important points:
1)Information about the area to be classified must be available
before the system can be appl ied.This information may be In
the form of historical data,aerial photographs,brief.
on-site inspection or detailed and intensive studies.The
system is designed for use at varying degrees of detail.
There will be few areas where sufficient information is
available to allow the most detailed application of the
system.If the level of detail provided by the classifica-
tion is not sufficient for the needs of the user,additional
data gathering is mandatory.
2)Below the level of class,the system is open-enoed and
incomplete.We give only examples of the vast number of
Dominance Types that occur.The user may idehtify additional
·68
dominance types and determine where these fit into the
classification hierarchy.It is also probable that as the
system is used the need for additional subclasses may become
apparent.
3)One of the main purposes of the new classification is,to
assure uniformity throughout the United States~It is
important that the user pay particular attention to the
definitions in the classification.Any attempt at modifica-
tion of these definitions will lead tb lack bf uniformity in
application.
4)One of the principal uses of the classification system will
be the inventory and mapping of wetlands and deep-water
habitats.A classification used in mapping is scale-specific,
both for the minimum size of units mapped and for the degree
of detail attainable.It is necessary for the user to
develop a specific set of mapping conventions for each
application arid to demonstrate their relationship to the
generalized classification described here.For example,
there are a number of possible mapping conventions for a
small wetland basin 50 m (164 ft)in diameter with concentric
rings of vegetation about the deepest zone.At a scale of
1:500 each zone may be classified and mapped;at 1:200,000 it
might be nec~ssary to map the entire basin as one 20ne and
ignore the peripheral bands;at 1:100,000 the entire wetland
bas in may be sma 11 er than the sma 11 est mappab 1e un i t.In the
latter case,maps will seldom be adequate for a detailed
69
inventory and must be supplemented by detailed information
gathered by sampling.In other cases ,i t may be necessary
to develop mapping conventions for taxa that cannot be easily
recognized;for instance,submergent aquatic beds in turbid
waters may have to be mapped simply as water.
HIERARCHICAL LEVELS AND MODIFIERS
We have designed the various levels of the system for specific
purposes and the relative importance of each will vary among users.The
systems and subsystems are most important in applications involving large
regions or the entire country.They serve to organize the classes into
meaningful assemblages of information for data storage and retrieval.
The classes and subclasses are the most importaht part of the
system for many users and are basic to wetland mapping.Most classes
should be easi1y recogniz~ble by users in a wide variety of disciplines.
However,the c;lass designations apply to average conditions over a period
of years,and since many wetlands are dynamic and subject to rapid
changes in appearance,it will frequently be necessary to have data that
span a per)od of years and several seasons in each of those years in
order to place a wetland in its proper class.
The dominance type is most important to users interested in
detailed regional studies.It may benecessa rytoidentifydom i nance
types in order to determine which modifying terms are appropriate because
plants and animals present in an area tend to reflect environmental
conditions over a period of time.Water regime can be determined from
long term hydrologic studies where these are available.The more common
70
procedure will be to estimate this parameter from the dominance types.
Several studies have related water regimes to the presence and distribu-
tion of plants or animals (e.g.,Chapman 1960,Stephenson and Stephenson
1972,Stew~rt and Kantrud 1972).
Similarly,we do not intend that salinity measurements be made for
all wetlands except where these data are required;in many cases,plant
species or associations can be used to indicate broad sal inity classes.
Lists of halophytes have been prepared for both coastal and inland areas
(e.g.,Duncan 1974,MacDonald and Barbour 1974,Ungar 1974),and a
number of floristic and ecological studies have described plants ~hat are
indicators of salinity (e.g.,Penfound and Hathaway 1938,Moyle 1945,
Kurz and Wagner 1957,Dillon 1966,Anderson 1968,Chabreck 1972,Stewart
and Kantrud 1972,Ungar 1974).
In some areas the dominance types to be expected under different
wate·r regimes and types of water chemistry conditions have not been
identified,and detailed regional studies will be required before the
classification can be applied in detail.In areas where detailed s6il
~aps are available,it is also possible to infer water regime and water
chemistry from soil s~ries (U.S.Soil Conservation Service,Soil Survey
Staff 1975).
