HomeMy WebLinkAboutUSFWS 1984 Habitat Suitability Model Mercenaria mercenariaFWS/OBS-82/10,77
AUGUST 1984
HABITAT SUITABILITY INDEX MODELS:
HARD CLAM
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19970318 096
Fish and Wildlife Service
U.S. Department of the Interior
DTIC QUALITY IKBP; RPWGTii
This model is designed to be used by the Division of Ecological Services in
conjunction with the Habitat Evaluation Procedures.
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U.S. Fish and Wildlife Service
1010 Gause Boulevard
SI i dell, LA 70458
Thank you for your assistance.
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FWS/OBS-82/10.77
August 1984
HABITAT SUITABILITY INDEX MODELS:
HARD CLAM
by
Rosemarie Mulholland
Florida Cooperative Fish and Wildlife Research Unit
U.S. Fish and Wildlife Service
School of Forest Resources and Conservation
117 Newins-Ziegler
University of Florida
Gainesville, FL 32611
Project Officer
Michael S. Brody
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
1010 Gause Boulevard
SIidell, LA 70458
Performed for
National Coastal Ecosystems Team
Division of Biological Services
Research and Development
Fish and Wildlife Service
U.S. Department of the Interior
Washington, DC 20240
This report should be cited as:
Mulholland, R. 1984. Habitat suitability index models: hard clam. U.S.
Fish Wild!. Serv. FWS/OBS-82/10.77. 21 pp.
PREFACE
The habitat suitability index (HSI) model in this report on the hard
clam is intended for use in the habitat evaluation procedures (HEP) developed
by the U.S. Fish and Wildlife Service (1980) for impact assessment and
habitat management. The model was developed from a review and synthesis of
existing information and is scaled to produce an index of habitat suitability
between 0 (unsuitable habitat) and 1 (optimally suitable habitat).
Assumptions involved in developing the HSI model and guidelines for model
applications, including methods for measuring model variables, are described.
This model is a hypothesis of species-habitat relationships, not a
statement of proven cause and effect. The model has not been field-tested.
For this reason, the U.S. Fish and Wildlife Service encourages model users to
convey comments and suggestions that may help increase the utility and
effectiveness of this habitat-based approach to fish and wildlife management.
Please send any comments and suggestions you may have on the HSI model to the
following address.
National Coastal Ecosystems Team
U.S. Fish and Wildlife Service
1010 Gause Boulevard
Slide!!, LA 70458
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CONTENTS
Page
PREFACE in
ACKNOWLEDGMENTS vi
INTRODUCTION 1
Distribution 1
Life History Overview 1
SPECIFIC HABITAT REQUIREMENTS 3
Embryo, Larva, Juvenile 3
Adult 4
HABITAT SUITABILITY INDEX (HSI) MODEL 7
Model Applicability 7
Model Description 7
Suitability Index (SI) Graphs for Model Variables .... 9
Component Index (CI) Equation and HSI Determination ... 13
Interpreting Model Outputs 13
Field Use of Model 14
LITERATURE CITED 16
ACKNOWLEDGMENTS
Development of the habitat suitability index model for the hard clam-was
reviewed and constructively criticized by Dr. R. Winston Menzel, Oceanography
Department, Florida State University, Tallahassee; and Dr. John J. Manzi,
Marine Resources Research Institute, South Carolina Wildlife and Marine
Resources Department, Charleston. Thorough evaluations of model structure
and functional relationships were provided by personnel of the U.S. Fish and
Wildlife Service's (FWS) National Coastal Ecosystems Team (NCET). Michael S.
Brody served as project officer at NCET. Supportive narrative and model
reviews also were provided by Regional personnel of the U.S. Fish and
Wildlife Service and the Florida Department of Natural Resources. Funding
for model development and publication was provided by the FWS. The cover
illustration was prepared by David S. Maehr, Florida Game and Fresh Water
Fish Commission.
vi
HARD CLAM (Mercenaria campechiensis, Mercenaria mercenaria)
INTRODUCTION
Two species of hard clams occur along the Atlantic and Gulf of Mexico
coasts of North America: the southern hard clam, Mercenaria campechiensis
Gmelin 1791, and the northern hard clam, Mercenaria mercenaria Linne 1758
(Wells 1957b). The latter species, also commonly known as the quahog, was
formerly named Venus mercenaria. The two species are closely related,
produce viable hybrids (Menzel and Menzel 1965), and may be a single species.
Throughout their ranges, hard clams support extensive commercial and
recreational fisheries (Ritchie 1977). Additionally, because adult clams are
sedentary, they may serve as biological indicators of changing environmental
conditions.
