FIELD VALIDATION OF MULTI-SPECIES LABORATORY TEST
SYSTEMS FOR ESTUARINE BENTHIC COMMUNITIES
by
Robert J. Diaz
Mark Luckenbach
Sandra Thornton
Morris H. Roberts, Jr.
Virginia Institute of Marine Science
The College of William and Mary
Gloucester, VA 23062
Robert J. Livingston
Christopher C. Koenig
Gary L. Ray
Loretta E. Wolfe
Department of Biological Sciences
Florida State University
Tallahassee, FL 32306-2043
CR 812053
Project Officer
Dr. Thomas W. Duke
Office of the Director
Environmental Research Laboratory
Gulf Breeze, FL 32561
ENVIRONMENTAL RESEARCH LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U. S. ENVIRONMENTAL PROTECTION AGENCY
GULF BREEZE, FLORIDA 32561
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FIELD VALIDATION OF MULTI-SPECIES LABORATORY TEST
SYSTEMS FOR ESTUARINE BENTHIC COMMUNITIES
by
Robert J. Diaz
Mark Luckenbach
Sandra Thornton
Morris H. Roberts, Jr-
Virginia Institute of Marine Science
The College of William and Mary
Gloucester, VA 23062
Robert J. Livingston
Christopher C. Koenig
Gary L. Ray
Loretta E. Wolfe
Department of Biological Sciences
Florida State University
Tallahassee, FL 32306-2043
CR 812053
Project Officer
Dr. Thomas W. Duke
Office of the Director
Environmental Research Laboratory
Gulf Breeze, FL 32561
ENVIRONMENTAL RESEARCH LABORATORY
OFFICE OF RESEARCH AND DEVELOPMENT
U. S. ENVIRONMENTAL PROTECTION AGENCY
GULF BREEZE, FLORIDA 32561
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ABSRTACT
The aim of this project was to evaluate the validity of
using multi-species laboratory systems to assess the response of
eatuarine benthic communities to an introduced stress. Over a 5-
year period experiments in Apalachicola Bay, Florida, and the
York River, Virginia, sought to (1) develop criteria for
microcosm tests for evaluating the capacity of microcosms to
model natural communities in the presence and absence of a
pollution-induced stress, and (2) assess the validity of
extrapolating test results from one location to another.
Procedures for constructing, maintaining and sampling microcosms
were tested and refined over the study period. A large number of
laboratory and field tests were conducted synoptically over this
period, including experiments in which microcosms and field sites
were dosed with toxicants (mixed hydrocarbons in some and
pentachlorophenol in others). We have investigated various
methodologies for analysing and interpreting data derived from
microcosm tests.
The most promising results were achieved with medium-sized
2
microcosms (approximately 0.1 m ) in relatively short-term
experiments (5 weeks). Individual species response patterns in
the microcosms were highly variable and seldom showed good
agreement with patterns in the field. Species richness in the
microcosms and field showed good temporal agreement and provided
a conservative indicator of community response to toxic stress.
An eco1ogica 11y-based guild approach to grouping species proved
to be a powerful and reliable method of extrapolating from
microcosm test results to responses of field communities. Our
findings suggest that results from estuarine benthic-derived
microcosm toxicity tests may be used to predict some aspects of
community response to toxic stress. Further, the results
indicate some generality in these predictions which should permit
cautious extrapolation to other field sites.
This report was submitted in fulfillment of contract number
CR 812053 by the Virginia Institute of Marine Science and Florida
State University under the sponsorship of the U.S. Environmental
Protection Agency. This report covers a period from October 1981
to October 1985 and work was completed as of 1 March 1987.