Some of the modifiers are an integral part of this system and
their useisessential;othersare-Lisedonlyfor detailed applications or
for special cases.Modifiers are never used with.systems and subsystems;
however,at least one water regime modifier,one water chemistry modifier,
and one soil modifier must be used at all lower levels in the hierarchy.
Use of the modifiers listed under mixosaline and mixohaline (Table 1)is
71
optional but these fihercategories should be used where data are avail-
able.The user is cautioned not to rely on single observations of w~ter
regime or water chemistry.Such measUrements will give misleading results
in all but the most stable wetlands.The soil modifiers,mineral or
organic,must also be used.If a more detailed modifier,such as soil
order or suborder (U.S.Soil Conservation Service,Soil Survey Staff
1975)can be obtained,it should be used in place of the modifiers,
mineral and orgariic.Sp~cial modifiers are ~sed where appropriate.
RELATIONSHIP TO OTHER WETLAND CLASSIFICATIONS
There are numerous wetland classifications in use in the United
States.Here we relate this system to three published classifications
that have gained widespread acceptance.It is not possible to equate
these syStems directly for several reasons:1)The criteria selected for
establ ishing categories differ,2)some of the classifications are not
applied consistently in different parts of the country,and 3)the
elements classified are not the same in various classifications.
The most widely used classification system in the United States
is that of Martin et al.(1953)which was republ ished in IICircular 39 11
(Shaw and Fredine 1956).The wetland types are based on criteria such as
water depth and permanence,water chemistry,life form of vegetation and
dominant plant species.Table 3 compares some of the major components of
our system with the type descriptions listed in Circular 39.
In response to the need for more detailed wetland classification
in the glaciated northeast,Golet and Larson (1974)refined the fresh
water wetland types of CirCUlar 39 by writing more detailed descriptions
Table 3.Comparison of Wetland Types Described in Circular 39 with Some of the Kajor Components of this
Classification Sy:stem.
Classification of Wetlands and Deep-Water Habitats
Circular 39
Type
Type
Type 2
Type 3
Type 4
Type 5
Type 6
Type 7
References for Examples
Typical Vegetation
Wet Meadow (Stewart &Kantrud
1972,Dix &S~ins 1967
Bottomland Hardwoods (Braun 1950)
Sha 11ow-FreshNater Swamps
(Pen found 1952)
Fen (Heinselman 1963)
Fen,Northelin Sedge Meadow
(Curt is 1959)
Shallow Marsh (Golet &Larson
197~,Stewart &Kantrud 1972)
Deep Marsh (Golet &Larson
1974,Stewart &Kantrud 1972)
Open.Water (Golet &Larson 1974)
Submerged Aquatic (Curtis 1959)
Shrub Swamp (Golet &Larson 1974)
Shrub-Carr,Alder Thicket
(Curtis 1959)
Wooded S10IiIlIlp (Golet &Larson 1974)
SwaIlIPS (Penfound 1952.