Distribution
Mercenaria mercenaria is found in intertidal and subtidal areas along
the Atlantic and gulf coasts. Its range extends from the Gulf of St.
Lawrence to Texas (Belding 1912; Johnson 1934; Abbott 1954, 1974), but it is
most abundant from Massachusetts to Virginia (Stanley and DeWitt 1983).
Mercenaria campechiensis ranges from New Jersey (Merrill and Ropes 1967) to
St. Lucie Inlet, Florida (Godcharles and Jaap 1973) along the Atlantic coast,
and from Florida to Texas in the Gulf of Mexico (Ladd 1951).
Life History Overview
Spawning. Hard clams spawn from spring to fall depending on latitude
and temperature (Stanley and DeWitt 1983). In northwest Florida, the
spawning period extends over at least 6 months and has a peak in the spring
and a smaller peak in the fall (R. W. Menzel, Florida State University,
Tallahassee; pers. comm.). In more northern latitudes, clams spawn from May
to August (Stanley and DeWitt 1983). Not all clams in a given population
spawn at the same time (Loosanoff 1937); rather, spawning is spread over a
period of 8 to 10 weeks (Ansel 1 1967a). Water temperature is the determining
factor in final maturation of the gametes. When stimulated by the
appropriate temperature, males release semen containing pheromones._ Water
currents transport the pheromones to the females, which are in turn
stimulated to release eggs (Nelson and Haskin 1949). Fertilization occurs in
the water column. Carriker (1961) found that spawning occurred more
frequently during neap than spring tides and suggested that the higher
temperatures during neap tides triggered spawning. Bayne (1976) reported
that M. mercenaria usually spawned at or just after low tide, which was
coincident with maximum daily water temperature.
For M. mercenaria in Great South Bay, New York, Bricelj and Malouf
(1980) noted a significant correlation between size (length) and egg
production; 15% to 25% of the variation in fecundity was attributable to
differences in clam size. The estimated maximum production for one female
during a single spawning season was 16.8 million eggs. The sex ratio was
about 1:1. There was no evidence of a decline in egg production with
increasing age.
Egg. When first discharged by the female, the planktonic eggs of M.
mercenaria appear grayish and granular (Carriker 1961). Kraeuter et al.
(1982) found that survival was higher among large eggs (44 ym) than among
small ones (25 ym). Eggs of hard clams differ from those of some other
lamellibranchs in being surrounded by a thick gelatinous membrane (Loosanoff
and Davis 1950). The fertilized egg reaches the two-celled stage in about 45
minutes and the four-celled stage in about 90 minutes. At 22°C (72°F), the
trochophore stage is reached about 12 hours after fertilization (Loosanoff
and Davis 1950).
Larva. The pear-shaped trochophores actively propel themselves through
the water with a strongly ciliated velum (Loosanoff and Davis 1950). The
trochophore begins to form a primitive mouth and develop a shell gland. It
continues its planktonic existence after entering the shelled veliger stage,
which consists of two forms: straight-hinged and umboned. In the
straight-hinged phase a small thin shell secreted by the shell gland covers
the entire animal. The highly developed velum enables the larva to become a
proficient swimmer. The straight-hinged veliger phase lasts 1 to 3 days.
The veliger then enters the umboned veliger phase, characterized by a gently
sloping umbone projecting above the middle of the hinge line (Carriker 1961).
After 6 to 20 days the veliger reaches the pediveliger stage, in which it has
a foot and alternates between swimming in the water and crawling on the
bottom. This stage terminates when the velum is lost. The pediveliger then
enters the plantigrade benthic stages, and locomotion is limited to crawling
on the bottom. Initially the larva becomes a byssal plantigrade and affixes
itself to the substrate with a byssus. This is the setting or spattinq
stage. For a number of weeks, until the clam is about 9 mm (0.4 inches)
long, the byssal plantigrade alternates between byssal attachment and
crawling. The larva next enters the juvenile plantigrade stage, when the
byssus gland is no longer functional and the byssus is lost; the clam then
maintains its position beneath the sediment surface by means of its foot.
The siphons are fully developed at this stage (Carriker 1961). The juveniles
continue to grow and, as their siphons lengthen, they burrow deeper into the
sediment and complete their development (Carriker 1956). The adult clam
remains in much the same location for the rest of its life (Belding 1911).
Chestnut (1952) reported that adult clams moved up to 15 cm (6 inches)
laterally in 38 days, and Kerswill (1941) observed movements of up to 30 cm
(12 inches) in 2 months.
