&EPA
United States
Environmental Protection
Agency
Office of Research and
Development
Washington, DC 20460
EPA/600/R-06/132
October 2006
www.epa.gov
A Framework
for the Assessment of
the Wildlife Habitat Value
of New England Salt Marshes
Wildlife Habitat Value
of New England Salt Marshes
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EPA/600/R-06/132
October 2006
A Framework for the Assessment of the
Wildlife Habitat Value of New England
Salt Marshes
Richard A. McKinney
Cathleen Wigand
U.S. Environmental Protection Agency
Office of Research and Development
National Health and Environmental Effects Research Laboratory
Atlantic Ecology Division
27 Tarzwell Drive
Narragansett, RI 02882 USA
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Wildlife Habitat Value of New England Salt Marshes
Notice
The Office of Research and Development (ORD) has produced this document to provide a framework
for assessing the wildlife habitat value of New England salt marshes. Assessment protocols can be
used to provide information on the habitat value of coastal wetlands to aid in protection, restoration,
and mitigation of salt marsh habitats. This document should be cited as:
U.S. EPA. 2006. A Framework for the Assessment of the Wildlife Habitat Value
of New England Salt Marshes. EPA/600/R-06/132. Office of Research and Development.
Washington, DC 20460.
The research described in this report has been wholly funded by the US Environmental Protection
Agency and has been subjected to external peer review, however, it does not necessarily reflect the
views of the Agency.
Mention of trade names or commercial products does not constitute endorsement or
recommendation.
Abstract
Resource managers are frequently asked to make decisions that affect the protection and
restoration of wetland habitats. The desire is often to base at least some part of the decision on an
assessment of one or more wetland functions, such as wildlife habitat value. While protocols
currently exist to evaluate wildlife habitat value in freshwater wetlands, there is a lack of stand-alone
methods to assess this function for coastal salt marshes, a class of wetlands that are increasingly under
development pressure from urbanization. In this report, we provide a framework for assessing the
wildlife habitat value of New England salt marshes by identifying the habitat characteristics that
influence the presence and abundance of wildlife species. We identify these characteristics from
available information on the habitat requirements of 79 bird, 20 mammal, and 6 reptile and
amphibian species that use New England salt marsh habitats. The characteristics are incorporated
into wetland and landscape components (e.g., salt marsh size, salt marsh landscape setting) that we
feel are important for determining habitat suitability for wildlife species. For each component, we
identify several categories that provide a means for ranking habitat value. The wetland and
landscape components, along with their associated categories, can be used as the basis of an
assessment protocol to estimate salt marsh wildlife habitat value.
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Wildlife Habitat Value of New England Salt Marshes
Foreword
Since the late 1970's, most wetlands have been considered "waters of the U.S." and regulated under
the Clean Water Act (CWA). Under the CWA the U.S. Environmental Protection Agency, States,
and Tribes develop programs for protecting the chemical, physical, and biological integrity of the
nation's waters, including wetlands. A necessary step towards protecting and restoring the biological
integrity of wetlands is to ascertain the relative habitat value of wetlands in a landscape. This
manuscript presents a framework for assessing the wildlife habitat value of coastal wetlands by
identifying the habitat characteristics that influence the presence and abundance of wildlife species.
The framework is based on relevant life history traits and habitat requirements of (terrestrial) wildlife
species that use salt marshes. We identify eight wetland components that we feel would be
important to assess wildlife habitat value, such as the presence of habitat types (e.g. marsh -upland
border, pools, tidal flats), marsh morphology, size, and extent of anthropogenic modification. We
then propose categories within each component that relate to the habitat value of the marsh.
This manuscript is the first phase of developing the assessment protocol, consisting solely of the
scientific basis for developing the assessment indicators. In a subsequent manuscript we will present
specific ranking and scoring protocols for New England salt marshes. Once established, an
assessment protocol can be used to provide information on the habitat value of coastal wetlands to aid
in protection, restoration, and mitigation of salt marsh habitats.
This is the Office of Research and Development, National Health and Environmental Effects Research
Laboratory, Atlantic Ecology Division contribution number AED-06-054.
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Wildlife Habitat Value of New England Salt Marshes
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IV
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Wildlife Habitat Value of New England Salt Marshes
Contents
Notice ii
Abstract ii
Foreword iii
Introduction 1
- Salt Marsh Wildlife Habitat Requirements 2
- Wetland and Landscape Assessment Components 6
I. Salt Marsh Size Class 8
II. Salt Marsh Morphology 11
III. Salt Marsh Habitat Type 16
IV. Extent of Anthropogenic Modification 22
V. Salt Marsh Vegetation 27
VI. Salt Marsh Vegetative Heterogeneity 29
VII. Surrounding Land Cover and Land Use 30
VIII. Connectivity and Associated Habitat 32
Conclusion 33
Literature Cited 35
Appendix 1 47
Appendix 2 51
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Wildlife Habitat Value of New England Salt Marshes
Tables
Table 1. Birds known to inhabitant New England salt marshes or use salt marshes
as foraging or shelter habitat 3
Table 2. Mammals, amphibians, and reptiles known to inhabitant New England salt marshes
or use salt marshes as a foraging habitat 5
Table 3. Habitat types, edge habitats, and adjoining habitats of value to sale marsh wildlife 6
Table 4. Most commonly reported habitat types, edge habitats, and associated habitats
used by salt marsh breeding and foraging birds 7
Table 5a. Most commonly reported habitat types, edge habitats, and associated habitats
used by grazing, predator, and breeding salt marsh mammals, as well as mammals
that use salt marshes 7
Table 5b. Most commonly reported salt marsh habitat types, edge habitats, and associated
habitats used by amphibians and reptiles 7
Table 6. Wetland and landscape assessment components of New England
Salt marshes and their associated categories 8
Table 7. Body mass, optimal foraging water depth, and tarsus length of wading birds
In the family Ardeidae that utilize New England salt marshes 19
Figures
Figure la. Salt meadow marsh - Back-barrier or basin marsh with extensive creek systems interspersed
with salt meadow marsh interior 13
Figure Ib. Meadow / fringe marsh - Typically consists of areas of salt meadow marsh
interspersed with narrow or wide fringe marsh 13
Figure Ic. Wide fringe marsh - Typically dominated by low marsh but can contain
Some patches of high marsh vegetation 14
Figure Id. Narrow fringe marsh - Consists of a narrow belt of vegetation dominated
primarily by the low marsh Spartina alterniflora with few creeks 14
Figure le. Marine fringe marsh - A narrow fringe marsh that is bordered on the seaward
edge by unprotected open water 15
VI
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Wildlife Habitat Value of New England Salt Marshes
Figure 2. Salt marsh habitat types and the occurrence within a typical
New England salt marsh 17
Figure 3a. Extent of ditching in New England salt marshes - Little or no ditching 24
Figure 3b. Extent of ditching in New England salt marshes - Moderate ditching 24
Figure 3c. Extent of ditching in New England salt marshes - Severe ditching 25
Figure 4a. Extent of tidal restrictions in New England salt marshes
- No to low tidal restriction 25
Figure 4b. Extent of tidal restrictions in New England salt marshes
- Moderate tidal restriction 26
Figure 4c. Extent of tidal restrictions in New England salt marshes
- Severe tidal restriction 26
Figure 5. Occurrence of varying degrees of vegetative heterogeneity in
New England salt marshes 30
VII
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Wildlife Habitat Value of New England Salt Marshes
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VIM
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Wildlife Habitat Value of New England Salt Marshes
Introduction
Environmental stewards and managers
acknowledge the importance of assessing the
value of wetlands for the purposes of protection,
restoration, and mitigation. These assessments
may be particularly important for coastal salt
marshes, a class of wetland that by the nature of
their location are increasingly under
development pressure from urbanization.
Wetland assessment protocols typically either
add an evaluation of wetland function to existing
habitat classification systems or include wetland
functional assessments as one component of an
overall classification (Bartoldus 1999). Examples
of the latter include the hydro geomorphic
approach (Brinson 1993), those based on national
wetland classification protocols (e.g., Tiner 2003),
and stand-alone wetland assessment techniques
and protocols (e.g., Adamus et al. 1987). These
assessments address many wetland functions
including water quality improvement, flood
control, ground-water recharge, and wildlife
habitat value.
Of the wetland functions addressed by
assessment protocols, wildlife habitat value has
garnered particular attention and led to the
development of several regional classification and
assessment protocols for freshwater and inland
wetlands. These assessments often rely on
general vegetative characteristics to estimate
habitat value without consideration of the
specific habitat requirements of wildlife species
known to inhabit the wetlands (e.g., Schroeder
1996). Notable exceptions are the classifications
developed by J. S. Larson and coworkers to assess
the wildlife habitat value of freshwater wetlands
in the northeast U.S. (e.g., Golet and Larson
1976, Whitlock et al. 1994). These classifications
are based on dominant vegetation, but they also
incorporate wildlife habitat requirements.
A number of protocols have been developed
to assess the wildlife habitat value of freshwater
marshes (Bartoldus 1999). However, to our
knowledge there are no species-specific, stand-
alone assessment protocols to assess the wildlife
habitat value of coastal salt marshes. The
objective of this report is to present a framework
Snowy egret Egretta thula
(Photo by Ryan Hagerty, US FWS)
for the development of assessment protocols for
wildlife habitat value in coastal salt marshes in
New England that are based on the presence of
marsh habitat types, marsh morphology, and
landscape setting and incorporate the specific
habitat requirements of resident wildlife species.
The report identifies terrestrial wildlife species
(birds, mammals, reptiles, and amphibians)
known to use salt marshes during some part of
their life histories and compiles habitat use data
from published life history accounts, unpublished
reports, and anecdotal information from
wetlands ecologists. Habitat requirements of
species are organized into the a series of wetland
components that provide a framework for
assessing wildlife habitat value for New England
1
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Wildlife Habitat Value of New England Salt Marshes
salt marshes. For each component, we propose
several categories that can be used to classify salt
marshes for wildlife habitat value. The different
categories within each component range from
Great egret aldea alba
(Photo by Lee Karney, US FWS).
those that imply that wildlife species would
obtain the full benefit or habitat value of the
component to those implying that the species
would obtain less than full value. However,
when utilized in an actual assessment, the
weighting and ranking of components will
depend upon the target wildlife species under
consideration and the overall intent of the
assessment.
In this report we focus on New England salt
marshes, defined as those occurring from Maine
to New Jersey (Chapman 1940). New England
salt marshes are typically small and receive low
suspended sediment loads from relatively small
drainage basins, resulting in predominately
organic peat substrates (Roman et al. 2000). Salt
marsh morphology in this region reflects the
relatively steep slope of New England estuarine
coastlines, as well as the influence of
development and modification by humans (Kelly
1987, Kelly et al. 1988). Traditionally, studies on
New England salt marsh habitat value have
emphasized marine species that depend on salt
marsh habitats during a portion of their life
cycle. For example, mummichog Fundulus spp.
and several shrimp species (e.g., Paleomontes
spp.) are resident in salt marshes (Cross and
Stiven 1999, Halpin 2000), and others use salt
marsh habitats for egg laying (Harrington and
Harrington 1961, Daiber 1962) and foraging
(Vince et al. 1976, Daiber 1982), However, in this
report we focus on terrestrial wildlife and present
a framework for the assessment of salt marsh
habitat value solely for these species.
SALT MARSH WILDLIFE HABITAT REQUIREMENTS
We identified 79 bird, 20 mammal, and 6
amphibian and reptile species that use New
England salt marshes at some point in their life
history (Tables 1 and 2). Wildlife habitat
requirements were identified from accounts in
the Birds of North America (Poole and Gill
1992), an atlas of New England wildlife (DeGraaf
and Yamesaki 2001), literature surveys of salt
marsh bird species (e.g., Reinert and Mello 1995,
Benoit and Askins 2002, Shriver et al. 2004),
mammalian species accounts published by the
American Society of Mammalogists (e.g., Bekof
1977), unpublished reports, anecdotal
information from wetlands ecologists, and
personal observations. Salt marsh birds were
categorized as breeding (those species that have
been observed to nest in salt marshes) or foraging
(those which spend at least some portion of their
life histories feeding in salt marshes). Foraging
species are further divided into year-round,
summer-only, migrant, or winter-only foragers.