Heinsehaan 1963)
Classes
Emergent Wetland
Forested Wet land
Emergent Wetland
Emergent Wetland
Emergent Wet I and
Aquat ic Bed
AquiJtic Bed
Unconso II dated Bot tom
Scrub/Shrub Wet I and
Forested Wetland
Water Regimes
Te.porarily Flooded
Intermittently Flooded,
Saturated
S_ipermanently Flooded
Seasonally Flooded
Perwanent I y Flooded
Intermittently Flooded
Senlpermanently Flooded
Pe~nently Flooded
Intermittently Exposed
All Nontldal Regimes
Except Permanently
Flooded
All Nontldal Regimes
Except Permanently
Flooded
Wate'r Chemistry
Fresh
Mlxosallne
Fresh
Mi,xosal ine
Fresh,
Mixosal ine
Fresh
Mlxosa 1'1 ne
Fresh
Mlxosaline
Fresh
Fresh
-.....J
N
Table 3.(continu~d)
Circular 39
Type
Type 8'
Type 9
Type 10
Type II
Type 12
Type 13
Type I"
Type 15
References for Examples
Typical Vegetation'
Bog (Heinselman 1963,
Dansereau &Segadasvianna 1952)
Pocosin (Kologiski 1977)
Peaty Fresh-Water Swamps
(Pen found 1952)
Intermittent Alkali Zone
(Stewart &Kantrud,1972)
Inland Salt Marshes
(Ungar 197%)
InIand Sa line.Lake COIIIlIUn i ty
(Ungar 197%)
Marsh (Anderson 1968)
Estuarine Bay Marshes,Estuarine
River Marshes (Stewart 1962)
Fresh &Inter1llediate Marshes
(Chabreck 1972)
Harsh (Anderson 1968)
Estuarine Bay Marshes,Estuarine
River Marshes (Stewart 1962)
Fresh &Iiltermediate Marshes
(Chabreck 1972)
Estuarine B3ys
•(Stewart &Kantrud,1972)
Panne,Harsh Slough
(Redfield 1972)
Marsh Pans (Pestrong 1965)
Classes
Scrub/Shrub Wetland
Forested Wetland
Moss/Lichen Wet land
Flat
Emergent Wetland
Aquatic Bed
Unconsolidated Bottom
Emergent Wetland
Emergent Wetland
Aquatic:Bed
Unconsolidated Bottom
Flat
Water Reg imes
Saturated
Seasonal Iy Flooded
Intermittently Flooded
Temporarily Flooded
Seasona II y FJ,ooded
Semipermanently Flooded
Permanently Flooded
Intermittently Flooded
Regularly Flooded
I rregularly Flooded
Semipermanently Flooded-
Tidal
Regularly Flooded
Semlpermanently Flooded-
Tidal
Subtidal
Permanently Flooded-Tidal
ReguJarly Flooded
Irregularly Flooded
Water."Chemi st ry
Fresh
(acid only)
Eusal ine
Hypersal ine
Eusal ine
Eusal i ne
Mi xoha I ine
Fresh '.
Mixoha.1 ine
Fresh
Hixohallne
Fresh
Hyperha line
Euhal ine
Table 3.(continued)
Ci rcular 39
Type
Type 16
Type 17
Type 18
Type 19
Type 20
References for Examples
Typical Vegetation
Salt Karsh (Chapman 1960,
Redfield 1972)
Sal'tKarsh (Chapman 1960)
Saline,Brackish &Intermediate
(Ele~terfus 1972)
Salt·Karsh (Chapman 1960)
Kelp Beds,Temperate Grass Flats
(Phillips 1974)
Trop'i ca I Kar i ne Meadows
(Odin 1974)
Eelgras·sBeds
(Akins &Jefferson 1973.
E.leuterius 1973
KanQrove Swamps (Wa Ish 1974)
Kangrove Swamp Systems
(Kuenzler 1974)
Kangroves(Chapman 1976)
Classes Water Regimes Water Chemistry
.Emergent Wetland I rregu I ar Iy Flooded Euhal ine
"ixohal i ne
Emergent Wet land I rregu lar Iy Flooded tuhal ioe
"ixohal ine
Emergent Wetland.Regularly Flooded Euhaline
"ixohaline
Unconsolidate~Bottom.Subti dal Euha.line
Aquatic Bed I rregul'arly ExpOsed.Hixohal ine
Flat Regu lady Floc;>ded
Irregularly Flooded
""""-'='"
Scrub/Shrub Wetland Irregularly Exposed Hyperha line
Forested.Wetland Regularly Flooded Euhal ine
Irregularly FloOded "i xohal i ne·
..
75
and subdividing classes on the basis of finer differences in plant life
forms.Golet ahd Larson's classes are roughly equivalent to Types 1-8 of
Circular 39,except that they restricted Type 1 to river floodplains.