Juvenile and adult. W. mercenaria secrete shell material daily and grow
rapidly (Kennish and Loveland 1980). However, growth begins to slow at age 2
(Carriker 1961; Rhoads and Pannella 1970), when energy is diverted into
reproductive processes (Kennish and Loveland 1980).
Recruitment throughout the range of the two species of Mercenaria is
erratic and unpredictable (Hibbert 1976; Menzel 1976). It is probably
determined by the amount of predation occurring after the juveniles settle
(Menzel 1976). A major predator of small clams is the blue crab, Callinectes
sapidus (Haven and Andrews 1957; Menzel and Sims 1962; Castagna 1970a,1970b;
Castagna and Kraeuter 1977). Other major predators are whelks, Busycon
spp.; moon snails, Polinices duplicatus; and stone crabs, Menipe mercenaria
(Menzel et al. 1976).
SPECIFIC HABITAT REQUIREMENTS
Embryo, Larva, Juvenile
£H. Calabrese (1972) observed that the successful recruitment of M.
mercenaria requires that the pH of estuarine waters not fall below 7.0; he
found no significant decrease in the number of clam embryos developing
normally within the pH range of 7.0-8.75, but that number was greatly
reduced at pH 9.0. Survival of clam larvae was normal at pH 6.25-8.75, but
the range for normal growth was 6.75-8.50. Although clam larvae can survive
at pH 6.25, a pH of 7.0 is required for normal development of the embryo.
Levels of pH below 7.0 limit recruitment of the species (Calabrese 1972).
Dissolved oxygen. Morrison (1971) found that growth of shelled veligers
of M. mercenaria was normal when dissolved oxygen concentration was 4.2 mg/1
or greater. Growth essentially ceased at concentrations of 2.4 mg/1 and
less. Larvae survived extended exposures (14 days) to 1 mg/1 dissolved
oxygen but grew little. Prolonged exposure to levels of less than 4.0 mg/1
lengthened the clam's planktonic stage and decreased its probability of
survival. Embryos developed normally at oxygen levels as low as 0.5 mg/1;
however, 100% mortality occurred at 0.2 mg/1.
Salinity. Salinity appears to be most critical for M. mercenaria during
the egg and larval stages (Stanley and DeWitt 1983). At Long Island Sound,
New York, eggs developed into straight-hinged veligers only within the
relatively narrow salinity range of 20.0 to 32.5 parts per thousand (ppt).
The optimum for development of clam eggs was about 26.5 to 27.5 ppt (Davis
1958). Growth of larvae, once they attained the straight-hinged stage, was
comparatively good at salinities as low as 20 ppt (Davis 1958), but Chanley
(1958) found that growth of juvenile M. mercenaria was retarded at
salinities of 22.5 ppt or lower. Castagna and Chanley (1973) found that
metamorphosis of M. mercenaria from veliger to seed clam (byssal plantigrade
stage) was inhibited below 17.5 to 20 ppt.
Temperature. Davis and Calabrese (1964) noted that laboratory-reared
straight-hinged veligers of M. mercenaria were capable of ingestion, but not
digestion, at 10°C (50°F), and consequently did not grow. Growth was
positively related to temperature at 18.0° to 30.0°C (64° to 86°F). Growth
of straight-hinged veligers of M. mercenaria was little affected by
temperature differences within the range of 20° to 30°C (68° to 86°F).
Although the optimum temperature for growth of M. mercenaria larvae was not
well defined, growth was optimum at the following temperature/salinity
combinations: 30°C (86°F)/22.5 ppt and higher, 27.5°C (81.5°F)/17.5 and 20.0
ppt, and 25°C (77°F)/15.0 ppt. The larvae appeared to be more sensitive to
differences in salinity than to differences in temperature (Davis and
Calabrese 1964). Kennedy et al. (1974) observed that temperature tolerance
increased with age; cleavage stages were the most sensitive to high
temperatures and straight-hinged larvae the least sensitive.
Substrate. The nature of the bottom substrate seems to be the main
factor responsible for settling of larvae and for the qualitative composition
of bottom communities (Thorson 1955). Keck et al. (1974) reported from
laboratory studies that significantly higher (P < 0.05) numbers of M.
mercenaria larvae set in sand than in mud; they suggested that the addition
of organic material to the sediment may be responsible for reduced setting
because of increased bacteria levels, reduced dissolved oxygen, and increased
production of hydrogen sulfide. Carriker (1959) recommended that the
substrate be firm and free of excessive organic mud for larval clam culture;
muddy bottoms can be surfaced with shells, sand, or gravel.