Birds in the latter two foraging categories use salt
marshes sporadically and some are rarely
encountered in marsh habitats. Mammals are
categorized as foragers (i.e., those species that
feed on salt marsh vegetation), predator
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Wildlife Habitat Value of New England Salt Marshes
Table 1. Birds known to inhabit New England salt marshes or use salt marshes as foraging or shelter habitat.
Group
Common Name
Species
Breeders
Foragers - year round
American oystercatcher
lapper rail
common tern
killdeer
laughing gull
least bittern
mallard
marsh wren
mute swan
red-winged blackbird
salt marsh sharp-tailed sparrow
seaside sparrow
swamp sparrow
Virginia rail
willet
American crow
American robin
bald eagle
belted kingfisher
black-bellied plover
Bonaparte's gull
cedar waxwing
common grackle
double-crested cormorant
European starling
fish crow
gray catbird
great black-backed gull
great horned owl
herring gull
house sparrow
mourning dove
northern cardinal
northern flicker
northern mockingbird
red-shouldered hawk
red-tailed hawk
rough-legged hawk
ring-billed gull
ring-necked pheasant
semipalmated sandpiper
short-eared owl
song sparrow
Haematopus palliatus
Rallus longirostris
Sterna hirundo
Charadrius vociferus
Lams atricilla
Ixobrychus exilis
Anas platyrhynchos
Cistothorus palustris
Cygnus olor
Agelaius phoeniceus
Ammodramus caudacutus
Ammodramus maritimus
Melospiza georgiana
Rallus limicola
Catoptrophorus semipalmatus
Corvus brachyrhynchos
Turdus migratorius
Haliaeetus leucocephalus
Ceryle alcyon
Pluvialis squatarola
Lams Philadelphia
Bombycilla cedrorum
Quiscalus quiscula
Phalacrocorax auritus
Sturnus vulgaris
Corvus ossifragus
Dumetella carolinensis
Lams marinus
Bubo virginianus
Lams argentatus
Passer domesticus
Zenaida macroura
Cardinalis cardinalis
Colaptes auratus
Mimus polyglottos
Buteo lineatus
Buteo jamaicensis
Buteo lagopus
Lams delawarensis
Phasianus colchicus
Calidris pusilla
Asio flammeus
Melospica melodia
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Wildlife Habitat Value of New England Salt Marshes
Foragers - summer American goldfinch
bank swallow
barn swallow
black-crowned night heron
chimney swift
common yellowthroat
eastern kingbird
glossy ibis
great blue heron
great egret
greater yellowlegs
green heron
least tern
lesser yellowlegs
little blue heron
osprey
snowy egret
spotted sandpiper
tree swallow
yellow-crowned night heron
Foragers - migration cattle egret
least sandpiper
semipalmated plover
semipalmated sandpiper
sora
Foragers - winter American black duck
American coot
American wigeon
blue-winged teal
brant
Canada goose
dunlin
green-winged teal
northern harrier
northern pintail
ring-necked duck
sanderling
snowy owl
Carduelis tristis
Riparia riparia
Hirundo rustica
Nycticorax nycticorax
Chaetura pelagica
Geothlypis trichas
Tyrannus tyrannus
Plegadis falcinellus
Aldea herodias
Aldea alba
Tringa melanoleuca
Butorides virescens
Sterna antiUamm
Tringa flavipes
Egretta caerulea
Pandion haliaetus
Egretta thula
Actitus macularia
Tachycineta bicolor
Nyctanassa violacea
Bubulcusibis
Calidris minutilla
Charadrius semipalmatus
Calidris pusilla
Porzana Carolina
Anas rubripes
Fulica americana
Anas americanus
Anas discors
Branta bernicla
Branta canadensis
Calidris alpina
Anas crecca
Circus cyaneus
Anas acuta
Aythya collaris
Calidris alba
Nyctea scandiaca
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Wildlife Habitat Value of New England Salt Marshes
Table 2. Mammals, amphibians, and reptiles known to inhabit New England salt marshes or use salt marshes as
foraging habitat.
Group1
Common Name
Species
Mammals
Foragers
Predators
Breeders
Amphibians / reptiles
black-tailed jackrabbit
eastern cottontail
least shrew
masked shrew
raccoon
Virginia opossum
white-tailed deer
coyote
fisher
long-tailed weasel
mink
red fox
river otter
striped skunk
meadow jumping mouse
meadow vole
muskrat
New England cottontail
Norway rat
woodland vole
common snapping turtle
eastern painted turtle
green frog
northern diamondback terrapin
northern water snake
spotted turtle
Lepus californicus
Sylvilagus florianus
Cryptotis parva
Sorex cinereus
Procyon lotor
Didelphis virginiana
Odocoileus virginianus
Canis latrans
Manes pennanti
Mustela frenata
Mustela vison
Vulpes vulpes
Lontra canadensis
Mephitis mephitis
Zapus hudsonius
Microtus pennsylvanicus
Ondatra zibethicus
Sylvilagus transitionalis
Rattus norvegicus
Microtus pinetorum
Chelydra serpentina serpentina
Chrysemys picta picta
Rana clamitans melanota
Malaclemys terrapin terrapin
Nerodra sipedon sipedon
Clemmys guttata
foragers are those who consume indigenous salt marsh flora or fauna; e.g., marsh grasses or resident
invertebrates such as bivalves. Predators will take advantage of prey when present; e.g., small mammals, birds
and eggs. Breeders are those that will potentially nest in some part of the marsh.
(i.e., those who will venture onto a salt marsh
to take advantage of prey when present), and
breeders (i.e., those that will potentially nest in
some part of the marsh).
While our framework as a whole uses
maximum wildlife species diversity and
abundance as a standard by which to assess salt
marsh habitat value, categorization of bird and
mammal species allows for flexibility in its
application. For example, to assess habitat value
for salt marsh foraging birds, one would first
identify the relevant species from Table 1, then
refer to Appendix 1 and the appropriate passages
in the text for specific habitat types and
component categories that are important for
these species. These categories could then be
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Wildlife Habitat Value of New England Salt Marshes
emphasized in an assessment by weighting their
values appropriately.
We identified common habitat types
associated with New England salt marshes, or
those that were reported as being used by at
least 3 bird or mammal species in published life
history accounts, unpublished reports, and
anecdotal information from local wetlands
ecologists (Table 3). The most commonly
reported habitat types, edge habitats, or
adjoining habitats for each bird and mammal
category, based on the published literature
including species life history accounts in the
Birds of North America (Poole and Gill 1992)
and Mammalian Species reports (e.g., Bekof
1977), are summarized in Tables 4 and 5. These
habitat types, as well as the habitat requirements
of salt marsh fauna, form the basis of the salt
marsh assessment components described in this
report.
WETLAND AND LANDSCAPE ASSESSMENT
COMPONENTS
Below we describe eight wetland and
landscape assessment components of New
England salt marshes (Table 6). Several of the
components, such as Salt Marsh Habitat Type,
Salt Marsh Vegetation, Salt Marsh Vegetative
Heterogeneity, and Connectivity and Associated
Habitat are directly based on or composed of the
different habitat types on the salt marsh
landscape or ecosystems that are linked to the
salt marsh. Other components, such as Degree
of Anthropogenic Modification, and
Surrounding Land Cover and Land Use reflect
the alteration of these habitats. The remaining
Table 3. Habitat types, edge habitats, and adjoining habitats of value to salt marsh wildlife.
Habitat types
Open water (< 60 cm)1
Tidal flat
Low marsh2
High marsh3
Pools
Pannes
Trees overhanging water
Wooded islands
Marsh-upland border
Phragmites
Edge habitats
Marsh-water edge
Tidal creek edge
Marsh-pool edge
Marsh-upland edge
Associated habitats
Sand or cobble beach
Coastal dunes or overwash
Other salt marsh wetland
Brackish wetland or pond
Freshwater wetland or pond
Upland meadow
Upland forest
Shallow open water less than 60 cm in depth
2Smooth cordgrass (Spartina a/fer/2/$ara)-dominated low marsh
3Salt meadow often dominated by Spartina patens and forbs
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Wildlife Habitat Value of New England Salt Marshes
Table 4. Most commonly reported1 habitat types, edge habitats, and associated habitats used by salt marsh
breeding and foraging birds.
All birds Breeders Foragers - year round
1) High marsh 1) High marsh 1) High marsh
2) Low marsh 2) Marsh-upland edge 2) Marsh-upland border
3) Tidal flats 3) Low marsh 3) Low marsh
4) Shallow open water 4) Tidal flats 4) Upland forest
5) Upland forest 5) Shallow open water 5) Sand or cobble beach
Foragers - summer Foragers - migration Foragers - winter
1) High marsh 1) Low marsh 1) Shallow open water
2) Shallow open water 2) Tidal flats 2) Marsh-upland edge
3) Low marsh 3) Shallow open water 3) Low marsh
4) Marsh-upland edge 4) Sand or cobble beach 4) Tidal flats
5) Marsh-water edge 5) Marsh-upland edge 5) Upland meadow
'Sources for avian wildlife habitat information include Birds of North America (Poole and Gill 1992),
DeGraaf and Yamesaki 2001, and literature cited in Appendix 1.
Table 5a. Most commonly reported1 habitat types, edge habitats, and associated habitats used by grazing,
predator, and breeding salt marsh mammals, as well as all mammals that use salt marshes.
All mammals Grazers Predators Breeders
1) High marsh 1) High marsh 1) Low marsh 1) High marsh
2) Marsh-upland border 2) Marsh-upland border 2) Freshwater wetland 2) Marsh-upland border
3) Low marsh 3) Low marsh 3) High marsh 3) Upland meadow
4) Upland meadow 4) Upland meadow 4) Upland meadow 4) Upland forest
5) Freshwater wetland 5) Upland forest 5) Tidal flats 5) Low marsh
Sources for mammalian wildlife habitat information include mammalian species accounts published by
the American Society of Mammalogists, DeGraaf and Yamesaki 2001, and literature cited in Appendix 2.
Table 5b. Most commonly reported1 salt marsh habitat types, edge habitats, and associated habitats used by
amphibians and reptiles.
All amphibians and reptiles
1) Freshwater wetland or pond
2) Brackish wetland or pond
3) Marsh-upland border
4) Marsh-water edge
5) Tidal flat
Sources for amphibian wildlife habitat are given in Appendix 2.
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Wildlife Habitat Value of New England Salt Marshes
components (Salt Marsh Size, Salt Marsh
Morphology) take into account the size,
morphology, and landscape position of the
marsh, which may be important to territorial
species and those that require adjacent upland
habitats. Salt marsh size and morphology may
also be useful in pre-classifying marshes prior to
assessment.
Together these eight wetland and landscape
assessment components comprise a framework
that can be used to assess and evaluate salt
marsh wildlife habitat value.
I. Salt Marsh Size Class
Salt marshes along the New England coast
include narrow, discrete fringe marshes less than
10 ha in area and salt meadow complexes of up
to 2000 ha. Mean salt marsh size ranges from
40.2 ha for marshes in southern New England to
174.8 ha for marshes in the Gulf of Maine
(Shriver et al. 2004). In general, large wetlands
are considered to be of greater value to wildlife
as habitat, although smaller marshes may in
some cases provide important habitat for
endemic species or those with specific habitat
requirements. Several studies have reported a
positive relationship between the number of
bird species and wetland area (Brown and
Dinsmore 1986, Craig and Beal 1992), and others
have documented area dependence for species
richness of salt marsh breeding birds,
particularly those that are short
Table 6. Wetland and landscape assessment components of New England salt marshes and their associated
categories. The categories represent habitat, morphological, vegetation or land use types, or classes
that represent a marsh characteristic (size class, degree of anthropogenic modification, level of
heterogeneity). Criteria are those parameters that may be used in an assessment protocol to rank
marshes, e.g., a marsh with a greater number of salt marsh habitat types may rank above a marsh
with fewer types, depending on goal of the assessment protocol.