The Golet and Larson system does not recognize the coastal (tidal)fresh
wetlands of Circular 39 (Types 12-14)as a separate cate~ory,but
classifies these areas in the same manner as nontidal wetlands.In
addition to devising 24 subclasses,they also created five size categories;
six site types,giving a wetland's hydrologic and topographic location;
eight cover types (modified from Stewart and Kantrud 1971)expressing the
distribution and relative proportions of cover and water;three vegetative
interspersion types;and six surrounding habitat types.Since this
system is based on the classes of Martin et aL (1953),Table 3 may also
be used to compare the Golet and Larson system to the one described here.
Although our system does not include size categories and size types,this
information will be available fro~the results of the new inv~ntory of
wetlands and deep-water habitats of the United States.
Stewart and Kantrud (1971)devised a new classification system ·to
better serve the needs of researchers and wetland managers in the
glaciated prairies.Their system recognizes seven classes of wetlands
which are distinguished by the vegetational zone occupying the central or
deepest part and covering five percent or more of the wetland basin.The
classes thus reflect the wetland's water regime;for example,temporary
ponds (Class I I)are those where the wet-meadow zone occupies the deepest
part of the wetland.Six possible subclasses were created,based on
differences in plant species composition that are correlated with
variations in average salinity of surface water.The third component of
76
classification in their system is the cover type which represents
di ffercnc~5 in the spatial relation of emergent cover to open water or
exposed bottom soil.The zones of Stewart and Kantrud's system are
readily related to our water regime modifiers (Table 4),and the 5ub-
classes are roughly equivalent to our·water chemistry modifiers
(Table 5).
A classification system is most easily learned through use.To
illustrate the application of this system,we have classified a repre-
sentative group of wetlands and deep-water habitats of the United
States.1
l The final pubilcation wl11 include a classification of a repre-·
sentat ive group ofwe.t 1ands and.deep-water habi tats of the Un i ted States.
Photographsofth~sehabitats wIll also be provided.
77
Table 4..Comparisoll of the Zones of Stewart and Kan rud1s (1971)
Classification with the Water Regime ·Modif ersused in
this Classification System
Zone
(Stewart and Kant rud)
Wetland-I ow-pra i de zone
Wet Meadow Zone
Shallow Marsh Zone
Deep Marsh Zone
Permanent Deep Water Zone
Intermlttent-alkaliZone
Fell (Alkaline Bog)Zone
Water Regime
Modi fier
non wetland by one definition
Temporarily Flooded
Seasonally Flooded
Semipermanently Flooded
Intermittently Exposed
Permanently Flooded
Intermittently Flooded (with
saline or hypersaline water)
Saturated
78
Tab le 5
Cbmparisonofthe Water Chemistry Subclasses of
Stewart and Kantrud (1972)with the Water
Chemistry Modifiers used in thts
Classification System
This c111ssif ieation
Approximate
sp'ecific
conductances
(Mh
Stdwart and Kantrud
(·972),..'I ~J os)
100.000
Saline
60.000
H }'persal ine
45.000
Eusaline
Pol}saline
Subsaline 30.000
15.000 Mesosalint"
Brack;sh 8.000 Mixosaline
5.000
Modeiately Brackish Oligosaline
2.000
Slightly Brackish
800
500
Fresh
Fresh
40
n
79
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soil surveys.Preliminary Abridged Text.U.S.Soil Conserv.
Serv.,Washington,D.C.330 pp.
89
Walsh,G.E.1974.Mangroves:a review.Pages 51-174..!1!R.J.
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90
APPENDIX A
SCIENTIFIC AND COMMON.~AMES OF PLANTS
SCIENTIFIC NAME
Acer rubrum L.
Alnus spp.
A.rugosa (DuRoi)Spreng.
Andromeda glaucophylla Link
Aulacomnium palustre (Hedw.)Schwaegr.
Avicennia germinans (L.)L.
(~.nitida Jacq.)
Azolla spp.
Baccharis halmifolia L.
Beckmannia syzigachne (Steud.)Fern.
Betula pumila L.
Brasenia schreberi
Campyliu~stellatum (Hedw.)C.Jens.
Carex spp.
Cephalanthus occidental is L.
Ceratophyllum spp.
C.demersum L.