Suspended solids. Suspended solids affect both the eggs and larvae of
hard clams. Davis (1960) found that clam eggs did not develop normally at
silt concentrations of 3.0 or 4.0 g/1. Growth of straight-hinged veligers
was normal at a silt concentration of 0.75 g/1, retarded at 1.0 to 2.0 g/1,
and negligible at 3.0 and 4.0 g/1.
Adult
pH. In mortality experiments with adult M. mercenaria, Calabrese (1972)
observed that the pH of tidal estuarine waters should not fall below 7.0 even
though larvae can survive at lower pH levels. The species could not
reproduce successfully in waters where pH remained appreciably above 9.0
since at pH 9.50 to 9.75 there was virtually no development of embryos.
Dissolved oxygen. Fluctuations in dissolved oxygen do not affect hard
clams as much as do fluctuations in temperature and salinity (Stanley and
DeWitt 1983). The burrowing ability of M. mercenaria was neither severely
nor permanently impaired by exposure to reduced oxygen levels (less than 1
mg/1 seawater) for up to 3 weeks (Savage 1976). Pratt and Campbell (1956)
found no correlation between growth rates and various concentrations of
dissolved oxygen. All life stages tolerate nearly anoxic conditions for long
periods, though they may cease growing (Stanley and DeWitt 1983). Greenfield
and Crenshaw (1981) reported that M. mercenaria has evolved several metabolic
responses to anoxia; the type of response depends on the length of exposure.
Salinity. The effects of salinity and temperature on clams are diffi-
cult to interpret because of the interaction between these two factors.
Woodburn (1961, 1962) reported that near oceanic salinity (35 to 36 ppt) was
best for M. campechiensis and recommended 20 ppt as a minimum level. The
experimentally determined nonlethal minimum salinity for adult M. mercenaria
was 12.5 ppt (Castagna and Chanley 1973). The range of salinities at a
northwest Florida site supporting both species was 26 to 35 ppt (Menzel
1961). Menzel and Sims (1962) recommend a salinity in excess of 25 ppt for
M. mercenaria plantings in Florida. Adult hard clams are capable of
withstanding low salinity by closing their shells. In South Carolina, M.
mercenaria mortality was less than 5% during 2- and 3-week periods when
salinity was less than 10 ppt (Burrell 1977). In the laboratory, Pearse
(1936) found that adult M. mercenaria could survive for 114 hours in
freshwater.
Temperature. The Joint Subcommittee on Aquaculture (1983) reported that
hard clam gonadal development begins at 8° to 10°C (46° to 50°F), spawning
occurs between 22° and 28°C (72° and 82°F), and that growth continues between
8° and 28°C (46° and 82°F). Kennish and Olsson (1975) reported 21° to 25°C
(70° to 77°F) as the preferred or required temperature range for spawning of
M. mercenaria in Barnegat Bay, New Jersey. Carriker (1961) reported spawning
at temperatures of 22° to 30°C (72° to 86°F) in New Jersey, and Mitchell
(1974) reported spawning at 18° to 20°C (64° to 68°F) in England. Menzel
(1976) reported that spawning peaks in Florida when water temperatures
approach 22° to 24°C (72° to 75°F). In northwest Florida, shell growth was
greatest in spring and fall. Menzel (1961, 1962) demonstrated that growth of
M. campechiensis was negligible during winter at temperatures approaching 10°
to 12 C (50° to 54°F). _M. campechiensis continued to grow until temperatures
approached 35°C or 95°F (R. W. Menzel, Florida State University, Tallahassee;
pers. comm.). Burrowing-rate response curves for M. mercenaria suggested a
preferred range of 21° to 31°C (70° to 88°F) in Rhode Island (Savage 1976);
growth was negligible at temperatures below 10°C or 50°F (Belding 1931; Pratt
and Campbell 1956) and at temperatures above 27° to 28°C or 81° to 82°F
(Menzel 1961, 1962). In his review paper, Ansel! (1967b) concluded that 20°C
(68°F) was the optimum temperature for growth of M. mercenaria and that the
rate dropped off symmetrically at higher and lower temperatures, ceasing
below 9°C (48°F) and above 31°C (88°F). In the laboratory, Storr et al.
(1982) observed that maximum overall shell growth of M. mercenaria occurred
at 12.8° to 15.6°C (55° to 60°F) and 23.9°C (75°F).