Component
Categories
Criteria
I. Salt Marsh Size Class
II. Salt Marsh Morphology
III. Salt Marsh Habitat Types
Very small (under 5 ha)
Small (5 - 25 ha)
Medium-sized (26 - 125 ha)
Large (126 - 200 ha)
Very large (over 200 ha)
Salt meadow marsh
Meadow / fringe marsh
Wide fringe marsh
Narrow fringe marsh
Marine fringe marsh
Shallow open water
Tidal flats
Low marsh
Trees overhanging water
High marsh
Pools
Marsh area
Marsh morphology
Presence or abundance
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Wildlife Habitat Value of New England Salt Marshes
Component
Categories
Criteria
IV. Extent of Modification
V. Salt Marsh Vegetation
VI. Vegetative Heterogeneity
VII. Surrounding Land Cover
VIII. Connectivity
Pannes
Wooded islands
Marsh-upland border
Phragmites
Little to no ditching
Moderate ditching
Severe ditching
Little to no tidal restriction
Moderate tidal restriction
Severe tidal restriction
Aquatic plants
Emergents
Shrubs
Trees
Vines
High heterogeneity
Moderate heterogeneity
Low heterogeneity
Open water
Natural land
Maintained open land
Developed land
Sand or cobble beach
Coastal dunes or overwash
Other salt marsh wetland
Brackish wetland or pond
Freshwater wetland or pond
Upland meadow
Upland forest
Degree of modification
Presence or abundance
Number of habitat edges
Presence or area
Presence or area
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Wildlife Habitat Value of New England Salt Marshes
grass meadow specialists (Benoit and Askins
1992, Shriver et al. 2004). These findings imply
that larger salt marshes may provide greater
relative habitat value for some species of
breeding birds. They also point to the
importance of habitat fragmentation in
determining species richness. The negative
effects of habitat fragmentation on bird species
richness has been demonstrated for forest and
grassland birds, where it has been reported that
area sensitive species tend to have lower
densities in small habitat patches versus larger
blocks of continuous habitat (Askins et al. 1990,
Vickery et al. 1994). Fragmentation has been
shown to influence bird distribution in New
England salt marshes, with larger habitat
patches generally supporting more species
(Clarke et al. 1984, Benoit and Askins 1999,
2002). Larger and less fragmented marshes may
provide greater habitat value to wildlife that are
sensitive to human activities, since peripheral
disturbances will have less of an effect on the
inner part of the marsh (Golet and Larson 1974).
Larger marshes will also have less relative edge
habitat per marsh area, which may mitigate
processes such as nest predation that may be
correlated with marsh edge (Johnson and
Temple 1990). Large, contiguous blocks of
wetland will tend to contain a greater diversity
of habitat types, and are therefore more likely to
meet all species' habitat requirements (Burke
and Nol 1998). However, even small or fringe
salt marshes have habitat value, particularly for
foraging species. For example, a study of salt
marsh habitat use in Narragansett Bay, RI
showed consistent densities of foraging herons
and egrets at sites ranging from 2 - 70 ha (Trocki
2003).
Benoit and Askins (2002) reported minimum
area requirements for six bird species that breed
in Connecticut salt marshes. They found that
when they considered salt marsh fragments to
be defined as those separated by broad barriers
(>500 m of open water, or >50 m of upland
habitat), minimum area requirements ranged
from 8 to 138 ha (Benoit and Askins 2002).
Seaside sparrow territories of <1 ha were
reported in ditched marshes in Massachusetts
(Marshall and Reinert 1990), but nonetheless
these species were absent in marshes of less than
67 ha in the Connecticut study. Similarly,
sharp-tailed sparrows have reported home-
ranges of 1.2 - 5.7 ha (Wolfenden 1956,
Greenlaw and Rising 1994), but were not
reported in Connecticut marshes less than 10 ha.
Willet Catoptrophorus semipalmatus were the
most area sensitive, absent in marshes of less
than 138 ha, but this may have been confounded
by recent recolonization of salt marshes after
extirpation from hunting and egg collection
(Bevier 1994).
Mammals that utilize salt marshes exhibit a
wide range of home range sizes, depending upon
Black-crowned night heron Nycticorax
nycticoras (Photo by Lee Karney, US FWS).
whether they forage near nests and burrows or
follow and chase mobile prey across larger areas.
For example, meadow jumping mouse Zapus
hudsonius and meadow vole Microtus
pennsylvanicus have home ranges of less than 1
10
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Wildlife Habitat Value of New England Salt Marshes
ha, while home ranges of the wide ranging
coyote Cam's latrans, red fox Vulpes vulpes, and
mink can extend for thousands of hectares
(Whitaker 1972, Harrison et al. 1989, Reich
1996, Lariviere 1999).
We adopted the mean of the minimum area
requirements for salt marsh breeding birds
(about 60 ha) reported by Benoit and Askins
(2002) as the mid-point of our middle salt marsh
size category. We then divided the range of
areas between 5 and 200 ha among three size
classes to derive the following salt marsh size
categories:
1. Very small:
2. Small:
3. Medium-sized:
4. Large:
5. Very large:
under 5 ha.
5 - 25 ha.
26 -125 ha.
126 - 200 ha.
over 200 ha.
Use of this component in a wildlife habitat
assessment
Based on the available information about
species habitat requirements, an assessment of
salt marsh wildlife habitat value should include
a consideration of marsh size. Since for a
majority of species habitat value increases with
marsh size, a ranking scheme should value larger
over smaller marshes. The five categories
presented above could be used to rank salt
marshes by assigning increasing value to the
ranking as size class increases. However, we
reiterate that even small or fringe salt marshes
may have significant habitat value for wildlife
species. For example, a given salt marsh
regardless of size may provide important habitat
for an endemic or endangered species. Smaller
marshes have also been shown to support
significant numbers of foraging herons and
egrets (Trocki 2003); quite possibly these
marshes may be appealing to these species
because their small size discourages use by
potential avian and mammalian predators.
Situations of this sort can be mitigated to some
extent by including this assessment framework
as one component in a multivariate decision-
making model such as that proposed by Larson
(1976) for fresh-water wetlands. Models of this
sort will first determine whether a wetland
possesses out-standing or unique attributes (e.g.,
uncommon geomorphological features,
archaeological value). This approach can
identify marshes that may rank low in an overall
assessment of wildlife species diversity but
nonetheless may have important intrinsic value.
II. Salt Marsh Morphology
In addition to its size, the morphology of a
salt marsh may affect habitat value. For
example, a fringing salt marsh is by definition
narrow, but may cover a long extent of a
shoreline and hence have a large area.
However, because it provides little buffer from
peripheral human disturbance and is often
dominated by low marsh with few additional
marsh habitat types, it may be of limited value
to wildlife. Conversely, a meadow marsh of
equal area may buffer wildlife in its interior
from peripheral disturbance, and is also more
likely to consist of several salt marsh habitat
types. It is therefore important to consider salt
marsh morphology along with the area of the
marsh when determining wildlife habitat value.
The Salt Marsh Morphology component is
derived from the concept of wetland cover type
first introduced by Stewart and Kantrud (1971)
for prairie pothole wetlands and adapted by
Golet and Larson (1974) for freshwater wetlands
in the northeast. Cover type acknowledges the
importance of the proportion of vegetative cover
and open water to wetland wildlife, with the
11
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Wildlife Habitat Value of New England Salt Marshes
most important factor being the length of edge
between cover and water per unit area of
wetland. This element is particularly important
for species that utilize open water for foraging
but need the presence of nearby vegetative
cover for shelter. In freshwater wetlands, cover
type is important for breeding waterfowl
because the edge between vegetative cover and
water provides isolation of breeding pairs,
protection from exposure to strong winds, and
greater production and diversity of food
organisms (Baldassare and Bolen 1994). The
marsh-water edge may provide similar functions
for wintering waterfowl in New England salt
marshes. In addition, the marsh-water edge
may be important for species that forage in
shallow water and occasionally use nearby
vegetated areas for protection. For example,
plovers and sandpipers feed on exposed tidal
flats at the marsh border and dart in and out of
sparse Spartina alterniflora for protection or to
pursue prey organisms (Recher 1966, Johnsgaard
1981). Other species, including willet and
killdeer Charadrius vociferus, may take
advantage of the increased prey abundance and
diversity at the marsh-water edge (Danufsky and
Colwell 2003, Maimone-Celorio and Mellink
2003). Wading birds may occasionally forage at
the marsh-water edge when it is flooded to take
advantage of the camouflaging effect of
vegetation (Hancock and Kushlan 1984).
Marsh cover type as defined for freshwater
wetlands may not be an appropriate metric for
evaluating the wildlife habitat value of salt
marshes. While the impact of marsh-water edge
may be similar for salt marsh species, cover type
is confounded somewhat by tidal inundation
and marsh geomorphology. Salt marshes are by
definition bordered by estuarine or marine open
water; defining what proportion of the adjoining
open water is to be considered when
determining cover type by estimating percent
vegetative cover (i.e., what percentage of the
wetland area is occupied by open water) can be
problematic. We therefore propose an alternate
classification based on the geomorphology of salt
marshes along the New England coast. Classes
of salt marsh morphology will represent varying
amounts of marsh-water edge and marsh-upland
edge in relation to wetland area. This
classification acknowledges that edge habitat,
which may be beneficial to some species, needs
to be balanced by sufficient interior area to
buffer wildlife from unfavorable edge processes
(e.g., increased predation risk, human
disturbance). Five classes of salt marsh
morphology are shown in Figure 1 and
described below:
1.) Salt-meadow marsh: The salt meadow marsh
is generally a back-barrier or basin marsh
with extensive systems of wide and narrow
creeks interspersed with large expanses of salt
meadow marsh interior. Wide, basin-like
marshes typically have a distinct bank
between open water and marsh and support a
greater diversity of habitat types and features,
including high marsh and border plant
communities, marsh pannes and pools, and
inter- and sub-tidal creeks. Salt meadow
marshes may be ditched or un-ditched (Figure
la). This salt marsh type is generally of the
greatest value to wildlife species, because of
the potential for the existence of a number of
habitat types, and the degree of protection
and buffering afforded from the surrounding
landscape.
12
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Wildlife Habitat Value of New England Salt Marshes
Figure 1. Salt marsh morphology categories of New England marshes.
a) Salt-meadow marsh. Back-barrier or basin marsh with extensive creek systems
interspersed with salt meadow marsh interior. May be ditched or un-ditched.
b) Meadow / fringe marsh. Typically consists of areas of salt meadow marsh interspersed
with narrow or wide fringe marsh.
13
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Wildlife Habitat Value of New England Salt Marshes
c) Wide fringe marsh. Typically dominated by low marsh but can contain some patches of
high marsh vegetation, with a small number of narrow creeks (Figure Ic). Marsh width
ranges from 10-50 m in width from seaward to landward marsh edge.
/ :^i
-*-•" : '
d) Narrow fringe marsh. Consists of a narrow belt of vegetation dominated primarily by the
low marsh Spartina alterniflora with few creeks. Generally less than 10 m in width from
seaward to landward marsh edge.
14
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Wildlife Habitat Value of New England Salt Marshes
e) Marine fringe marsh. A narrow fringe marsh that is bordered on the seaward edge by
unprotected open water; i.e., not located within a cove or embayment. Typically have
significant edge exposed to open water and high exposure to waves and prevailing winds.
2.) Meadow / fringe marsh: This marsh type
consists of areas of salt meadow marsh
interspersed with fringe marsh. Fringe marsh
may be narrow or wide, and predominantly
consists of low marsh. These marshes may be
ditched or un-ditched (Figure Ib). Because
meadow / fringe marshes can contain a
number habitat types, they can provide
significant wildlife value to most bird and
mammal species.
3.) Wide fringe marsh: Fringe marshes form in
bands along shorelines where there is some
protection from wave and wind but slope
limits the landward extent of the marsh.
Wide fringe marshes are often dominated by
low marsh but can contain some patches of
high marsh vegetation, typically grade from
open water to upland, and have a small
number of narrow creeks (Figure Ic).