Chamaecyparis thyoides (L.)BSP.
Chamaedaphne caljculata (L.)Moench
Chara spp.
Chenopodium rubrum L.
Chi losyphus fragi 1 is (Roth)Schiffn.
Cladium jamaicens Crantz
Cladonia spp.
Conocarpus erecta L.
Decodon verticillatus (L.)E11-
Desmatodon heimii (Hedw.)Mi tt.
Drepanocladus (C.Mue 11 . )Roth
COMMON NAME
Red Maple
Alders
Speckled Alder
Bog-rosemary
Black Mangrove
Mosquito-fern
Sea-myrtle
Ameri~an Sloughgrass
Swamp Birch
Water;-sh ie ld
Sedges
Buttonbush
Hornwort
Coonta i1
White Cedar
Leather-leaf
Stonewort
Coast-bl ight
Sawgrass
Reindeer Moss
Buttonwood
Water-wi 11 ow
Aquatic Moss
91
APPENDIX A (continued)
SCI ENT I Fie NAME
Echinochloa crusgalli (L.)Beauv ..
Eichhornia crassipes (Mart.)Solms.
Eleocharis aclcularis (L.)R.&S.
Equisetum fluviatile L.
Fissidens spp.
f.jul ianus (Mont.)Schimp.
Fontinalis
Fraxinus nigra Marsh
f.pennsylvanica Marsh
Glyceri a spp.
Go rdon ia I as i an th us (L.)EI lis
Halodule beaudettei (de~Hartog)den Hartog
Halophi la spp.
Iva frutescens L.
Juncus roemerianus Scheele
Kalmia polifolia Wang.
Laguncularia racemosa Gaertn.f.
Ledum groenlandicum Oeder
Larix laricina (Du Roi)K.Koch
Lemna spp.
Leucothoe axillaris (Lam.)D.Don
Lyonia lucida (Lam.)K.Koch
Lythrum salicaria L.
Macrocystis spp.
Magnolia virginiana L.
Marsupella spp.
Myriophyllum spp.
Najas spp.
Nitella spp.
Nuphar spp.
N.advena (Ait.)f.
N.variegatum
COMMON NAME
Wild Millet
Water-hyacinth
Sp i ke-rush
Water Horseta i I
Water Moss
Water Moss
Water Moss
Black Ash
Red Ash
Manna-grasses
Loblolly Bay
Shoal grass
Sea-grass
Marsh Elder
Needle Rush
Pale or Bog-laurel
White Mangrove
Labrador-tea
American Larch or Tamarock
Duckweeds
Leucothoe
Tetter-bush
Purple Loosestrife
Brown Algae
Sweet Bay
Li verwort
Water-mi I foi I
Naiads
Nitella
Spatter-dock
Ye I low Pond-lily
Variegated Pond-lily
92
APPENDIX A (continued)
SCIENTIFIC NAME
Nymphaea spp.
Nymphoides aquatica (Walt.)o.Kuntze
Nyssa aquatica L.
~.sylvatica Marsh
Peltandra virginica (L.)Kunth
Pelvetia spp.
Persea borbonia (L.)Spreng.
Phragmites communis Trin.
Phyllospadix scouleri Hook.
~.torreyi Wats.
Picea mariana (Mill.)BSP
Pinus serotina Michx.f.
Pistia stratiotes L.
Podostemum ceratophyllum Michx.
Polygonum spp.
~.amph i bi urn L.
Pontederia cordata L.
Potamogeton spp.
~.epihydrus Raf.
P.natans L.
Quercus bicolor Willd.
Q.lyrata Wal t.
Q.michauxii Nutt.
Rhizophora mangle L.
Rumex mexicanus Meisn.
Ruppia maritima L.
Sagi ttari a spp.
Sa~icornia pacifica StandI.
Sal ix spp.
Salvinia spp.
S.rotund i fo I ia Will d.