11
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CONTENTS
Abstract ii
Figures iv
Tables vi
1. Introduction 1
2. Conclusions 2
3. Objectives 3
4. Study Sites 4
5. Ecological Characterizations ... 5
6. Methods 7
Experimental protocols 7
Dosing procedures 8
Toxicant levels 9
Guild assignments 10
Data analysis 12
7. Results and Discussion 13
Physical/Chemical data 13
Synopsis of test results 14
Recruitment patterns 18
Response variables 19
Predicting response to toxic stress 22
References 24
ill
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FIGURES
Number Page
1 Study Sites
a. Apalachicola Bay, FL 47
b. York River, VA 48
2 Mean Abundance in Weekly Samples from 1981-1986 49
3 Total Macrofauna and Species Richness -
Spring 1982 Test, Controls 50
4 Total Macrofauna and Species Richness -
Fall 1982 Test, Controls 51
5 Total Macrofauna and Species Richness -
Spring 1983 Test, Controls 52
6 Total Macrofauna and Species Richness -
Fall 1983 Test, Controls 53
7 Total Macrofauna and Species Richness -
Spring 1985 Test, Controls 54
8 Total Macrofauna and Species Richness -
Fall 1985 Test, Controls 55
9 Abundances of Dominant Guilds in Individual and
Combined Tests, Controls 56
10 Lab and Field PCP Levels - Spring 1985 Test
a. Low Dose 65
b . High Dose 66
11 Lab and Field PCP Levels - Fall 1985 Test
a. Low Dose 67
b. High Dose 68
12 Response to PCP - Total Macrofauna -
Spring 1985 Test 69
13 Response to PCP - Species Richness -
Spring 1985 Test 70
14 Response to PCP - Total Macrofauna -
Fall 1985 Test 71
IV
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15 Response to PGP - Species Richness -
Fall 1985 Test 72
16 Response to PCP - Selected Guilds -
Spring 1985 Test 73
17 Response to PCP - Selected Guilds -
Fall 1985 Test 78
v
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TABLES
Numb e r Page
1 Generalized Protocol for Laboratory Microcosm/
Field Validation Studies 26
2 Sampling Schedules for the Combined (FSU-VIMS)
Experimental Program (1981-1985) 27
3 Functional Group Assignments for Taxa Collected
a. Florida 28
b. Virginia 34
4 Species Composition of Dominant Guilds 38
5 Percent of Total Individuals in the Top 5 Guilds
in Each Test 41
6 Guilds Which Showed Good Agreement Between Temporal
Trends in the Lab and Field 45
7 Evaluation of Concordance Between Laboratory and
Field Results for PCP-dose Experiments 46
VI
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SECTION 1
INTRODUCTION
A priority of environmental toxicology is to predict the
ecological effects of a toxic substance by extrapolating from
controlled laboratory experiments. Until recently such
experiments have generally been restricted to sing 1 e-species
acute tests. Much of the rationale for this approach has been
based upon the assumption that acute tests with the most
sensitive species provide conservative estimates of environmental
impact, an assumption which has recently been criticized (Kimball
and Levin, 1985; Cairns, 1983; 1986a). Despite the fact that
arguments can still be made for the utility of single-species
testing (Wies, 1985), there is growing recognition of the need
for multi-species toxicity testing (Cairns, 1985).
As the use of multi-species laboratory test systems
(microcosms) increases, a requisite part of the development must
be field validation. We accept here the definition of validation
offered by Cairns (1986b) as the testing of "the ability to
predict the relationship between the response of the artificial
laboratory system and the natural system." There are several
components to any such evaluation. The first involves
establishig criteria for conducting microcosm tests which are
specific enough to reduce undesirable laboratory artifacts and
general enough to be of utility in a range of habitats. Second,
it is necessary to evaluate the capability of the laboratory
system to model temporal patterns in the natural system in the
absence of toxic stress. Only after this does it become
appropriate to compare the response of the microcosm and field
communities to a po1 lution-induced stress. Finally, if microcosm
tests are to have applicability outside of the site-specific
system in which they are conducted, it is necessary to evaluate
the validity of extrapolating between systems.
Towards the end of validating an estuarine benthic microcosm
test system, we initiated a 5-year program in two estuaries.
Using macroinvertebrate and microbial communities from
unvegetated, soft-sediment habitats in Apalachicola Bay, Florida
and the York River tributary of the Chesapeake Bay, Virginia, we
conducted a series of combined 1aboratory/fieId experiments to
address the questions posed above. The details of the individual
experiments have been reported earlier (Diaz et al., 1984, 1986;
Livingston et al., 1985a, 1985b, 1985c, 1985d, 1986) and we will
not dwell on those details here but rather summarize the overall
project, its findings and draw conclusions regarding the use of
benthic microcosms for predicting environmental consequences of
toxic stress.