Generally, these marshes range from 10-50
m in width from seaward to landward marsh
edge. This salt marsh type has less habitat
value to most species, although a wide fringe
marsh may provide important foraging habitat
for low marsh foraging and breeding birds
and mammals.
:.) Narrow fringe marsh: This marsh type
consists of a narrow belt of vegetation
dominated primarily by the low marsh
Spartina alterniflora with few creeks. Narrow
fringe marshes are characterized by high
amounts of both marsh-water and marsh-
upland edge per wetland area. These marshes
are generally less than 10 m in width from
seaward to landward marsh edge. This marsh
type is characteristic of areas impacted by
urbanization where a marsh has been filled to
accommodate adjacent development (Figure
Id), but can also be found in undisturbed
areas. Narrow fringe marshes provide the
least value to wildlife species, because they
are generally composed of only a few habitat
types, and offers little protection and
buffering from the surrounding landscape.
.) Marine fringe marsh: narrow fringe marsh
that is bordered on the seaward edge by
15
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Wildlife Habitat Value of New England Salt Marshes
unprotected open water; i.e., not located
within a cove or embayment. These marshes
have significant edge exposed to open water
and gain little to no protection from upland
environments. Marine fringe marshes have
high exposure to waves and prevailing winds,
and hence their habitat value to wildlife may
be limited (Figure le).
Use of this component in a wildlife habitat
assessment
In an assessment of salt marsh wildlife
habitat value, salt marsh morphology and size
class may be used to stratify salt marshes under
consideration (i.e., pre-classify a set of marshes
into categories) such that, for example, salt
meadow marshes are compared and ranked
relative to other salt meadow marshes and
separate from wide fringe marshes.
Alternatively, since we can assign relative
habitat value to the salt marsh morphology
categories, these could be used in an assessment
by weighting the categories with salt meadows
marshes having the most and narrow fringe
marshes the least habitat value.
III. Salt Marsh Habitat Type
Interaction of tidal inundation with the
geomorphology of salt marshes results in belts of
halophytic vegetation from the seaward edge of
the marsh toward the upland (Miller and Egler
1950, Redfield 1972, Nixon 1980). Chapman
(1940) first described this general pattern of
zonation in New England salt Marshes as one
consisting of: i) submergent sub-tidal vegetation,
e.g., Zostera marina, ii) tidal flats; iii) low marsh
dominated by smooth cordgrass Spartina
altemiffora, iv) high marsh dominated by salt
meadow cordgrass Spartina patens, and v)
marsh-upland border dominated by Juncus spp.
We retain these five zones as distinct micro-
habitat types in New England salt marshes,
replacing "submergent sub-tidal vegetation"
with a shallow open water habitat that may or
may not be vegetated. We also identify five
microhabitat types that arise from differences in
the geomorphology, tidal inundation, and the
composition and complexity of salt marsh
vegetation: pannes, marsh pools, trees
overhanging water, wooded islands, and
Phragmites australis (Figure 2). Below we
describe wildlife use of these habitat types in
their order of occurrence from the seaward to
the landward edge on the salt marsh.
Shallow open water (<60cm depth)
Shallow open water consists of estuarine
water seaward of the low marsh edge or tidal
waters that are part of large creeks within the
marsh itself. This habitat is used by foraging
herons and egrets during the breeding season
(Willard 1977, Ramo and Busto 1993).
Additionally, migrating herons and egrets rely
heavily on these foraging habitats as stopover
sites during spring and summer migration
(Chavez-Ramirez and Slack 1995). Water height
is particularly important to these species. Based
on Birds of North America species accounts and
other studies a maximum water depth of 60 cm
for is suggested for herons and egrets foraging in
New England salt marshes (Custer and Osborn
1978, DuBowy 1996, Matsunaga 2000; Table 7).
Shallow open water is also important for
wintering waterfowl, particularly dabbling
ducks that use these areas for foraging on
submerged macroalgae or submergent
vegetation (Erwin et al. 1994, Mowbray 1999,
Longcore et al. 2000, Drilling et al. 2002).
Maximum foraging depths may differ for these
species. Several species of diving ducks
16
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Wildlife Habitat Value of New England Salt Marshes
Figure 2. Salt marsh habitat types and the occurrence within a typical New England salt marsh.
Not shown trees overhanging water.
(including bufflehead Bucephala albeola and
scaup Aythya spp.) may also use shallow sub-
tidal areas to forage for benthic macro-
invertebrates (Gauthier 1993). Mammals
including mink Mustela vison and fisher Maries
pennanti utilize this habitat as well as the
adjacent tidal flats when feeding on fish and
birds (Powell 1984, Lariviere 1999).
Tidal flats
Tidal flats are areas of mud or sand on the
seaward edge of a marsh or creek that are
exposed at low tide. Tidal flats are important
foraging areas for a number of salt marsh bird
species, including foraging, breeding and
wintering species (Appendix 1). In our review of
the published literature, tidal flats are used by 17
of the 79 bird species. Tidal flat substrate
includes both mud and fine sediments and sandy
areas of course grain sediments. Each substrate
has a unique assemblage of benthic fauna,
consisting primarily of invertebrates that reside
in or on the sediment. Although there is some
overlap in benthic species between the two
substrates, each provides a unique foraging
habitat for different assemblages of marsh bird
species (Appendix 1). For example, yellowlegs
Tringa spp. feed almost exclusively on exposed
mud flats on a diet that includes amphipods and
other small crustaceans (Elphick and Tibbitts
1998, Tibbitts and Moskoff 1999). Other birds
using mud flats include rails, sparrows, several
duck species, willets Catoptrophorus
semipalmatus, and occasionally herons and
egrets. Sandpipers and plovers will preferentially
forage on more sandy sediments, feeding on
polychaetes, gastropods, and small bivalves.
Oystercatchers Haematopus palliatus also forage
on tidal flats that contain sufficient densities of
bivalves (Nol and Humphrey 1994). As a
foraging strategy, common snapping turtles
Chelydra serpentina serpentina will burrow in
17
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Wildlife Habitat Value of New England Salt Marshes
salt marsh tidal flats and wait for prey to pass
near (Babcock 1971).
Low marsh
The low marsh in New England salt marshes
is described as the belt of emergent vegetation at
the seaward edge of the marsh that is typically
dominated by Spartina alterniflora (Miller and
Egler 1950, Niering and Warren 1980, Nixon
1980). The landward edge of the low marsh is
often defined by the extent to which the marsh
is consistently flooded by tides, i.e., mean high
water (Redfield 1972). Low marsh habitat is
used for nesting or foraging by 43 of the 79 bird
species (Appendix 1), and is the second most
frequently used of all salt marsh habitats (Table
4). Low marsh vegetation is important breeding
habitat for seaside sparrow Ammodramus
maritimus, willet Catoptrophorus semipalmatus,
and on occasion salt marsh sharp-tailed sparrow
Ammodramus caudacutus. The relatively low
stem density of stands of Spartina alterniflora,
combined with its wide, tall leaves, provide an
ideal microhabitat for nests of these species:
sturdy stems are used to support nests above the
substrate and also to dissipate winds and
maintain high temperatures and humidity.
Seaside sparrows require relatively large (> 0.5
ha) expanses of tall form Spartina alterniflora,
and build their nests several centimeters above
the substrate in an attempt to avoid flooding
Greenlaw and Rising 1994). The low stem
density of Spartina alterniflora, along with the
scouring action of daily tides, helps to keep the
underlying sediments clear of debris. This gives
smaller species that forage in the low marsh
access to the bare sediment (and resident benthic
invertebrates) between stems, while still
providing protective cover. When flooded, the
low marsh is also occasionally used as forging
habitat by larger birds such as herons and egrets
(Appendix 1).
Trees overhanging water
Although more common in marshes in the
southeastern U.S., trees can occasionally be
located sufficiently close to the marsh-water
edge such that tree limbs will overhang open
water. This provides a preferred foraging habitat
for cattle egrets Bubulcus ibis, green herons
Butorides virescens, black-crowned night herons
Nycticorax nycticora, and belted kingfishers
Ceryle alcyon (Davis and Kushlan 1994; Hamas
1994; Appendix 1).
High marsh
In contrast to low marsh vegetation, Spartina
patens, which dominates the high marsh in New
England salt marshes, is a short fine grass with
high stem density. The high marsh may also be
populated with several other salt marsh grasses
and several species of forbs. The combination of
dense vegetation, vegetative diversity and
infrequent flooding results in a habitat that
supports a greater diversity and abundance of
invertebrates, particularly insects. This
vegetative heterogeneity also results in a
favorable habitat for foraging bird species,
particularly those that feed on flying insects
(Appendix 1). Swallows, red-winged blackbirds
Agelaius phoeniceus, and sparrows, as well as
other occasional passerines, utilize high marsh
habitats for foraging. Furthermore, the dense
vegetation characteristic of the high marsh,
along with less frequent flooding, provides
nesting habitat for sharp-tailed sparrows,
waterfowl and least bittern Ixobrychus exilis.
Sharp-tailed sparrows reportedly will locate
nests where they will only be flooded by
extreme spring tides, and often successfully
18
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Wildlife Habitat Value of New England Salt Marshes
Table 7. Body mass, optimal foraging water depth, and tarsus length of wading birds in the family Ardeidae that utilize New England salt marshes.
Common Name Species
Black-crowned night heron Nycticorax nycticorax
Cattle egret Bubulcus ibis
Great egret Egretta alba
Great blue heron Aldea herodias
Green heron Butorides virescens
Little blue heron Egretta caerulea
Snowy egret Egretta thula
Yellow-crowned night heron Nyctanassa violacea
'Average male ± SD (when reported) / Average female ± SD
2Tarsus = lowest segment of leg, before toes; average male ±
3 Average adult mass.
4References
A) Dunning 1993.
B) Palmer 1962.
C) Browderl973.
D) Gross 1923.
E) Quinney and Smith 1979.
Optimal
Body mass1 Water Depth Tarsus length2 Reference4
913±115g/827±69g -- 18.3±0.5 mm / 17.7±0.3 mm D
371.8 g/ 359.8 g -- 78.6 mm / 80.4 mm C
935±134g/812g 20 -40 cm 167 mm/ 137±14mm A,B
2230±760g3 25 -60 cm 179±12 mm/ 171±12 mm E
241 g3 <5cm 53.0 mm /51.2mm F
364±47g/315g 5 - 15 cm 96.2 mm/ 88.1 mm B,G
369 g3 8cm 97.1 mm /89.6mm B
716±18g/649±16g 15 -25 cm 99 mm /97mm H,I
(when reported).
SD (when reported) / Average female ± SD (when reported).
F) Niethammer and Kaiser 1983.
G) Rodgers and Smith 1995.
H) Blake 1977.
I) Hartmanl955.
19
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Wildlife Habitat Value of New England Salt Marshes
re-nest immediately after the first flooding,
allowing young to fledge before the next spring
tide (Post and Greenlaw 1982, DeRagon 1988).
Several species (e.g., black-bellied plover
Pluvialis squatarola) use the high marsh for
roosting during high tide when feeding grounds
are covered (Paulson 1995). Overall, 47 of the
79 bird species use the high marsh, and it is the
most frequently used of all salt marsh habitats
by birds (Table 4).
Coyote Canislatrans
(Photo by R.H. Barrett, US FWS).
A number of mammal species, including
black-tailed jackrabbit Lepus californicus,
eastern cottontail Sylvilagus florianus, and
meadow jumping mouse Zapus hudsonius feed
on forbs that are found in the high marsh
(Currie and Goodwin 1966, Chapman et al.
1980, Whitaker 1972; Appendix 2). Glossy ibis
Plegadis falcinellus often forage extensively on
the high marsh, particularly in marshes that are
adjacent to or near agricultural fields (Trocki
2003). Herons and egrets have also been
observed to forage in high marsh vegetation at
high tide (Hancock and Kushlan 1984, Custer
and Osborn 1978). All told, this diverse habitat
type is reportedly used for nesting and foraging
by 17 of the 79 marsh bird species, and 14 of the
20 mammal species (Tables 7 & 8).