COMMON NAME
Water-lily
Floating-heart
Water Gum or Tupelo Gum
Black Gum
Arrow-arum
Slim Brown Rockweed
Red Bay
Reed
Surf Grass
Surf Grass
Black or Bog-spruce
Pond or Marsh-pine
Water-lettuce
Threadfoot
Smartweeds
Water-smartweed
Pickerelweed
Pondweeds
Ribbonleaf Pondweed
Floating-leaf Pondweed
Swamp White-oak
Ove r-cup Oak
Basket Oak
Red Mangrove
Widgeon-grass
Arrowhead
Glasswort or Common
Pickleweed
Wi llow
Water Fern
Water Fern
....
93
APPENDIX A (continued)
SCIENTIFIC NAME
Scirpus acutus Muhl.
Scolochloa festucacea (Willd.)Link
Spartina alterniflora Loisel.
S.cynosuroides (L.)Roth
S.foliosa Trln.
S.patens (Ait.)Muhl.
Sphagnum spp.
Spirodela polyrhiza (L.)Schleid.
Suaeda californica Wats.
Syringodium filiformis Kuetz
Taxodium ascendens Brogn.
T.distichium (L.)Richard
Thalassia testudinum Koenig &Sims
Thuja occidentalis L.
Trapa spp.
Triglochin maritima L.
Typha angustifolia L.
T.I at i fo I i a L.
Ul mus arne r i cana
Ulva spp;
Utricularia vulgaris L.
Vallisneria americana Michx.
Xanthium italicum Moretti
Zenobia pulverulenta (Bartr.)Pollard
Zizania aquatica L.
Zostera marina L.
Zizaniopsis miliacea (Michx.)Ddl &Asch.
COMMON NAME
Hard-stem Bulrush
White-top
Salt-water Cordgrass
Big Cordgrass
Cordgrass
Saltmeadow Cordgrass
Peat Moss
Big Duckweed
Sea-b lite
Manatee-grass
Pond Cypress
Bald Cypress
Turt Ie-grass
White Cedar or Arbor Vitae
Water-nut
Arrow-grass
Narrow-leaved Cat-tail
Common Cat-tail
American or White Elm
Sea lettuce
Bladderwort
Wi Id Ce lery
Cocklebur
Zenobia
Wi Id Rice
Grass-wrack or Eelgrass
Water Mi llet
SCIENTIFIC NAME
Acmaea
Acropora
Amphitrite
Amphiura
Anodonta
Anopheles
Arenicola
Baetis
Balanus
Caenis
Callianassa
Cerianthiopsis
Cerithium
Chaeopterus
Chaoborus
Chiredotea
Chironomus
Chthamalus
Cnemidocarpa
Crassostrea
Dendraster
Diamesa
Donax
Echinocardium
Emeri ta
Ephemera
Eukiefferiella
Gammarus
94
APPENDIXB
SCiENTIFIC AND COMMON NAMES OF ANIMALS
COMMON NAME'"
Fa 1se Li mpet
Coral
Terebellid Polychaete
Starfi sh
Freshwater Clam
Mosqui to
Lugworm
Mayfly
Barnacle
Mayfly
Burrowing Shrimp
Sea Anemone
Sna i1
Polychaete
Diptera
Isopod
Midge
Barnacle
Ascidian
Oyster
Sea Star
Dipte ra
Wedge She 11
Sea Urchin
Sand Bug
May Fly
Diptera
Amphipod
..
95
APPENDIX B (continued)
SCIENTIFIC NAME
Haustori us
Herpobdella
Hi ppospong ia .
Hydropsyche
Leptosynapta
Limnea
Li mnodr i 1us
Littorina
Lumbriculus
Lygia
Lymnaea
Macoma
Me 11 ita
Mitel1a
Modio Ius
Montipora
Muri cea
Mya
Mytilus
Nassar ius
Nemoura
Nereis
Ned ta
Ocypode
01 iva
Orchest ia
Paract is
Parastenocaris
Patella
Pecten
Penaeus
""
",',,:
COMMON NAME'"
.{\mp~ipo?
Leech
Encrusting Sponges
Caddis Fly
Holothurian
01 i gochaete
Periwinkle
01 i gochaete
Isopod
Freshwater Snail
Te 11 in She 11
Amphipod,.