Substrate. Hard clams inhabit a variety of sediment types (Joint
Subcommittee on Aquaculture 1983). Hibbert (1976) found that growth rates of
M. mercenaria were similar on bottom substrates of 3% to 93% mud. Sims and
Stokes (1967) could not establish a consistent correlation of sediment
particle size with M. campechiensis population density for Tampa Bay,
Florida; however, Pratt (1953) showed that the population density of M.
mercenaria in Narragansett Bay, Rhode Island, was inversely correlated with
the particle size of the major sediment constituent. Average concentrations
of clams were greatest on predominantly muddy bottoms, less on sand bottoms,
and least on rocky bottoms. Clams were most abundant in predominantly fine
sediments, but in these sediments their abundance was generally a function of
the coarseness of the minor constituents. Clams do not grow well in silty
substrates. Pratt and Campbell (1956) found an inverse relationship between
growth of M. mercenaria and the fineness of the sediment (expressed as
percentage of silt and clay). The inferior growth was attributed to frequent
gill clearing, which expended energy and interfered with feeding. Johnson
(1977) also reported slower growth of M. mercenaria in finer sediment due to
increased expulsion of pseudofeces.
Soft sediment was given as the principal factor limiting the abundance
and diversity of benthic mollusks in bayfill canals in Tampa Bay, Florida
(Sykes and Hall 1970). Sediment types can be roughly correlated with current
velocities. A muddy bottom normally indicates calm water, whereas a coarse
bottom generally indicates more turbulent water (Thorson 1955). Wells
(1957a) found that the average density of M. mercenaria in Chincoteague Bay,
Maryland, was highest where the water current was 30 to 50 cm/s. The Joint
Subcommittee on Aquaculture (1983) reported that clams grow well where
currents average 50 cm/s or less.
Suspended solids. Little information is available on the effects of
suspended solids on adult clams. As clams filter feed, the uningestible
materials are sorted, accumulated, and expelled as pseudofeces (Pratt and
Campbell 1956). The rate of pseudofeces generation increases with turbidity
(Pratt and Campbell 1956). Menzel (1961) suggested that high turbidity may
have inhibited growth at one Florida site.
Food. Adult hard clams feed by filtering plankton and microorganisms
(Chestnut 1951). They may depend on an abundant supply of plankton before
and during spawning to provide sufficient energy to ripen the gonads (Ansell
1967a). In the laboratory, food concentrations of 300 mg carbon per liter of
seawater were optimal for feeding of M. mercenaria (Tenore and Dunstan 1973).
Robinson and Langton (1980) found that digestion was nearly continuous in a
subtidal population of M. mercenaria, regardless of time of day or tidal
stage.
Vegetation. Studies conducted in Florida's west coast estuaries have
reported the association of M. campechiensis with stands of turtle grass,
Thalassia testudinum (Schroeder 1924; Woodburn 1962; Sims and Stokes 1967;
Taylor and Saloman 1968, 1970; Godcharles 1971). This association with
rooted vegetation persists in inland waters but not offshore (Godcharles and
Jaap 1973; Menzel 1976). In Bogue Sound, North Carolina, Peterson (1982)
found a significantly higher (P < 0.01) average density of M. mercenaria in
partly vegetated plots than in unvegetated plots and a significantly higher
(P < 0.01) density in thickly vegetated than in partly vegetated plots.
Although seagrasses may provide protection from predators and stabilize the
sediment (Godcharles and Jaap 1973; Peterson 1982), they are apparently not
essential for the well-being and survival of clams.
Water depth. Hard clams seem to prefer relatively shallow water,
although they also are found in the open ocean. Along the South Carolina
coast, Anderson et al. (1978) found that 50% of all the M. mercenaria and M.
campechiensis were collected at a depth of about 2 m (7 ft), and less than
10% at depths greater than 5 m (16 ft). Cummins (1966) found commercial
concentrations of M. campechiensis off the South Carolina coast at depths of
7.3 to 11 m (24 to 36 ft). Along the west coast of Florida the most
productive beds were at 5.5 to 7.3 m (18 to 24 ft). M. campechiensis were
generally most abundant at depths of 4.7 to 9.2 m (15.4 to 30.2 ft) in Tampa
Bay, Florida (Godcharles and Japp 1973). Off Woods Hole, Massachusetts,
Robinson and Langton (1980) collected M. mercenaria at 6 to 8.5 m (20 to 28
ft). In Maine, Gustafson (1955) noted that M. mercenaria lives between tide
levels to depths of at least 15 m (49 ft).
HABITAT SUITABILITY INDEX (HSI) MODELS
Model Applicability
Geographic area. The model is applicable to intertidal and subtidal
estuarine habitats as defined by Cowardin et al. (1979) along the Atlantic
and gulf coasts and can be used for both M. mercenaria and M. campechiensis.