Pools
Marsh pools are a common feature in New
England salt marshes, although their abundance
in the marsh landscape may be tied to the extent
to which the marsh has been subjected to
mosquito ditching (Adamowicz and Roman
2005). Miller and Egler (1950) describe pools as
shallow (seldom deeper than 30 cm), typically
containing submergent vegetation (Rupia
maritime), and inhabited by a variety of nekton
species. Pools will generally form in depressions
in the marsh surface that can retain tidal waters,
and would therefore be expected to contain
many of the same prey species as are found in
the surrounding open water habitat (Raposa and
Roman 2001). However, varying water depths
and different pool water salinities may alter
community composition. For example, pools
that are located some distance from tidal waters
and therefore experience only infrequent
flooding may take on the characteristics and of
brackish/freshwater ponds. Pools in the salt
marsh landscape are therefore a diverse habitat
type that seemingly could provide foraging
habitat for a number of bird species.
Interestingly, only 3 species have been
specifically identified in life history accounts as
using marsh pools (glossy ibis, lesser yellowlegs
Tringa flavipes, and snowy egrets Egretta thula),
although we have observed on numerous
occasions many of the same heron and egret
species that feed in shallow water foraging in
salt marsh pools. A study of bird use of ditched
versus unditched marshes in Narragansett Bay
showed greater bird use in unditched marshes,
which may have been related to the greater
density of marsh pools (Reinert et al. 1981).
However, an important consideration may be
the amount of available foraging habitat within
a pool (i.e., water depth <60 cm). Depending on
20
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Wildlife Habitat Value of New England Salt Marshes
marsh geomorphology, pools may have steep,
erosional edges and depths that are too great for
use by wading birds. Several marshes during a
recent survey of salt marshes in Narragansett
Bay were found to have average depths of
greater than 60 cm in many of their pools, and
foraging by herons and egrets was not observed
in the deeper pools (K. Raposa and T. Kutcher,
personal communication). Therefore, available
foraging habitat should be considered in
addition to the presence of pools when
determining wildlife habitat value.
Pannes
Particularly in the high marsh, slight
depressions in the salt marsh surface may retain
water that subsequently becomes highly saline
as a result of evaporation. These areas may
develop into pannes, or bare, exposed
depressions in the marsh that can at times be
filled with shallow water (Wiegert and Freeman
1990). The habitat value of pannes results from
their being devoid of vegetation and therefore
providing foraging areas for species that prefer
low-lying, un-vegetated substrates. However,
pannes are physically harsh habitats
characterized by high soil salinities and frequent
flooding and drying, and little is known of their
benthic communities. Additionally, mid-
elevation pannes in northern New England salt
marshes are typically colonized by a number of
stress-tolerant forbs, owing to differences in
climate (less solar radiation and cooler
temperatures results in less potential for high
soil salinities) and a lesser extent of ditching and
draining (Ewanchuk and Bertness 2004a,b).
Forb pannes in northern New England marshes
may not provide the same wildlife habitat value
as un-vegetated southern marsh pannes. Species
known to forage in un-vegetated or low-lying
areas include snowy egret, lesser yellowlegs,
glossy ibis, sharp-tailed sparrow, and seaside
sparrow. Additionally, species that forage on
tidal flats or exposed mud may utilize pannes
(Appendix 1).
Wooded islands
Wooded islands are elevated areas within the
high marsh dominated by trees. Species may
include red maple Acer rubrum, black cherry
Prunus seratina, black oak Quercus velutina,
pitch pine Pinus rigida, black gum Nyssa
sylvatica, willow Salixspp., and alder Alnus spp.
Although small in area, wooded islands function
as habitat for several species, particularly as
roost sites for great egrets Aldea alba, great blue
herons Aldea herodias, and black-crowned night
herons Nycticorax nycticorax (Appendix 1).
These areas have the potential to provide
breeding habitat for herons and egrets, although
it is unclear whether they would be of sufficient
area to provide this function, particularly for
colonial breeders (Butler 1992, McCrimmon et
al. 2001).
Marsh-upland border
In New England salt marshes, the habitat
located at the upland margin of the marsh is
dominated by salt marsh shrubs Iva fructescens
and sea myrtle Baccharis halimifolia, as well as
brackish / upland sedges (Carex spp., Scirpus
spp.), rushes (Juncusspp.), and forbs (e.g., marsh
mallow Althaea officinalis, salt marsh aster Aster
spp.). The marsh-upland border can be rather
broad depending on marsh topography, and is of
value to a number of species for foraging and
nesting. Least bittern, clapper rail Rallus
longirostris, and Virginia rail Rallus limicola are
known to use marsh shrubs as breeding habitat
(Gibbs et al. 1992, Conway 1995, Eddleman and
21
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Wildlife Habitat Value of New England Salt Marshes
Conway 1998). Several passerine species utilize
this habitat type for foraging, including gray
catbird Dumetella carolinensis, willow
flycatcher Empidorax traillii, and eastern
kingbird Tyrannus tyrannus. Waterfowl,
including American black duck, mallard, and
Canada geese Branta canadensis, may also use
this habitat type for roosting. The presence of
waterfowl and marsh bird nests make this
habitat attractive to mammals and reptiles (e.g.,
northern water snake Nerodra sipedon sipedoii)
that feed on breeding birds and their eggs
(Appendix 2). For example, coyote Canis
latrans, red fox Vulpes vulpes, and striped skunk
Mephitis mephitis have been known to feed on
waterfowl and their eggs (Verts 1967, Bekoff
1977, Lariviere and Pasitschniak-Arts 1996).
Mink Mustela vison
Phragmites
Dense stands of common reed Phragmites
australis at the upland edge of salt marshes are a
widespread feature of southern New England
marshes, particularly in areas subject to high
nutrient inputs. This tall, erect perennial was
long thought to have little or no wildlife habitat
value, however recent studies have shown that
some bird species will nest in Phragmites stands
(e.g., Benoit and Askins 1999). This may be a
result of adaptation: for example the marsh wren
Cistothorus palustris and swamp sparrow
Melospiza georgiana are both marsh specialist
that nest in tall, reedy vegetation, preferably
cattail Typha angustifolia, but have been found
to nest in Phragmites stands that have replaced
cattails (Mowbray 1997, Benoit and Askins
1999). All told, we identified 10 species that use
Phragmites for nesting or foraging habitat. In
addition to marsh wren and swamp sparrows,
little blue heron Egretta caerulea, least bittern,
and mallard have been documented to nest in
Phragmites (Gibbs et al. 1992, Rodgers and
Smith 1995, Drilling et al. 2002). Recently, tree
swallows have been observed foraging for
insects over Phragmites stands on Cape Cod salt
marshes (J. Portnoy, personal communication).
Use of this component in a wildlife habitat
assessment
Salt marsh habitat type can be included in an
assessment of wildlife habitat value of New
England salt marshes by assigning a relative
value to the presence of each habitat type, or
assigning a value to a marsh based on the
number of habitat types present. How these
components are ranked or scored would depend
on the goal of the assessment. For example, if
the goal is to assess salt marsh habitat for
maximum species diversity, the presence of
many habitat types in a wetland would be
emphasized. Alternatively, if habitat value was
assessed for a guild of species, presence of
suitable habitat for the species under
consideration would be given more weight in an
overall assessment.
IV. Extent of Anthropogenic Modification
A majority of the salt marshes in New
England have been subject to some degree of
human modification (Adamovicz and Roman
2005). Human impacts at the local scale include
22
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Wildlife Habitat Value of New England Salt Marshes
those that directly modify or destroy salt marsh
habitat such as dredging, diking, spoil dumping,
grid ditching, canal cutting, and salt hay farming
(Kennish 2001). Salt marshes in New England
have been extensively ditched, and by 1938 an
estimated 90% of the salt marshes from Maine to
Virginia had been ditched in order to reduce
breeding habitat for the marsh mosquito
Ochlerotatus sollicitans (Bourn and Cottam
1950). Ditching typically leads to lowered water
table levels and draining of the marsh surface,
which in turn alters marsh habitat. In addition
to ditching, restriction of tidal flow to the marsh
caused by under-sized culverts or bridges,
causeways, manmade dikes, naturally occurring
berms or shelves can lead to large-scale changes
in marsh topography and vegetation patterns
(Esselink et al. 1998, Sturdevant et al. 2002).
Ditching and tidal restriction may lead to a
reduced density of pools in ditched salt marshes
(Adamowicz and Roman 2005), decreases in low
marsh vegetation (Sun et al. 2003), and increases
in the number of un-vegetated pannes and in
the extent of Phragmites australis (Ewanchuk
and Bertness 2004b). These changes in the
topography and vegetative structure of the
marsh may in turn influence patterns of
utilization by wildlife, and hence affect salt
marsh wildlife habitat value (Wolfe 1996).
Ditching and tidal restriction may differ in
the degree to which they influence salt marsh
wildlife habitat value. As described, most
ditching diminishes wildlife habitat value,
particularly for those species which rely on
marsh pools. However, ditching may in some
cases increase the occurrence of un-vegetated
pannes, and therefore increase foraging
opportunities for species that utilize panne
habitats. Tidal restriction can cause a decrease
in vegetative heterogeneity, but can also lead to
the formation of new marsh habitats such as
semi-permanent brackish ponds favored by
several species. We therefore classify ditching
and tidal restriction from least impact to highest
in the following categories.
Degree of ditching
1.) Little to no ditching. Marsh supports as
intact and natural system of wide and
narrow creeks, and generally have a
density of marsh pools (Figure 3a).
2.) Moderate ditching. Ditches are present
and may be numerous, but natural creeks
still intact and present. Marshes have a
moderate density of marsh pools
(Figure 3b).
3.) Severe ditching. Marshes show extensive
regular pattern of man-made ditches,
contain few or no natural creeks, and are
characterized by low density of marsh
pools (Figure 3c).
Degree of tidal restriction
1.) Little to no tidal restriction. Salt marsh
has significant contact with marine waters
(Figure 4a).
2.) Moderate tidal restriction. Moderate
contact with marine waters, though
configuration (channels not notably wide
or deep, not open to embayment, some
drainage creeks and ditches) or man-made
restrictions may present some obstacle to
flushing (Figure 4b).
3.) Severe tidal restriction. Little contact with
tidal waters as a result of man-made
restrictions. Noticeable changes in
topography and vegetative structure
(Figure 4c)
23
-------
Wildlife Habitat Value of New England Salt Marshes
Figure 3. Example of a New England salt marshes.
a) Little or no ditching. Most un-ditched marshes are characterized by an intact and natural
system of wide and narrow creeks and a high density of marsh pools.
:iE^'$p?&f»
iM *^;«itea
b) Moderate ditching. Ditches are present and may be numerous, but natural creeks are still intact
and present. There is generally a moderate density of marsh pools.
24
-------
Wildlife Habitat Value of New England Salt Marshes
c) Severe ditching. Note the extensive regular pattern of man-made ditches with few or no natural
creeks. The marsh also has a low density of marsh pools.
Figure 4. Extent of tidal restriction in New England salt marshes.
a) No to low tidal restriction. The marsh has significant contact with marine waters.
-------
Wildlife Habitat Value of New England Salt Marshes
,..'«: ..• ",••.,• • * n-*.*
''". . •vf.'J'JS''
V.- .•**-'>
"'^ ';v-i,v
,.;V
b) Moderate tidal restriction. The marsh has moderate contact with marine waters, though
configuration or man-made restrictions may present some obstacle to flushing.
c) Severe tidal restriction. There is little contact with tidal waters as a result of man-made
restrictions.
26
-------
Wildlife Habitat Value of New England Salt Marshes
Use of this component in a wildlife habitat
assessment
The degree of ditching is primarily related to
the extent of surface water on the marsh
(Reinert et al. 1981, Adamovicz and Roman
2005). In general, salt marshes with lesser
degrees of ditching or extent of tidal restriction
would be expected to have greater habitat value.