1
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SECTION 2
CONCLUSIONS
Variability in natural estuarine systems is high,
necessitating large numbers of experimental replicates and
samples to observe even major responses. Careful attention must
be paid to physical/chemical features of the microcosms
throughout the tests to insure that conditions remain as close to
those in the natural field sites as possible. Monitoring of
toxicant levels and distribution within the microcosms throughout
the experiment is necessary to evaluate dissipation and breakdown
of the toxicant. Concurrent with laboratory testing, samples
from the field sites are required to assess natural fluctuations
in the benthic populations. Temporal variation in recruitment
adds year to year and site to site variation in community
responses in microcosm tests. To overcome this problem it is
mandatory that microcosm tests be properly timed to corresepond
with known stages in recruitment cycles. Furthermore it is
necessary that only community components which show good
agreement between laboratory systems and field sites be used to
evaluate response to toxins. In this respect species richness of
the community and the numerical abundances of certain guilds
(listed in Table 6) appear to be the best components to use.
We advocate an approach of categorizing species into
"ecological types" or guilds which has several advantages. This
categorization gives a managable number groupings — enough to
provide some detail but few enough to permit reasonable detection
of patterns. The emphasis on species groupings reduces the
dependence of the predictions upon single species which may be
highly variable in their occurrence from year to year. Those
guilds which are observed to behave aberrantly in the laboratory
may be excluded from the analyses a priori. And, the use of
"ecological types" facilitates comparisons among sites which have
different species compositions. However, this approach requires
good ecological characterization of the species comprising the
benthic community used in the testing. These ecological data are
often difficult to obtain.
We conclude that laboratory microcosms can provide a
valuable tool for assessing natural benthic community responses
to toxic stress, provided that the caveats and conditions stated
in this report are heeded.
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SECTION 3
OBJECTIVES
The primary objectives of this project were:
(1) the development of criteria for conducting
microcosm tests and interpreting the results;
(2) the evaluation of the capacity of a benthic
microcosm system to simulate natural field
communities in the absence of a toxicant;
(3) the comparison of response patterns of
laboratory and field communities to a
pollution-indueed stress; and
(4) the determination of the validity of
extrapolating from microcosm tests conducted
in one locale to natural communities in
anothe r .
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SECTION 4
STUDY SITES
The study sites in the Apalachicola Bay system (East Bay and
St. George Sound) were located in polyhaline and oligohaline
areas, and those in the York River in the meso-polyhaline portion
of the estuary (see Fig. 1). All sites were shallow (1-2 m) ,
unvegetated areas. Sediments in the oligohaline site were silty
sand, and sediments in the polyhaline and meso-polyha1ine sites
were predominately fine sands. Each of the study sites are
considered representative of extensive portions of temperate
estuaries. For both the Virginia and Florida experiments, the
laboratory microcosms were located near the field study sites.
More details of the study sites are given in earlier reports
(Diaz et al., 1984, 1986; Livingston et al., 1985, 1986).
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SECTION 5
ECOLOGICAL CHARACTERIZATRIONS
An essential part of this program was an understanding of the
ecological backdrop against which the experiments were conducted.
Weekly monitoring programs for infaunal macroinvertebrates have
been ongoing at the Virginia site since 1979 (Diaz, 1984) and at
the Florida site since 1981. Ten replicate samples per week were
collected with 5.0 cm and 7.5 cm diameter hand-held corers in the
York River and Apalachicola Bay sites, respectively. These
samples were processed on a 250 um and a 500 um sieve series and
all macrobenthic invertebrates identified to the lowest possible
taxon and enumerated. Figure 2 shows weekly mean abundances of
total macrofauna from the Apalachicola Bay and York River sites
from October 1981 through April 1986 and indicates the dates of
the laboratory/field experiments. An important point of
comparison between these sites is the timing of recruitment
events. In Florida peak recruitment generally occurred in the
fall and the greatest abundances and species richness were
observed in the winter. In Virginia the pattern was temporally
reversed with recruitment peaks occurring in the spring. The
relationship between the timing of the experiments and seasonal
patterns of recruitment is crucial to the interpretation of
variability in the data.
In addition to these background data on faunal abundances we
have found that an appreciation of trophic structures and
physical disturbance processes at each site is necessary for
interpreting our experimental results. Predation by bottom-
feeding fishes and decapods appears to be an important process
shaping benthic communities at each site (Virnstein, 1977; Dugan
and Livingston, 1982). Physical disturbance, both periodic
(waves) and aperiodic (storms) impact on these communities.