It is not applicable in the open ocean. The effect of pollution has not been
considered in model development; accordingly, the model is not applicable to
heavily polluted waters.
Season. The model is structured to account for seasonal variations in
habitat requirements of hard clams and, accordingly, to estimate the ability
of an area to sustain a population year-round.
Minimum habitat area. Hard clams can grow in relatively high densities.
The minimum area required for a self-sustaining population of hard clams is
not known.
Verification level. The model has not been field-tested. The
acceptable model output is an index value between 0.0 and 1.0. Dr. R.
Winston Menzel (Oceanography Department, Florida State University,
Tallahassee) and Dr. John J. Manzi (Marine Resources Research Institute,
South Carolina Wildlife and Marine Resources Department, Charleston) reviewed
and evaluated the hard clam model. Although their comments have been
incorporated, the author is responsible for the final version of this model.
Model Description
Overview. The structure of the hard clam HSI model is depicted
graphically in Figure 1. The habitat suitability index model applies to the
entire life cycle of hard clams. It uses two life requisites (water quality
and substrate-suspended solids) to evaluate an area. It is based on six
habitat variables: dissolved oxygen concentration (Vi), salinity (V2), water
temperature (V3), percent silt-clay concentration in the substrate (V4),
water current (V5), and suspended solids (V(>). A food component was not
included because of insufficient information on food availability in
estuaries. Vegetative cover was omitted because it is not an important
habitat characteristic throughout the range of hard clams. This model does
not apply to areas where clams are incapable of burrowing in the substrate.
Water quality component. Both adult and embryonic hard clams are
capable of withstanding low dissolved oxygen concentrations. The dissolved
oxygen requirements of the larval stage (V1 ) are used to evaluate
suitability. Dissolved oxygen is most important from April to September,
the period when most spawning occurs. Prolonged exposure to less than 4.0
mg/1 dissolved oxygen decreases the probability of survival of M. mercenaria
larvae. The optimal dissolved oxygen concentration is 4.0 mg/1 or higher.
Salinity (V2) affects growth and survival of all life stages. The
optimal salinity range for adult M. campechiensis is 24 to 35 ppt and 20 to
30 ppt for adult M. mercenaria. Growth of juvenile M. mercenaria is
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retarded at salinities of 22.5 ppt and lower. The optimal salinity range for
hard clams throughout their range is assumed to be 22 to 35 ppt.
Water temperature (V3) requirements for all life stages are combined.
Growth of adult M. mercenaria and M. campechiensis is negligible below 10°C
(50°F), and straight-hinged veligers of M. mercenaria are unable to digest
food at 10°C (50°F). Accordingly, temperatures of 10°C (50°F) or less are
considered unsuitable. Growth is negligible above 28°C (82°F) for M.
mercenaria and above 35°C (95°F) for M. campechiensis (Menzel 1961, 1962,
~$ers~. cömm.). Temperatures of 35°C (95°F) and higher are considered
unsuitable. The optimal temperature range for growth is assumed to be 20° to
31°C (68° to 88°F). Water temperature shou'd be measured only during the
growing season, where growing season is defined as the period of time when
the mid-depth water temperature is greater than 10°C (50°F).
The pH requirements were not included because marine and estuarine
waters normally have a high buffering capacity. An influx of pollutants,
such as hydrogen sulfide, might alter the pH, but the present model is not
designed to evaluate highly polluted areas. Water depth is not included
since hard clams are distributed throughout estuaries at various depths.
Substrate-suspended solids component. Hard clams are found in almost
any bottom type into which they can burrow. The percent silt and clay (V4)
in the substrate affects clam growth; as that percent increases, clams must
more frequently clear their filtering apparatus, which expends energy and
interferes with feeding.
Water currents (V5) affect the type of sediment found in an area and the
stability of the bottom; 30 to 50 cm/s is considered optimal.
Suspended solids (Ve) affect larval clam development. Growth of larvae
is normal at silt concentrations as high as 0.75 g/1, but growth decreases at
higher levels and ceases at 3.0 to 4.0 g/1. Turbidity should be measured
during the spawning season from April to September.
Suitability Index (SI) Graphs for Model Variables
Graphic representations of the relationship between the habitat vari-
ables and hard clam habitat quality are given here. Optimum suitability is
indicated by an SI value of 1.0 and unsuitability by a value of 0.0. The SI
graphs are based on the assumption that the suitability of a particular
habitat variable can be represented by a two-dimensional response surface and
is independent of other variables that contribute to habitat suitability.
Data sources and assumptions associated with SI graphs are listed in Table 1.
Table 1. Variable sources and assumptions for hard clam suitability indices.