This could be captured in a quantitative
assessment by weighting the categories with the
"little to no" categories having the greatest value
and "severe" categories the least value. While it
is difficult to directly relate the extent of tidal
restriction to habitat value, tidally restricted
marshes may offer fewer resources to wildlife
species (e.g., Raposa and Roman 2001). This
component could therefore be included in an
assessment in a manner similar to that of the
degree of ditching (i.e., the "little to no"
categories to "moderate" categories having the
greatest habitat value and "severe" categories the
least value).
V. Salt Marsh Vegetation
While vegetation has been proposed as the
most important component of wildlife habitat in
freshwater marshes (Golet and Larson 1974),
New England salt marshes contain fewer species
of plants, trees, and shrubs than freshwater
wetlands because of their harsh physical regimes
determined in part by salt water inundation,
high soil salinities, and nutrient limitation.
Tiner (1987) describes five life forms of New
England tidal marshes (including tidal fresh
marshes): aquatic plants, emergents, shrubs,
trees, and vines. However, not all life forms
may be present in estuarine and coastal salt
marshes. For example, aquatic plants include
three sub-forms (submergents, free-floating
plants, and plants with floating leaves), but only
submergents are regularly found in salt marshes.
Trees, while generally not capable of growing in
hyper-saline soils, may be found occasionally in
isolated patches within the salt marsh of
sufficient elevation to avoid regular tidal
inundation (wooded islands). Vines are limited
to one species: common dodder Cuscuta
gronovii that is only occasionally found in salt
marshes, usually parasitizing marsh elder Iva
frutescens. While vegetative life forms and sub-
forms still have important wildlife habitat value,
the lack of vegetative heterogeneity may
decrease the relative importance of this category
to the overall habitat value of a salt marsh.
Below we list five life forms and nine sub-
forms of vegetation found in salt marshes and
important to wildlife. Sub-form categories are
derived from Golet and Larson (1974). Latin
names are taken from Tiner (1987).
Aquatic plants
Found in permanently flooded pools or sub-
tidal waters. In salt marshes, consist of rooted
submerged plants.
Sub-form:
1. Rooted submergent
- Widgeon grass Ruppia maratima
- Eelgrass Zostera marina
- Pondweeds Potamogeton spp.
Emergents
Rooted, erect herbaceous plants that have all or
part of their growth above water, or that grow
in regularly flooded inter-tidal areas.
Sub-forms:
1. Robust emergents (Erect emergents up to 4 m tall)
- Common reed Phragmites australis
- Cattail Typha spp.
- Fireweed Erechtites hieracifolia
27
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Wildlife Habitat Value of New England Salt Marshes
2. Short meadow emergents (Sedge-like
emergents, less than 1.5 m tall)
- Sedges Sdrpusspp.; Carexspp.
- Spike-rush Eleocharis spp.
- Black grass Juncus gerardii
- Baltic rush Juncus balticus
3. Narrow-leaved emergents (Narrow-
leaved graminoids less than 2 m tall)
- Smooth cordgrass Spartina alterniflora
(tall form up to 2.5 m tall)
- Salt meadow grass Spartina patens
- Spike grass Distichlis spicata
- Switchgrass Panicum virgatum
- Red fescue Festuca rubra
- Goose grass Puccinellia maritime
4. Forbs (herbaceous plants other than grasses
having little or no woody material)
- Seaside goldenrod Solidago sempervirens
- Salt marsh asters Aster spp.
- Seaside plantain Plantago maritime
- Sea lavender Limonium nashii
- Sea milkwort Glaux maritime
- Rose mallow Habiscus moscheutos
- Marsh mallow Althaea officinalis
- Sea rocket Cakile edentula
- Sea blite Suaeda linearis
- Glasswort Salicornia spp.
- Marsh orach Atriplexpatula
- Silverweed Potentilla anserine
- Marsh pink Sabatia spp.
- Seaside gerardia Agalinis maritime
- Annual salt marsh fleabane
Pluchea purpurascens
- Seaside arrow grass
Triglochin maritimum
- Saltwort Salsola kali
- Marsh fleabane Pluchea odorata
Shrubs
Woody vegetation less than 7 m in height
usually with multiple stems.
Sub-forms:
1. Low compact shrubs (generally less
than 1.5 m tall, with dense foliage
- Marsh elder Iva frutescens
- Sea myrtle Baccharis halimifolia
- Sweet gale Myricagale
Trees
Woody plants 7 m or greater in height
having a single main stem.
Sub-forms:
1. Deciduous trees:
- Black willow salix nigra
- Alder Alnus spp.
- Red maple Acer rubrum
- Black gum Nyssa sylvatica
- Trembling aspen Populus tremuloides
- Black oak Quercus velutina
- Black cherry Prunus serotina
2. Coniferous trees:
- American white cedar
Chamaecyparis thyoides
- Pitch pine Pinus rigida
- Eastern red cedar
Juniperus virginiana Vines
Woody plants or herbaceous plants that intertwine
around stems of other plants.
Sub-form:
1. Vines
- Common dodder Cuscuta gronovii
Use of this component in a wildlife habitat
assessment
The presence of these salt marsh vegetation
forms and sub-forms may have most utility
when assessing habitat value for a particular
wildlife species for which specific vegetative
habitat requirements are known. In this case,
the optimal vegetation type for that species
would be given more weight in the overall
assessment. Alternatively, greater relative value
could be placed on the presence of a number of
vegetative life forms and sub-forms when
assessing habitat value for overall wildlife
species diversity.
28
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Wildlife Habitat Value of New England Salt Marshes
VI. Salt Marsh Vegetative Heterogeneity
New England salt marshes are typified by
regular zonation among bands of differing
species of emergent vegetation (Miller and Egler
1950, Neiring and Warren 1980). Much of the
vegetative heterogeneity in salt marshes arises
from the interspersion of different sub-forms of
emergent vegetation. However, to a lesser
degree emergent vegetation is interspersed with
other forms of vegetation, for example shrubs on
the marsh-upland edge, and with water, as at
the edge of tidal creeks and pools. In this sense,
vegetative heterogeneity in salt marshes can be
represented by the abundance and diversity of
vegetative edge habitats (Table 3). We define
salt marsh vegetative edge habitat as the
interface between two adjacent vegetative life
forms, or between a vegetative life form and a
marsh habitat type.
A currently held paradigm in conservation
biology is that wildlife species diversity
increases with increasing number of types of
edge habitat, inasmuch as increases in edge
habitat represent an increase in habitat
heterogeneity (Ries and Sisk 2004, Ries et al.
2004, Cramer and Willig 2005). Edge habitat
may also be beneficial to some species by
providing increased prey abundance and
diversity (Whaley and Minello 2002, Albrecht
2004, Horn et al. 2005). However, some studies
have shown that habitat edge may be
detrimental, for example to breeding birds by
exposing nests to predation and parasitism
(Batary and Baldi 2004, Wolf and Batzli 2004,
Fletcher 2005).
Several species of breeding birds, including
waterfowl, marsh wren, and clapper rail will
utilize the marsh/upland edge, possibly to take
advantage of increased foraging opportunities
(Gibbs et al. 1992, Eddleman and Conway 1998,
Drilling et al. 2002). Foraging species may use
marsh/water edge habitat, and this edge may
also be of value as protection from exposure for
wintering waterfowl. Tidal creek edge may be
important for sharp-tailed sparrows and clapper
rails (DeRagon 1988, Eddleman and Conway
1998).
While a majority of the vegetative
heterogeneity in salt marshes arises from the
interspersion of different sub-forms of emergent
vegetation, we have seen little evidence either
in the literature or anecdotally of use of this
edge by wildlife species. Emergent plants
species are often interspersed in New England
marshes, and when present in monotypic stands
the borders between species can be irregular and
indistinct. This along with the similar physical
structure of the plants in different emergent
zones may diminish habitat value. We therefore
omit emergent/emergent edge from considera-
tion, and propose three life form edges
(emergent/shrub, emergent/tree, and shrub/tree)
as possibly enhancing salt marsh wildlife habitat
value. The emergent/shrub and shrub/ tree
edges will typify the marsh/upland edge in New
England salt marshes, and may provide habitat
value for some avian species when present. We
also add two life form / habitat type edges
(emergent/open water, emergent/tidal flat) that
were identified as being important for foraging
birds (Table 3). Three categories of salt marsh
vegetative heterogeneity are derived from the
presence of these 5 types of habitat edge (Figure
5):
1.) High heterogeneity: 5 habitat edges
present (Figure 5a)
2.) Moderate heterogeneity: 3 or 4 habitat
edges present (Figure 5b)
29
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Wildlife Habitat Value of New England Salt Marshes
3.) Low heterogeneity: 1 or 2 habitat edges
present (Figure 5c)
Use of this component in a wildlife habitat
assessment
New England salt marshes with greater
vegetative heterogeneity would be expected to
have greater wildlife habitat value. In an
assessment of habitat value, this could be
reflected in a weighting of the categories with
high heterogeneity having the greatest value
and low heterogeneity the least value. However,
we caution when assessing habitat value for a
single species or guild of species, habitat
heterogeneity may not be as important as the
presence of one or more favorable habitat types
for the species of concern.
VII. Surrounding Land Cover and Land Use
The importance of surrounding habitat type
to wetland wildlife value has been hypo-
thesized for many years. Early work
demonstrated the importance of adjacent
natural habitat for a number of species that
Figure 5. Occurrence of varying degrees of vegetative heterogeneity in New England salt marshes
a.
Open water
Tidal flats
Emergents
Wooded islands
Shrubs
Trees
c.
a) High habitat heterogeneity: 5 habitat edges present, b) moderate habitat heterogeneity: 3 or 4
habitat edges present; c) low habitat heterogeneity: 1 or 2 habitat edges present.
30
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Wildlife Habitat Value of New England Salt Marshes
prefer upland foraging and nesting sites. For
example, waterfowl often depend on the
presence of suitable upland habitat adjacent to
wetlands for nest sites and for roosting
(Baldassarre and Bolen 1994). Great blue herons
and great egrets will preferentially use large
canopy hardwood trees adjacent to salt marsh
foraging habitat for roosting and occasionally
nesting (Hancock and Kushlan 1984, Butler
1992). Several species, including glossy ibis
Plegadis falcinellus and red-winged blackbirds
will preferentially use salt marshes that are
adjacent to agricultural land because of
increased availability of food (Davis and Kricher
2000, Trocki 2003).
Recent studies in the landscape ecology of
wetlands have demonstrated the importance of
the complexity and degree of disturbance of
surrounding habitat (Freemark et al. 1995,
Riffell et al. 2003). The negative effects of
urbanization and alteration of adjacent uplands
on wildlife has been demonstrated for both
inland and coastal marshes (DeLuca et al. 2004,
Shriver et al. 2004, Traut and Hosteller 2004).
In urban settings, natural lands bordering salt
marshes may have a buffering effect and may be
important in mitigating the effects of human
disturbance.
Information about the proportion of land-use
types in a buffer around a salt marsh can be used
to classify the landscape setting of the marsh.
The size of the buffer will depend both on the
scale of the intended assessment (e.g., regional
comparisons over large geographic areas versus
local studies) and the species under
consideration. For a study at the scale of a
typical bay or estuary, we suggest quantifying
the proportion of land-use types in a 150 m
buffer around the marsh (Carslisle et a/2004,
McKinney et al. 2006). We propose nine land
use types aggregated into 4 broad categories for
the assessment. These land use types include
generally accepted land cover categories that
have been identified by or included in previous
classifications (e.g., Anderson 1976). In
assessing the value of landscape setting, we
recognize that 1) salt marshes bordered by
forested, open or other wetlands are more
valuable to wildlife; 2) depending on the species,
agricultural or certain maintained open lands
may be of wildlife habitat value; and 3) salt
marshes bordered by developed lands will be
less valuable as wildlife habitat.
Northern Water snake Nerodra sipedon sipedon
(Photo by Gary Stoltz, US FWS).