During the course of this project each site was impacted by at
least one major storm event which hit during the lab oratory/fieId
experiments. The timing of microcosm tests in relation to
predator utilization of the habitats and disturbance events in
these sites was a crucial component of proper experimental
design.
Another essential feature of our ecological characterization
of the field sites was an understanding of species - specific
functional roles in the community. Information on trophic,
mobility and reporductive modes was a central part of our
analysis effort. This is discussed in greater detail in the
section on guild assignments. We emphasize at this point,
however, that even with the extensive data which have been
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collected from each of these sites, much of this type of
detailed, species-specific information is lacking.
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SECTION 6
METHODS
EXPERIMENTAL PROTOCOLS
The focus of experiments conducted during 1982 and 1983 was
to establish criteria pertaining to microcosm construction,
microcosm maintenance, test duration, sampling procedures and
response variables. In addition treatments were employed to
assess the impact of predator exclusion and inclusion in the
field sites.
Microcosm communities were constructed of a series of cores
collected with diver-operated box cores (10 x 20 cm; 10 cm deep).
Cores were arranged contiguously on seawater tables in the same
spatial arrangement as in the field. A wide range of microcosm
sizes have been tested. During 1982 and 1983 experiments at both
2
sites were conducted in microcosms ranging from 0.8 to 1.0 m in
size. Additional experiments in Florida in 1983 compared three
22 2
microcosm sizes: 0.67 m , 0.067 m , and 0.0084 m . The spring
2 2
1985 experiment in Virginia compared 1.00 m and 0.11 m
microcosms. Our objectives here were twofold: (1) to assess
whether microcosm size affected the ability of laboratory
community dynamics to track those of the field, and (2) to
determine whether it was preferable to use larger microcosms
which could be sampled repeatedly or smaller ones which must be
destructively sampled. The details of results from these
experiments are given in earlier reports and are summarized as
2
follows. Small microcosms (0.0084 m ) contained fewer species
than the field sites and showed considerable divergence in
2
community parameters from the field. Medium (0.08 - 0.11 m ) and
2 .
large (0.67 - 1.00 m ) microcosms contained similar numbers of
species and generally showed the same degree of concordance
between laboratory and field populations. Replicate large-sized
microcosms were sampled repeatedly throughout the duration of
experiments, while individual replicates of medium-sized
microcosms were sampled at only one time period and discarded.
The disturbance associated with repeated sampling of large
microcosms was judged to have an impact on community and
population dynamics, so we settled on the medium-sized microcosms
2
(approximately 0.1 m ). With a microcosm of this size a large
number of replicates must be established at the initiation of an
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experiment and a portion destructively sampled at each sampling
t ime .
The sizes of the core samplers employed at each site (5 cm
Virginia; 7.5 cm Florida) were based upon our experience with the
field monitoring programs and were selected to provide adequate
sampling of most resident macrofauna. Throughout the experiments
the same size coring devices were used to collect laboratory and
field s ample s .
Test durations in 1982 and 1983 ranged from 5 to 9 weeks
during which time laboratory and field treatments were sampled
synoptically on a weekly or biweekly basis. Samples were sieved
on 250 um and 500 um mesh screens, and mac ro inve r t eb r a t e s were
identified to the lowest possible taxon and enumerated. A
variety of community and population statistics were considered
(see below) and most showed divergence between the laboratory and
field 5 weeks after initiation. On this basis we adopted a 5-
week duration for subsequent dosing experiments.
Containers with azoic sediments were placed in the seawater
table at both sites during the 1985 experiments. These
defaunated treatments were sampled and processed similarly to the
microcosms and were used to monitor recruitment into the
laboratory system through the seawater intakes.
Throughout the experiments physical and chemical
measurements were made in the laboratory and field. Temperature,
salinity, dissolved oxygen, sediment grain size and sediment
organic content were monitored regularly. Periodic measurements
of pH, sediment temperature and Eh were also made.
Field treatment locations were located haphazardly within
pre-selected sites and marked with metal frame structures (2 m x
2 m bottom area x 3 m high). These frames served as a means of
relocating sample sites and of holding a sample platform. The
sample platform had a gridded array of sample ports which
permitted individual core samples to be taken in pre-dete rmined ,
random locations within the treatments. Field treatments in the
various preliminary tests included (1) uncaged sites demarcated
only by the open metal frames, (2) caged sites in which the
frames were wrapped with screening to exclude predators, and (3)
caged sites with predators included. In addition field
treatments were dosed with toxin-laden sediments (see below).