Variable and source Assumption
Morrison 1971
Chanley 1958
Davis 1958
Woodburn 1961, 1962
Taylor and Saloman 1970
Castagna and Chanley 1973
R. W. Menzel, pers. comm.
Optimal dissolved oxygen concentra-
tion for larval M. mercenaria growth
and survival is 4.0 mg/1 or higher.
The optimal salinity range for growth
and survival of adult M.
campechiensis is 24 to 35 ppt while
optimal range for adult M. mercenaria
is 20 to 30 ppt. The optimal
salinity range for hard clams
throughout their range is assumed to
be 22 to 35 ppt.
Menzel 1961, 1962
Davis and Calabrese
Savage 1976
1964
Pratt and Campbell 1956
Johnson 1977
Growth of M. mercenaria and M.
campechiensis adults is neglTgible
below 10°C (50°F), and M. mercenaria
veligers cannot digest food at 10°C.
Growth is negligible above 28°C
(82°F) for M. mercenaria and above
35°C (95°F) for M. campechiensis.
Optimal range for growth is assumed
to be 20° to 31°C (70° to 88°F).
Clams must be capable of burrowing
in substrate. As percentage of
silt-clay content increases, growth
decreases.
Wells 1957a
Joint Subcommittee on
Aquaculture 1983
Davis 1960
Densities of clams are highest where
current velocities are 30 to 50 cm/s.
Larval clam growth is optimal at silt
concentrations of 0.75 g/1 or less.
10
Variable Description
Mean mid-depth
dissolved oxygen
concentration April
to September.
Suitability Graph
Mean annual mid-
depth salinity.
Mean mid-depth water
temperature during
growing season.
i n _ i . l.U - i Index 00 /
£ 0.6- •r— /
r— — /
1 0-4- (0
-M
•r- *^
^ 0.2- /
/
0.0-
10
~r
15 20 25
11
Variable Description Suitability Graph
Percentage silt-clay
in substrate sample.
0 25 50 75 100
Mean annual
surface water
current velocity.
Mean total suspended
solids April to
September.
12
Component Index (CI) Equation and HSI Determination
Development of an HSI for hard clams requires that the SI values for the
habitat variables be combined into component indices (CI) for water quality
and for substrate-suspended solids. Percentage of silt-clay (SI v4) is
squared because it is considered the most important habitat variable. The
suggested equation follows.
Component Equation
1/3 Water quality (WQ) (SI„ X SI„ X SIy )
Substrate-suspended solids (SS) (Slv?x SIvr X SIvJ "4 5 b
HSI = (WQ X SS)1/2
Sample data sets representing a range of hypothetical habitat values for
hard clams are presented (Table 2). The HSI values generated are believed to
reflect the relative potential of the habitats to support hard clams.
Interpreting Model Outputs
The hard clam HSI determined by use of these models does not necessarily
represent the population of hard clams in an area. Habitats with an HSI of 0
may contain some hard clams and habitats with a high HSI may contain only a
few. The proper interpretation of the HSI is one of comparison. On average,
habitats with high HSI's would be able to support higher populations of hard
clams than habitats with low HSI's. A close correlation between population
size and HSI is unlikely.
Table 2. Calculations of suitability indices (SI), component indices (CI),
and habitat suitability indices (HSI) for three hypothetical data sets on the
basis of habitat variables (V) and model equations.
"MTidel Data set 1 Data set 2 Data set 3
component
'6
Data SI Data SI
V, (mg/1) 6 1.0 3 0.67
V* (ppt) 20 0.0 26 1.0
yt. (°C) 25 1.0 15 0.5
V^ (%) 50 0.5 0 1.0
V* (cm/s) 0 0.6 25 0.93
V^ (g/1) 1 0.9 2 0.54
4 1.0
35 1.0
20 1.0
75 0.25
50 1.0
0. 5 1.0
1.0
0.5
WQ 0.0 0.69
SS 0.6 0.84
HSI 0.0 0.76 0.7
13
Field Use of Model
This model is designed for use in intertidal and subtidal estuarine
areas and not for open ocean areas. It is set up to evaluate habitat
suitability for both M. mercenaria and M. campechiensis. Inasmuch as
information on habitat variables was not always available for both species,
information pertaining to M. mercenaria was applied to M. campechiensis and
vice versa. Information was merged into a range of values when information
on both species was available. This adjustment may have caused the optimal
ranges of some variables to be wider than if each species was treated
individually. Since the two species interbreed and produce viable offspring,
this use of the habitat information seems to be justified.