The nine land-use types are:
Open water
Land-use type:
1)Water: marine sub-tidal habitat
Natural land
Land-use types:
2) Forest: deciduous forest, coniferous forest,
brushland
3) Wetland
4) Barren land: beaches, sandy areas, rock
outcrops
31
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Wildlife Habitat Value of New England Salt Marshes
Maintained open land
Land-use types:
5) Urban or built-up land: power lines
developed recreation, cemeteries, vacant land
6) Agricultural land: row crops, pasture,
orchards, cranberry bogs, confined feeding
operations, idle agriculture
7) Maintained open land: strip mines,
quarries, gravel pits, power lines
Developed land
Land-use types:
8) Disturbed open land: commercial and
industrial land, airports, rail line, roads and
highways, railroads, freight, storage,
stadiums, water and sewage treatment,
waste disposal facilities, marinas
9) Residential land: single or multi-family
homes, areas of high population density
characterized by multi-dwelling apartment
buildings
Use of this component in a wildlife habitat
assessment
In an assessment of wildlife habitat value,
landscape setting, or an assessment of
surrounding land use, could influence salt marsh
habitat quality with urbanization and human
alteration of adjacent uplands thought to have a
negative effect and surrounding natural lands a
mitigating or positive influence on habitat
quality. This could be reflected in an assessment
by calculating the proportion of developed
versus natural lands and open water and
assigning a rank or score to a marsh accordingly,
with for example marshes with a higher
proportion of natural land being ranked above
those with a greater proportion of developed
land. The "maintained open lands" category
would be assessed relative to the species under
consideration, but in general this category
would be expected to have a relative value
between that of developed and natural lands.
VIII. Connectivity and Associated Habitat
During the past decade, wildlife-habitat
studies have begun to encompass larger spatial
and temporal scales (Edwards et al. 1994,
Morrisey 1996). Ecologists continue to
formalize the importance of both landscape
structure (the patterns of habitat density,
distribution, shape and size) and landscape
connectivity, or the functional relationship
between adjacent habitats arising from their
spatial distribution and the movement of
organisms (With et al. 1997). This emphasis and
resulting studies serve to reinforce the long-held
hypothesis that a wetland's value as wildlife
habitat is greater if it is located near other
wetlands, and that its value increases with the
degree of connectivity to and complexity of
associated wetlands. There are many examples
of connectivity and the availability of associated
natural habitats enhancing a wetland's habitat
value, particularly for avian species. Specific
examples for salt marsh fauna include use of
adjacent foraging areas away from nest sites
(Ramo and Busto 1993, Bryan et al. 1995, Smith
1995), post-breeding movements (Rotella and
Ratti 1992, Mauser et al. 1994), and movements
within migration and winter sites (Goss-Custard
and Durell 1990, Rehfisch et al. 1996, Farmer
and Parent 1997).
The following categories of associated habitat
are of potential value to salt marsh wildlife
(Table 3):
1) Sand or cobble beach
2) Coastal dunes or overwash
3) Other salt marsh wetland
4) Brackish wetland or pond
5) Freshwater wetland or pond
6) Upland meadow
7) Upland forest
32
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Wildlife Habitat Value of New England Salt Marshes
The presence of these habitat types in close
proximity (e.g., within a 150 m buffer) to a salt
marsh will enhance connectivity and facilitate
movements between salt marsh and associated
habitats (Haig etaL 1998).
Use of this component in a wildlife habitat
assessment
As with landscape setting of a marsh, the
presence of associated habitat types could
influence salt marsh habitat quality. In a
general sense, the presence of associated habitats
(i.e., greater landscape heterogeneity) is thought
to have a positive influence on habitat quality
and hence would increase wildlife habitat value.
To include salt marsh habitat type in an
assessment of wildlife habitat value of New
England salt marshes, one could assign a relative
value to the presence of each associated habitat,
or assign a value to a marsh based on the
number of associated habitats. How these
components are ranked or scored could depend
on the goal of the assessment and the specific
habitat requirements of the species under
consideration. Alternatively, if the goal is to
assess salt marsh habitat for maximum species
diversity, the presence of many associated
habitats within a 150 m buffer surrounding the
marsh would be emphasized.
Conclusions
This report provides a summary of wildlife
(i.e., birds, mammals, amphibians, and reptiles)
found in New England salt marshes and some of
their respective habitat requirements. The
wetland and landscape components in the report
describe some aspects coastal wetlands and their
associated habitats, and form the basis of a
framework to assess wildlife habitat value of
New England salt marshes.
An assessment of salt marsh wildlife habitat
function will require data on the extent of the
various components listed in this report. While
much of this data can be gleaned from the
analysis of remote sensing data such as aerial
photos, some level of field work will be required
to determine the occurrence of salt marsh
Great blue heron Aldea herodias
(Photo by Lee Karney, US FWS).
habitat types and the extent of vegetative
heterogeneity. Alternatively, this data can come
from existing salt marsh assessment protocols
(e.g., Carlisle et al. 2004) that have a field
component.
In any assessment, the actual weighting of
the various components and a component's
relative contribution will depend upon the
species and habitat under consideration and the
stakeholder intent. For example, distinct
requirements of species under consideration
should be reflected in the assessment by
emphasizing the wetland and landscape
components that encompass those requirements.
Special weighting for rare species or those of
local, regional, or national interest, and rare
habitats (those that are not commonly found in
a region) should also be considered. Once
completed, a salt marsh wildlife habitat
33
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Wildlife Habitat Value of New England Salt Marshes
assessment could be used as a guide for
protective, restorative, and mitigation efforts for
New England salt marshes.
The overall value of a wetland is dependent
not only upon wildlife use and support but also
on the provision of many other ecosystem
services (e.g., water quality maintenance,
erosion control and flood abatement, recreation
and aesthetics). Other socioeconomic and
ecological factors that are not covered in this
report may also be important and enhance
ecosystem services provided by New England
salt marshes. In addition to wildlife habitat
value, consideration of special or needed
services (e.g., educational or recreational
resources; water quality maintenance; flood
abatement) will be an important part of
developing an overall salt marsh evaluation
model.
34
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Wildlife Habitat Value of New England Salt Marshes
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46
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Wildlife Habitat Value of New England Salt Marshes
Appendix 1. Habitat use, occurrence, and primary prey of birds
Common Name Species
that utilize New England salt marshes.
Habitat Type1 Occurrence2 Prey3
Reference4
Breeders
American oystercatcher
Clapper rail
Common tern
Killdeer
Laughing gull
Least bittern
Mallard
Marsh wren
Mute swan
Red-winged blackbird
Salt marsh sharp-tailed sparrow
Seaside sparrow
Swamp sparrow
Virginia rail
Willet Catoptrophoru
Foragers
American black duck
American coot
American crow
American goldfinch
American robin
American wigeon
Bald eagle
Bank swallow
Barn swallow
Belted kingfisher
Black-bellied plover
Black-crowned night heron
Blue-winged teal
Haematopus palliates
Rallus longirostris
Sterna hirundo
Charadrius vociferous
Lams atricilla
Ixobrychus exilis
Anas platyrhynchos
Cistothorus palustris
Cygnus olor
Agelaius phoeniceus
Ammodramus caudacutus
Ammodramus maritimus
Melospiza georgiana
Rallus limicola
semipalmatus
Anas rubripes
Fulica Americana
Corvus brachyrhynchos
Carduelis tristis
Turdus migratorius
Anas americanus
Haliaeetus leucocephalus
Riparia riparia
Hirundo rustica
ceryle alcyon
Pluvialis squatarola
Nycticorax nycticorax
Anas discors
TF, HM, MB O, S
TF, LM, HM O, S
SW, TF, HM, MB O, S
TF, LM, PL, PN O, S
SW, TF, HM, MB O, S
SW, TF, HM, MB, PH O, S
SW, LM, HM, PL F, Y
TF, HM, MB, PH O, S
SW, LM, HM, MB, PH F, Y
LM, HM, WI, MB, PH F, S
LM, HM F, S
LM, HM O, S
LM, HM, MB O, S
TF, LM, HM O, S
TF, LM, HM, PL, PN O, S
SW, LM, PL F, W
SW O,W
LM, HM O, Y
HM, MB, PH O, S
HM, MB O, Y
SW, LM O, W
LM, HM, MB O, Y
LM, HM, MB, PH O, S
LM, HM, MB, PH O, S
SW, LM, HM, MB O, Y
TF, LM O, Y
SW, TF, LM, HM, PL, WI F, S
SW, TF, LM O, W
Invertebrates (seeds)
Crustaceans (fish)
Fish (crustaceans)
Invertebrates (seeds)
Invertebrates (fish)
Fish (insects)
Vegetation
Invertebrates
Vegetation
Insects
Insects (seeds)
Insects (seeds)
Seeds (invertebrates)
Invertebrates (seeds)
Crustaceans (insects)
Vegetation
Vegetation (seeds)
Invertebrates (seeds)
Seeds (insects)
Invertebrates (seeds)
Vegetation (seeds)
Fish (birds, mammals)
Insects
Insects
Fish (invertebrates)
Invertebrates (bivalves)
Fish (crustaceans)
Invertebrates (seeds)
AT,AU
T
BF,BG
U
BCXBP^BR
V
W,X,Y,Z,AA
AB
AC,AD,AE
AF
AG,AH
AG,AI
AJ,AK
AL,AM
AN,AO
R, S
AP,AQ
AR
AS
AV
DD
AW
CF
CG
AX,AY,AZ
BA,BB
C,D,K,N
DE
47
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Wildlife Habitat Value of New England Salt Marshes
Appendix 1. Habitat use, occurrence, and primary prey of birds that utilize New England salt marshes (Cont'd).
Common Name
Species
Habitat Type1
Occurrence2
Prey3
Reference4
Bonaparte's gull
Brant
Canada goose
Cattle egret
Cedar waxwing
Chimney swift
Common grackle
Common yellowthroat
Double-crested cormorant
Dunlin
Eastern kingbird
European starling
Fish crow
Glossy ibis
Gray catbird
Great black-backed gull
Great blue heron
Great egret
Great horned owl
Greater yellowlegs
Green heron
Green-winged teal
Herring gull
House sparrow
Least sandpiper
Least tern
Lesser yellowlegs
Little blue heron
Mourning dove
Northern cardinal
Northern flicker
Northern harrier
Lams philadelphi
Branta bernicla
Branta canadensis
Bubulcus ibis
Bombycilla cedrorum
Chaetura pelagica
Quiscalus quiscula
Geothylpis trichas
Phalacrocorax auritus
Calidris alpina
Tyrannus tyrannus
Sturnus vulgaris
Corvus ossifragus
Plegadis falcinellus
Dumetella carolinensis
Lams marinus
Aldea herodias
Egretta alba
Bubo virginianus
Tringa melanoleuca
Butorides virescens
Anas crecca
Lams argentatus
Passer domesticus
Calidris minutilla
Sterna antillarum
Tringa spp.
Egretta caerulea
Zenaida macroura
Cardinalis cardinalis
Colaptes auratus
Circus cyaneus
SW,TF
SW, TF, LM
SW, LM, HM
SW, TF, LM, HM, PL
MB
HM, MB
LM, HM, MB, PH
MB
SW
SW,TF
HM, MB
LM, HM, MB, PH
LM, HM
SW, HM, PL
MB
SW, TF
SW, TF, LM, WI
SW, TF, LM, HM, PL
LM, HM, MB
SW, TF, LM, PL
SW, TR, PL
SW, TF, LM
SW, TF
LM, HM, MB, PH
TF, LM
SW,TF
SW, TF, LM, PL
SW, TF, PL
LM, HM, MB
HM, MB, PH
MB
LM, HM, MB
O,Y
O,W
O,Y
O, M
O,Y
0,S
O,Y
O, S
0,Y
O,W
0,S
O,Y
0,Y
F, S
O,Y
0,Y
F, S
F,S
O,Y
F,S
F, S
O,W
0,Y
O,Y
O, M
O, S
F,S
O, S
0,Y
O,Y
O,Y
O,W
Fish (invertebrates)
Vegetation
Vegetation
Fish (invertebrates)
Fruit (insects)
Insects
Insects (seeds)
Insects
Fish
Invertebrates
Insects (fruit)
Invertebrates (insects)
Invertebrates (seeds)
Invertebrates
Insects (fruit)
Fish (invertebrates)
Fish
Fish (crustaceans)
Mammals (birds)
Invertebrates (Small fish)
Fish
Invertebrates (seeds)
Fish (invertebrates)
Fish (invertebrates)
Invertebrates
Fish (invertebrates)
Invertebrates (Small fish)
Fish (crustaceans)
Seeds
Seeds (insects)
Insects (seeds)
Mammals (birds)
BC
DJ
DG
BD
CH
CI
BE
BH
BI
CL,CM,CN
CJ
BJ
BK
E
BL
BM
A,B,G,K,N,Q
B,C,G,K,L,Q
CT
H,P
F,N
DF
BN
BS
CO,CP
BT
H,P
G,I,K,N,O,Q.