All field treatments were established in triplicate and each
treatment replicate was sampled with 10-15 randomly located
replicate cores.
A generalized protocol of these methods is given in Table 1
and a schedule of experiments is presented in Table 2. For
greater details concerning the protocols for each test earlier
reports (cited above) should be consulted.
DOSING PROCEDURES
Experiments in which both laboratory and field sites were
dosed with toxicant-lad en sediments were conducted in the fall of
1983 and the spring and fall of 1985. In 1983 "naturally"
hydrocarbon contaminated sediments from the Elizabeth River, VA,
8
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were used to dose both laboratory and field treatments in the
York River and Apalachicola Bay. In both the spring and fall of
1985 unc ontaminated sediments were coated with pentachloropheno1
(PCP) to provide controlled-dose treatments for laboratory and
field sites. Our goal here was to evaluate the response of the
laboratory system to the stress relative to the response of the
field system (Objective 3).
During the spring 1985 experiment we tested dosing
procedures in which PCP contaminated sediments were added in
approximately 1 cm and 0.1 cm thick layers. No overt effects of
adding uncontaminated sediments were noted and we found that the
greater thickness of sediment provided more reliable dosing of
treatments, thus we adopted this procedure in the fall 1985
experiments. Laboratory dosing in each experiment was conducted
by spreading contaminated sediments uniformily over the microcosm
surface. Field dosing procedures involved two approaches. In
the fall 1983 and spring 1985 experiments in both Apalachicola
Bay and York River sites dosing was carried out by wrapping the
metal frames with plastic to reduce water flow, adding the
sediments to the enclosed water column, and removing the plastic
after sediments had settled to the bottom. This procedure was
successful in Apalachicola Bay, but not in the York River where
the plastic wrapping was insufficient to stop the stronger
currents (see Results and Fig. 10). During the fall 1985
experiment the same procedure was used in Apalachicola Bay and a
dosing box was used in the York River to apply toxin-laden
sediments. The dosing box was a large wooden box to which
sediments were added through a door on the top, the box was then
submerged and a false bottom removed to permit the sediments to
fall to the sediment-water interface. These methodologies were
successful at achieving dose equivalency between the field and
laboratory treatments (Fig. 11).
TOXICANT LEVELS
The hydrocarbon contaminated sediments from the Elizabeth
River used in the fall 1983 experiments were applied at nominal
concentrations; the wide variety of pollutants in these sediments
prevented the actual levels from being monitored. Lu (1982)
reported a detailed hydrocarbon analysis of the sediment at the
station from which contaminated sediments were obtained. In the
PCP-dosed experiments (spring and fall 1985) a high concentration
(nominally 10 ppm) and a low concentration (nominally 1 ppm) were
used. Actual concentrations of PCP in the laboratory and field
treatments were monitored throughout the test duration. These
analyses, which were carried out using methylene chloride
extraction and standard gas-liquid chromatography methods with
flame ionization and electron capture detection, proved to be
costly and time consuming but necessary. These data were
invaluable both for establishing when dose equivalency between
the laboratory and field was achieved and for tracking the time
course of the toxicant levels in each treatment.
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GUILD ASSIGNMENTS
In the latter portion of this project we became aware of the
need for grouping species for the purpose of analysis.