The reliability of the calculated HSI values can be only as good as the
data used for their calculation. Estimates of variables cannot replace
field measurements of variables. The HSI values are most useful when the
habitat variables are measured in the specific evaluation area. Existing
water quality information for the area should be used if it is available and
accurate. Adult clams are capable of withstanding suboptimal conditions by
simply closing their shells. Accordingly, temporary fluctuations in water
quality characteristics may not influence habitat suitability. Since
spawning occurs over an extended period and involves massive numbers of eggs,
temporary fluctuations in water quality may kill only a portion of the yearly
production of embryos and larvae. Accordingly, it is best to use long-term
data whenever possible to evaluate the suitability of an area for hard clams.
In the northern areas of the hard clam distribution, water temperatures may
remain below the optimal range of 20° to 31°C (68° to 88°F) throughout the
year. If hard clams are known to inhabit these areas, the temperature
suitability curve may require adjustment.
Suggested methods for measuring model variables in areas where data are
not available are described in Table 3. A valid sampling scheme must be
developed before field sampling is done. It is insufficient to take only
a few samples. Local fluctuations in water quality affect only portions of a
hard clam population and do not determine the overall suitability of an area.
If subjective estimates must be used, they should be made by experienced
professionals familiar with the evaluation area and be accompanied by full
documentation of the basis on which they were made. If further information
is required on hard clams, see the detailed annotated bibliography published
by McHugh et al. (1982).
14
Table 3. Suggested methods for measurement of variables used in hard clam
HSI model. For all variables, use existing data if possible.
Variable Method
V Mid-depth dissolved oxygen can be
1 measured by using Winkler titration or
an oxygen meter (American Public Health
Association 1976). Measure from April
through September.
V Mid-depth salinity can be measured by
2 titration, refractometer, or salinity
meter.
V Mid-depth temperature can be measured
3 by thermometer or temperature probe
(American Public Health Association
1976).
V Percentage of silt-clay can be determined
4 by washing a known weight of sediment
through a 63-ym (Tyler series No. 250)
sieve. Silt and clay pass through the
sieve.
V Surface water current velocity can be
5 measured with a flowmeter.
V Turbidity can be measured by direct
° measurement of suspended solids
(American Public Health Association
1976). Measure from April to September.
15
LITERATURE CITED
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16
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18
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.19
Menzel, R. W., E. W. Cake, M. L. Haines, R. E. Martin, and L. A. Olsen.
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20
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21
50272-lor
REPORT DOCUMENTATION
PAGE
l._REPORT NO.
FWS/OBS-82/10.77
3. Recipient's Accession No.
4. Title and Subtitle 5. Report Date
Habitat Suitability Index Models: Hard Clam August 1984
7. Author(s)
R. Mulholland
8. Performing Organization Rept. No.
9. Performing Organization Name and Address
Florida Cooperative Fish and Wildlife Research Unit, USFWS
School of Forest Resources and Conservation
117 Newins-Ziegler
University of Florida
Gainesville, FL 32611
10. Project/Task/Work Unit No.
11. Contracted or Grant(G) No.
(C)
(G)
12. Sponsoring Organization Name and Address
National Coastal Ecosystems Team
Division of Biological Services
Research and Development
Fish and Wildlife Service
U.S. Department of Interior
Washington, DC 20240
13. Type of Report & Period Covered
IS. Supplementary Notes
16. Abstract (Limit: 200 words)
for A review and synthesis of existing information were used to develop a model suitable
evaluating estuarine habitat for hard clams (Mercenaria mercenaria and Mercenaria
campechiensis). The model is scaled to produce an index of habitat suitability between
0 (unsuitable habitat) and 1 (optimally suitable habitat) for the Atlantic and Gulf
coasts of the United States. Habitat suitability indices are designed for use with
the Habitat Evaluation Procedures previously developed by the U.S. Fish and Wildlife
Service.
17. Document Analysis a. Descriptors
Mathematical models
Estuaries
Mollusca
Habitat
b. Identifiers/Open-Ended Terms
Bivalves
Habitat Suitability Index
Hard clam
Mercenaria mercenaria
c. COSATI Field/Group
Mercenaria campechiensis
Venus mercenaria
Quahog
18. Availability Statement
Unlimited
19. Security Class (This Report)
Unclassified
21. No. of Pages
vi + 21
20. Security Class (This Page)
Unclassified
(See ANSI-Z39.18) OPTIONAL FORM 272 (4-77)
(Formerly NTIS-35)
Department of Commerce
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Services, Washington, DC
Eastern Energy and Land Use Team
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♦ Locations of Regional Offices
Western Energy and Land Use Team
Ft, Collins, CO
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II. s.
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