Barnacle
Mussel
Coral
Al cyonari an
Long Neck Clam
Mussel
Mud Sna i 1
Plecoptera
Polychaete
Periwinkle
Gho?t -Crab
Amph i pop"
Oyster
Sea Anemone
Cope pod
Limpet
Sea llop
Shrimp
96
APP~NDI X B (cont i m.ted)
SCIENTIFIC NAME
Phyllognathus
Physa
Pi sas ter,
Pisidium
Pori tes
Pri st ina
Renilla
Sabellaria
Siphonaria
Spatangus
Spongi I la
Strongy 19centrotus
Tel Ii na
Tet racJ ita
Thais
Thyone
Thoracophel ia
Tivela
Tubifex
Uca
Urechis
COMMON NAME'"
Capepod
Freshwater Snail
Seastar
Mo 11 usk
Coral
01 i gochaete
Sea Pansy
Reefworm
False limpet
Heart Urchin
Sponge
Sea Urchin
Tellin Shell
Barnacle
Rock She 11
Sea Cucumber
Bamboo Worm
Pismo"Clam
Sewage Worm
Fiddler Crab
Echivrid
*Organisms are identified only to genus,so the common name refers to
general groupings.
..
97
APPENDIX C
CRITERIA FOR DISTINGUISHING ORGANIC SOILS FROM MINERAL SOILS
The criteria for distinguishing organic soils from mineral soils
in the United States (U.S.Soil Conservation Service,Soil Survey
Staff 1975:13-14,65)are quoted here so that those without ready
access to a copy of the Soil Taxonomy may employ this information in the
classification of wetlands:
For purposes of taxonomy,it is necessary,first,to define the
limits that distinguish mineral soil material from organic soil
material and,second,to define the minimum part of a soil that
should be mineral if the soil is to be classified as a mineral soil.
Nearly all soils·contain more than traces of both mineral and
organic components in some horizons,but most soils are dominantly
one or the other.The horizons that are less than about 20 to 35
percent organic matter by weight have properties that are more nearly
those of mineral than of organic soils.Even with this separation,
the volume of organic matter at the upper limit exceeds that of the
mineral material in the fine-earth fraction.
MINERAL SOIL MATERIAL
Mineral soil material either
1.Is never saturated wi th water for more than a few days and has
<20 percent organic carbon by weight;
or
2.Is saturated with water for long periods or has been arti-
ficially drained,and has
a.Less than 18 percent organic carbon by weight if 60 percent
or more of the mineral fraction is clay;
b.Less than 12 percent organic carbon by weight if the
mineral fraction has no clay;or
c.A proportional content of organic carbon between 12 and 18
percent if the clay content of the mineral fraction is between
zero and 60 percent.
Soil material that has more organic carbon than the amounts just
given is considered to be organic material.A full definition of :
organic soil materials is given in Chapter 4.
98
DISTINCTION BETWEEN MINERAL SOILS
AND ORGANIC SOILS
Most soils,are dominantlymineri:ll material,but many mineral
soils have horizons of organic material.For simplicity in writing
definitions of taxa,a distinction between what is meant by a mineral
soil and an organic soil is useful.In a mineral soil,the depth of
each horizon is measured from the top of the first horizon of mineral
material.In an organic soil,the depth of each horizon is measured
from the base of the aerial parts of the growing plants or,if there
is no continuous plant cover,from the surface of the layer of
organic materials.To apply the definitions of many taxa,therefore,
one must first decide whether the soil is mineral or organic.
If a soil has both organic and mineral horizons,the relative
thickness of the organic and the mineral soil materials must be con-
sidered.At some point one must decide that the mineral horizons
are more important.This point is arbitrary and depends in part on
the nature of the materials.A thick layer of sphagnum has a very
low bulk density and contains less organic matter than a thinner
layer of well-decomposed muck.It is much easier to measure thick-
ness of layers in the field than it is to determine tons of organic
matter per hectare.The definition of a mineral soil,therefore,is
based on thickness of the horizons or layers,but the limits of
thickness must vary with the kinds of material~.The definition
that follows is intended to classify as mineral soils those that have
no more organic material than the amount permitted in the histic
epipedon,which is defined later in this chapter.