Community-1eve1 statistics, though they provided some useful
information, obscured much of the details of response within the
community, and individual species population fluctuations were
too numerous and variable to permit clear interpretation of
community response. Grouping species according to higher
taxonomic levels (e.g. polychaete families, oligochaeta,
bivalvia) was attempted as a solution, but even closely related
species can play different functional roles within a community,
and the responses of species within these groups were often
heterogenous . Thus we classified each species into functional
groups based upon the manner in which they used resources, how
they lived and moved in the sediments, and their mode of
reproduction. The categories to which species were assigned are:
Trophic Mod e
sc aveng e r
deposit-feeder
suspension feeder
interface feeder
predator
scraper
unknown
Trophic Level
carnivore (>90% animal matter)
herbivore (>90% plant matter)
detritivore/omn ivore
unknown
Mob i1i ty Mod e
bur rowe r
mob i1e
sessile
tub e-builder
mob i 1 e
sessile
ep i faun a 1
mob i 1 e
sessile
Reproductive Mode
plank tonic larvae
demersa1 egg cases
b rcode r s
asexual
unknown
Assignments were made using published information (esp., Fauchald
and Jumars, 1979) and personal observations. In making these
assignments we took a limited view of the environment, choosing
10
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as our point of reference the spatial scales relevant to our
treatments. Therefore species which move on scales of em's to
m's were classified as mobile. The intent of the reproductive
mode category was to separate those species which have the
capability of reproducing and recruiting from within the
microcosms from those which do not. Therefore we pooled
categories to create a composite classification:
Dispersal Mode
limited dispersal
wide dispersal
variable dispersal
unknown
Here again the spatial scale is defined to reflect our interest
in processes relevant to the microcosms. For instance, maldanid
polychaetes (represented primarily by Ax iothella muc o s a at the
Apalichcola Bay site and by Clymene1 la torquat a at the York River
site) produce demersal egg cases which generally remain attached
to the tops of the adult tubes until hatching. Juvenile maldanid
polychaetes then crawl away and build tubes of their own. This
type of reproduction leads to limited dispersal in the context of
the microcosm since it permits these organisms to recruit from
within the microcosm. Another example of a limited disperser in
our categorization is Paranais litoralig , an asexually
reproducing oligochaete. The limited dispersal category is not
intended to imply that these species in nature do not exhibit
wide ranging dispersal, but merely that they clearly have the
capability of recruiting from within the microcosm. By contrast,
other species have obligate planktonic stages which preclude
successful development within the microcosms. These species are
categorized as wide dispersers to indicate their inability to
recruit from within the laboratory seawater tables. A few
species are variable in their reproductive modes both between and
within sites. The spionid polychaete S t reb 1 o sp io benedicti . for
instance, exhibits variable reproductive strategies ranging from
fully planktonic development to brooding (Levin, 1984). In the
York River estuary S.. b ened ic t i appears to be entirely planktonic
in its development and is therefore classified as a wide
disperser in Virginia, while in Apalachicola Bay both types of
development have been observed for £.. benedicti and it is
classified as a variable disperser in those experiments. Table 3
gives the functional group assignments for all species collected
from the Florida and Virginia study sites. We recognize the
tentative nature of some of these assignments and stress the need
for more ecological data to refine this approach.
Unique combiniations of these functional groupings were
used to define guilds, e.g depos it-feed ing , detritivore/omnivore,
mobile burrower, with wide dispersal. This approach yielded a
total of 59 guilds in the two study areas, of which only 17 were
composed of single species. The species compositions of dominant
guilds in each site are given in Table 4. At each location the
five most abundant guilds generally comprised >80% (and never
1 1
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less than 40%) of the total number of individuals collected.
Details of this for each test are given in Table 5.
This approach of categorizing species into guilds served two
purposes. First, it permitted us to identify those guilds of
organisms for which laboratory microcosm populations do not serve
as good analogs of natural populations in the absence of any
toxicant. These types of organisms can be excluded a. priori from
analyses to assess toxic impact. The second advantage to this
approach is that the identification of types of organisms which
act as ecological units facilitates comparisons between
microcosms and field sites from different locations. For
instance, while the species composition varies between the
Virginia and Florida sites, functionally similar ecological
groups are found in both sites and provide a basis for
c omp ar i s on.
DATA ANALYSIS
Throughout the course of this project we have made use of
large numbers of replicates and the robustness of Analysis of
Variance (ANOVA) to test for specific treatment effects in the
highly variable data sets. This approach has generally been a
powerful one and several significant treatment effects have been
identified. For instance, ANOVA can test for significant
differences in total abundance between laboratory field
treatments. However, the central question we have posed is not
so straightforwardly tested. In particular we ask, can microcosm
test results be used to predict the response of natural
communities? Cairns (1986b) pointed out that the absolute
response in a microcosm test need not be identical to that in the
natural system. It is simply necessary that we know the
relationship between the response in the laboratory and the
field. In this regard the temporal patterns of community, guild
or species response in the laboratory and field may be very
similar but of different magnitude and still be of utility for
predictive purposes. Statistical procedures which test for
differences between treatment means (such as ANOVA), but yield
nothing about the similarity of pattern, would miss this
similarity. Proper testing for similarity in such patterns would
require a non-parametric pattern analysis capable of dealing with
widely vairant data; we are not aware of such a test at present.