BU
BV
BW
CU,CV
48
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Wildlife Habitat Value of New England Salt Marshes
Northern mockingbird
Northern pintail
Osprey
Red-shouldered hawk
Red-tailed hawk
Rough-legged hawk
Ring-necked duck
Ring-necked pheasant
Sanderling
Semipalmated plover
Semipalmated sandpiper
Short-eared owl
Snowy egret
Snowy owl
Song sparrow
Sora
Spotted sandpiper
Tree swallow
Yellow-crowned night heron
Mimus polyglottos
Anas acuta
Pandion haliateus
Buteo lineatus
Buteo jamaicensis
Buteo laopus
Anas collaris
Phasianus colchicus
Calidris alba
Calidris semipalmatus
Calidris pusilla
Asio Hammeus
Egretta thula
Nyctea scandiaca
Melospiza melodia
Porzana Carolina
Actitus macularia
Tachycineta bicolor
Nyctanassa violacea
HM, PN, MB O, Y
SW, LM O, W
SW O, S
HM, MB O, Y
HM, MB O, Y
HM, MB O, Y
SW O,W
HM, MB O, Y
SW, TF O, W
TF O, M
TF, LM O, M
LM, HM, MB O, Y
SW, TF, LM, HM, PL F, S
LM, HM, MB O, W
LM, HM, MB O, Y
LM, HM O, M
SW, TF O, S
LM, HM, MB, PH O, S
SW, TF, LM, HM, PL, WI F, S
Insects (seeds)
Vegetation (Invertebrates)
Fish
Mammals (birds)
Mammals (birds)
Mammals (birds)
Seeds (invertebrates)
Seeds (vegetation)
Invertebrates (bivalves)
Invertebrates
Invertebrates
Mammals (birds)
Fish (crustaceans)
Mammals (birds)
Seeds (insects)
Seeds (invertebrates)
Invertebrates (fish)
Insects
Crustaceans
BX,BY
DH
CW
CX
CY
CZ,DA
DI
CA
CR
CB
CQ.
DB
B,C,G,K,M,Q
DC
CC
CS
CD
CK
C,K,N,R,CE
r = shallow water; TF = tidal flats; LM = low marsh; TR = trees overhanging water;
WI = wooded islands; MB = marsh-upland border; PH = phragmites.
2F = frequent; O = occasional; S = summer (breeding); W = winter (non-breeding); M =
3Primary prey (secondary prey in parentheses).
4References
A) Butler 1992.
B) Chavez-Ramirez and Slack 1995.
C) Custer and Osborn 1978.
D) Davis 1993.
E) Davis and Kricher 2000.
F) Davis and Kushlan 1994.
G) DuBowyl996.
H) Elphik and Tibbitts 1998.
I) Erwin etal. 1994.
J) Gibbs etal. 1992.
K) Hancock and Kushlan 1984.
L) McCrimmon etal. 2001.
M) Parsons and Master 2000.
N) Ramo and Busto 1993
O) Rodgers and Smith 1995.
P) Tibbitts and Moskoff 1999 .
Q) Willard 1977.
R) Longcore etal. 2000.
S) Morton etal. 1989.
T) Eddleman and Conway 1998.
HM = high marsh; PL = marsh pools; PN = pannes;
= fall/spring migration; Y = year-round
U) Jackson and Jackson 2000.
V) Gibbs etal. 1992.
W) Hill 1984.
X) Allen 1986.
Y) Mauser etal. 1991.
AA) Drilling etal. 2002.
AB) Kroodsma and Verner 1997.
AC) Ciaranca etal. 1997.
49
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Wildlife Habitat Value of New England Salt Marshes
AD) Holm 2002.
AE) Rees era/. 1997.
AF) Yasukawa and Searcy 1995.
AG) DeRagonl988.
AH) Greenlaw and Rising 1994.
AI) Marshall and Reinert 1990.
AJ) Reinert and Golet 1979.
AK) Mowbray 1997.
AL) Taylor 1975.
AM) Conwayl995.
AN) Douglas 1996.
AO) Lowther era/. 2001.
AP) Desrochers and Ankney 1986.
AQ) Brisbin era/. 2002.
AR) Verbeek and Caffrey 2002.
AS) Middleton 1993.
AT) Lauro and Burger 1989.
AU) Nol and Humphrey 1994.
AV) Sallabanks and James 1999.
AW) Buehler2000.
AX) Meyerreicks and Nellis 1967.
AY) Prose 1985.
AZ) Hamas 1994.
BA) Townshend era/. 1984.
BB) Paulson 1995.
BC) Burger and Gochfeld 2002.
BD) Telfair2006.
BE) Peer and Bellinger 1997.
BF) Andrews 1990.
BG) Nisbet2002.
BH) Guzy and Ritchison 1999.
BI) Hatch and Weseloh 1999.
BJ) Cabe 1993.
BK) McGowan2001.
BL) Cimprich and Moore 1995.
BM) Good 1998.
BN) Pierotti and Good 1994.
BO) Bongiorno 1970.
BP) Burger 1977.
BQ) Burger era/. 1978.
BR) Burger 1996.
BS) Lowther 2006.
BT) Thompson era/. 1997.
BU) Mirarchi and Baskett 1994.
BV) Halkin and Linville 1999.
BW) Moore 1995.
BX) Roth 1979.
BY) Derrickson and Breitwisch 1992.
BZ) Ryder 1993.
CA) Giudice and Ratti 2001.
CB) Nol and Blanken 1999.
CC) Arcese era/. 2002.
CD) Oring era/. 1997.
CE) Watts 1995.
CF) Garrison 1999.
CG) Brown and Brown 1999.
CH) Witmer era/. 1997.
CI) Cink and Collins 2002.
CJ) Murphy 1996.
CK) Robertson era/. 1992.
CL) Goss-Custard and Moser 1988.
CM) Mouritsen and Jensen 1992.
CM) Warnock and Gill 1996.
CO) Hicklinl987.
CP) Cooper 1994.
CQ) Gratto-Trevor 1992.
CR) MacWhirter 2002.
CS) Melvin and Gibbs 1996.
CT) Houston era/. 1998.
CU) Collopy and Bildtein 1987.
CV) MacWhirter and Bildstein 1996.
CW) Poole era/. 2002.
CX) Crocolll994.
CY) Preston and Beane 1993.
CZ) Bosakowski and Smith 1992.
DA) Bechard and Swem 2002.
DB) Wiggins 2006.
DC) Parmelee 1992.
DD) Mowbray 1999.
DE) Rowher era/. 2002.
DF) Johnson 1995.
DG) Mowbray era/. 2002.
DH) Austin and Miller 1995.
DI) Hohman and Eberhardt 1998.
DJ) Reed era/. 1998.
50
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Wildlife Habitat Value of New England Salt Marshes
Appendix 2. Habitat use, home range, and primary prey of mammals, amphibians, and reptiles that utilize New England salt marshes.
Common Name
Species
Habitat Type1
Home Range
Prey2
Reference3
Black-tailed jackrabbit
Eastern cottontail
Least shrew
Masked shrew
Raccoon
Virginia opossum
White-tailed deer
Coyote
Fisher
Long-tailed weasel
Mink
Red fox
River otter
Striped skunk
Meadow jumping mouse
Meadow vole
Muskrat
New England cottontail
Norway rat
Woodland vole
Common snapping turtle
Diamondback terrapin
Green frog
Northern water snake
Painted turtle
Spotted turtle
Lepus californicus
Sylvilagus florianus
Cryptotis parva
Sorex cinereus
Procyon lotor
Didelphis virginiana
Odocoileus virginianus
Canis latrans
Mattes pennant!
Mustela frenata
Mustela vison
Vulpes wipes
Lontra canadensis
Mephitis mephitis
Zapus hudsonius
Microtus pennsylvanicus
Ondatra zibethicus
Sylvilagus transitionalis
Rattus norvegicus
Microtus pinetorum
Chelydra s. serpentine
Malaclemys t. terrapin
Rana clamitans melanota
Nerodra s. sipedon
Chrysemys picta
Clemmys guttata
HM, WI, MU
HM, MU
HM, MU
HM, MU
LM, HM, PL, MU
HM, MU
HM, WI, MU, PG
LM, HM, WI, MU, PG
SW, TF, LM, HM
LM, HM, WI, MU
SW, TF, LM, HM
LM, HM, WI, MU, PG
SW, TF, LM
HM, MU
LM, HM, PL, MU, PG
HM, MU
SW, TF, LM
HM, MU
HM, MU
HM, MU
TF, LM
LM, HM
SW, TF
SW, TF, LM, MU, PG
HM, WI, MU
SW, HM, MU
20 -140 ha Forbs, succulents B,G
0.9 - 2.8 ha Forbs (grasses) F
170 - 280 ha Insects (crustaceans) AE
Insects
49 ha Invertebrates Q,Z
4.65 - 23.5 ha Insects (carrion) I,P,S
59 - 520 ha Grasses (forbs) R,X
1000 - 4900 ha Small mammals (crustaceans) A
900-1300 ha Small mammals (birds) T
16 - 160 ha Small mammals (birds) W
600 - 5600 ha Fish (small mammals) J,L
1450 - 2000 ha Birds (fish) K,M
Fish (crustaceans) N,O
200 ha Insects (small mammals) AB,AC
0.1 - 0.4 ha Forbs (insects) AD
0.01 - 0.4 ha Grasses (forbs) U,V
50 - 200 ha Aquatic plants (fish) AF
50 - 200 ha Aquatic plants (fish) E,H
7.8 ha Forbs (small mammals) C
1.1 ha Grasses (forbs) D,Y
Insects (crustaceans) AG
Crustaceans (insects) AH
0.01 ha Insects (crustaceans) AI
Amphibians (fish) AH
Insects (crustaceans) AG
Gastropods (insects) AG
1 SW = shallow open water; TF = tidal flats; LM = low marsh; TR = trees overhanging water; HM = high marsh; PL = marsh pools; PA = pannes;
WI =wooded islands; MU = marsh-upland border; PG = phragmites.
2 Primary prey (secondary prey in parentheses).
51
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Wildlife Habitat Value of New England Salt Marshes
3 References
A) Bekoff 1977.
B) Best 1996.
C) Calhoun 1962.
D) Cengel era/. 1978.
E) Chapman 1975.
F) Chapman era/. 1980.
G) Currie and Goodwin 1966.
H) Dalke 1937.
I) Fitch and Shirer 1970.
J) Gerrell 1970.
K) Harrison era/.1989.
L) Lariviere 1999.
M) Lariviere and Pasitschniak-Arts 1996.
N) Lariviere and Walton 1998.
O) Larsen 1984.
P) Lay 1942.
Q) Lotze and Anderson 1979.
R) McCullough 1984.
S) McManus 1974.
T) Powell 1984.
U) Reich 1996.
V) Riewe 1973.
W) Sheffield and Thomas 1997.
X) Smith 1991.
Y) Smolen 1981.
Z) Urban 1970.
AA) Van Vleck 1969.
AB) Verts 1967.
AC) Wade-Smith and Verts 1982.
AD) Whitaker 1972.
AE) Whitaker 1974.
AF)Willner era/. 1980.
AG) Babcock 1971.
AH) DeGraaf and Yamesaki 2001.
AI) Jensen 1967.
52
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