To determine whether a soil is organic or mineral,the thickness
of horizons is measured from the surface of the soil whether that is
the surface of a mineral or an organic horizon.Thus,any 0 horizon
at the surface is considered an organic horizon,if 'it meets the
requirements of organic soil material as defined later,and its
thickness is added to that of any other organic horizons to determine
the total thickness of organic soil materials.
DEFINITION OF MINERAL SOILS
Mineral soils,in this taxonomy,are soils that meet one of the
following requirements:
1.Mineral soil material <2mm in diameter (the fine-earth fraction)
makes up more than-half the thickness of the upper 80 cm
(31 in.)CsicJ;
2.The depth to bedrock is <40 cm and the layer or layers of
mineral soil directly above the rock either are 10 cm or more thick
or have half or more of the thickness of the overlying organic soil
mated a 1;or
3.The depth to bedrock is >40 cm,the mineral soil material
immediately above the bedrock is 10 cm or more thick,and either
a.Organic soil material is <40 cm thick and is decomposed
(consisting of hemic or sapric materials as defined later)or
has a bulk density of 0.1 or more;or
l'
•
99
b.Organic soil material is <60 cm thick and either is
undecomposed sphagnum or moss fibers or has a bulk density that
is <0.1.·
ORGANIC SOIL MATERIALS
Organic soil materials and organic soils
1.Are saturated with water for long periods or are artificially
drained and t excluding live roots t (a)have 18 percent or more
organic carbon if the mineral fraction is 60 percent or more clay,
(b)have 12 percent or more organic carbon if the mineral fraction
has .no clay,or (c)have a proportional content of organic carbon
between 12 and 18 percent if the clay content of the mineral
fraction is between zero and 60 percent;or
2.Are never saturated with water for more than a few days and
have 20 percent or more organic carbon.
Item 1 in this definition covers materials that have been called
peats and mucks.Item 2 is intended to include what has been called
litter or a horizons.Not all organic soil materials accumulate in
or under water.Leaf litter may rest on a lithic contact and
support a forest.The only soil in this situation is organic in the
sense that the mineral fraction is appreciably less than half the
weight and is only a small percentage of the volume of the soil.
DEFINITION OF ORGANIC SOILS
Organic soils (Histosols)are soils that
1.Have organic soil materials that extend from the surface to one
of the following:
a.A depth within 10 cm or less of a lithic or paralithic
contact,provided the thickness of the organic soil materials
is more than twice that of the mineral soil above the contact;
or
b.Any depth if the organic soil material rests on fragmental
material (gravel t stones,cobbles)and the interstices are
filled with organic materials t or rests on a lithic or paralithic
contact;or
2.Have organic materials that have an upper boundary within 40 cm
of the surface and
a.Have one of the following thicknesses:
(1)60 cm or·more if three-fourths or more of the volume is
moss fibers or the moist bulk density is <0.1 g per cubic
centimeter (6.25 Ibs per cubic foot);
(2)40 cm or more if
(a)The organic soil material is saturated with water
for long periods (>6 months)or is artificially drained;
and
(b)The organic material consists of sapric or hemic
materials or consists of fibric materials that are less
than three-fourths moss fibers by volume and have a
moist bulk density of 0.1 or more;and
100
b.Have organic soil materials that
(1)Do not have a mi nera 11 ayer as much as 40 cm th i ck
either at the surface or whose upper boundary is within a
depth of 40 cm from the surface;and
(2)Do not have mineral layers,taken cumulatively,as thick
as 40 cm wi th in the upper 80 cm..
It is a general rule that a soil is classed as an organic soil
(Histosol)either if more than half of the upper 80 cm
(32 in.)[sic]of soil is organic or if organic soil material of
any thickness rests on rock or on fragmental material having
interstices filled with organic materials.
Soils that do not satisfy the criteria for classification as
organic soils are mineral soils.
..