Therefore, to answer this final question we are forced to rely
upon subjective evaluations. The large number of experiments
together with the persistence of many of the patterns add
strength to these assessments.
12
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SECTION 7
RESULTS AND DISCUSSION
The data generated by this project are voluminous, and any
value gained by their complete inclusion here would be offset by
the drawbacks of such a massive document. Therefore complete
data files from the project have been archived in computer files
at FSU and VIMS and are available on request. Below we present a
summary of our findings emphasizing particularly those aspects
which address the primary objectives outlined above.
PHYSICAL/CHEMICAL DATA
Care was taken to maintain physical and chemical
characteristics of the microcosms as close to those of the field
as possible, yet some differences still arose. Eh profiles and
visual inspection of sediment color indicated that depth of the
oxygenated layer within the microcosm sediments decreased with
time. This effect was generally most pronounced after week 5 in
any given test and led to significant changes in the depth
distribution of organisms. Similar changes were not apparent in
the field over similar time courses.
Surface sediment composition in the microcosms also showed
differences from the field sites. Fine sediments (silts and
clays) and organic content increased in the microcosms with time.
These increases were the result of deposition of fine particles
brought into the laboratory in the seawater system and were not
observed in the field. In addition rapid changes in sediment
composition in the field were observed in association with storm
events which had no effect upon the microcosm sediment
characteristics.
Water and sediment temperatures in the microcosms were
slightly more variable than those in the field sites, but this
degree of variation apparently was not sufficient to pose
problems. Salinities in the laboratory and field treatments were
similar throughout all experiments.
We refer the reader to earlier reports for more information
regarding physio-chemical factors in each of the 1 aboratory/fie Id
experiments. Here we emphasize our finding that careful
attention to the parameters listed in Table 1(1.A) is an
important component of successfully conducting a microcosm
experiment. Divergence between the laboratory and field in one
or more of these parameters will lead to divergence of the
c ommun i t i e s.
13
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SYNOPSIS OF TEST RESULTS
Sprint? 1982 Experiment*
F lorida--
The field predator inclusion treatment followed the field
controls in terms of the response of infaunal numerical
abundance. The field exclusion treatment was characterized by
high numbers (primarily Med i oma s tu s amb i s e t a) . An increase in
total macrofaunal numbers was also observed in the laboratory,
but not in the field controls (Fig. 3); these results were
interpreted as the release of specific opportunistic polychaetes
from predation pressure. Mediomastus was one of the few
populations that was still recruiting at the time of the
experiment. Species richness was generally unaffected by
treatment (Fig. 3). The proportional abundance of functional
feeding groups was more conservative, showing no change in the
field controls and inclusion treatment and only slight changes in
the field exclusion and laboratory treatments.
Virg inia —
Species specific responses to treatments were variable. For
six of the 11 dominant species there were significant differences
in abundance among treatments, but only five species showed
significant variation with time. Paranais littoralis and newly
set bivalves were the only two forms to show effects of both
treatment and time. Streblosp io benedicti. E teone he t e ropod a .
and immature Capitellidae all increased with time. Polydora
1i g n i decreased and newly set bivalves increased and then
decreased with time. Variance to mean ratios for all eleven
numerically dominant species exceeded one. Total macrofaunal
abundances in the laboratory declined sharply between weeks 3 and
4, and by week 5 showed considerable divergence from the field
controls (Fig. 3). Species richness in the laboratory was
similar to the field treatments throughout most of the
experiment, but began to diverge slightly by the fifth week (Fig.
3).
Fall 1982 Experiment
Florida--
These experiments were conducted in the oligohaline site.
Abundance increased in the laboratory by week 4 (Fig. 4),
probably attributable to a release from predation. Trends in
total macrofauna abundance among the various field treatments
were similar, as were species richness values across all
treatments. When expressed as feeding modes and trophic groups,
the various field treatments showed comparable patterns through
time with a predominance of be 1 ow-surface, deposit-feeding
detritovores/omnivores. The laboratory treatments showed gradual
change to a predominance of browsing omnivores. By the fifth
week of the experiment laboratory treatments showed substantial
divergence from the field treatment.
14
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Virginia--
Total macrofaunal abundance began to diverge during the
first week of the test (Fig. 4). Low, but significant, levels of
recruitment into the field sites by Streblospio b ened ic t i and
Tub if ic oidea spp. contributed to this pattern. Recruitment into
the microcosm was essentially absent. These recruitment pulses
in the field however were dampened (presumably by predation) and
abundance levels in the laboratory and field appeared to be
converging at termination of the experiment (week 6). Species
richness values were similar in the microcosm and field
throughout the experiment (Fig. 4).
Spring 1983 Experiment
F lorida--
Results of the spring 1983 experiment (o1igoha1ine , station
3) indicate similar results in the various field treatments with
reduced numerical abundance in the laboratory microcosms (Fig.
5). Species richness trends were similar in all treatments. In
this experiment, feeding modes and trophic group proportions were
similar among all treatments in the field and laboratory. Both
mean faunal abundance and species richness were representative of
field conditions.
Virginia—
Macrofaunal recruitment occurred at the York River site
during this test, but only two species showed dramatic increases:
Strebloapio benedicti and E t e on e heteropoda . Both species
reached their greatest abundances in the field cage treatments
and remained low in abundances in the microcosms where their
recruitment was restricted. Again, we interpret the lack of
major population increases in the field control site as resulting
from p o s t-recruitment mortality (probably from predation). Both
total macrofauna abundance and species richness reflect
recruitment events which occurred in the field but not in the
microcosms (Fig. 5).
Fall 1983 Experiment
Flor id a-~
Results of this experiment (polyhaline, station ML) indicate
similar macrofaunal numbers in the field treatments whereas
numbers tended to be reduced in the laboratory treatments. A
comparison of macrofaunal abundance in the field and laboratory
(Fig. 6) reveals that recruitment occurred into the field sites
but not into the microcosm. Once again, temporal patterns of
species richness were similar in the various field and laboratory
treatments, although numbers of species were lower in the
laboratory microcosms. Functional feeding modes and trophic
organization of the invertebrate assemblages were similar in all
treatments; temporal variability of these indices was low with a
predominance of below-surface deposit feeders as detrita 1-feeding
omnivores. Toxic sediments did not appear to affect the field or
15
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laboratory numerical abundances or species richness. Once again,
functional feeding groups and the trophic organization appeared
similar in all treatments (laboratory and field). The toxic
sediments had no overt effect on the laboratory or field
microcosms when viewed as feeding or trophic entities.
Virg inia--
The microcosm treatments consistently had lower abundances
and species richness than their field counterparts (Fig. 6).
Increases in total abundance and species richness in the field by
week 3 are indicative of recruitment events which did not occur
in the laboratory. Individual species response in the control
treatments (laboratory and field) were highly variable as some
species increased and others declined over the period. The
addition of non-toxic York River sediments to laboratory and
field treatments did not substantially change either faunal
abundance or species richness. Toxic Elizabeth River sediments
caused declines in laboratory and field treatments, but the
magnitude of the response was greater in the laboratory- The
dose treatments altered total abundances, species richness and
guild makeup .
Spring 1984 Experiments
Virginia--
Total macrofaunal abundances in this test were similar in
the laboratory and field treatments until week 4 of the study
when recruitment peaks occurred in the field. Recruitment did
not occur in the microcosms at this time and result was a nearly
3-fold difference between abundances in the field and microcosm
controls. Decline in numbers of macrofauna after the field
recruitment peak was rapid and within one week abundances within
the laboratory and field controls were again similar- Species
richness was again a fairly conservative parameter and was
generally similar between the laboratory and field treatments.
Spring 1985 Experiments
Florida--
Figures lOa and lOb show the concentrations of PCP in
laboratory and field treatments during the time course of this
experiment. Good dose-equivalency was achieved in the Florida
experiments between laboratory and field concentrations. Dose-
specific effects on total macrofauna and species richness are
shown in Figures 12 and 13, respectively. The impact on field
assemblages was less severe than on microcosm assemblages, with
only slightly lowered abundances and small reductions in species
richness evident. The laboratory effects included a relative
increase carnivores. Laboratory controls showed increased
abundance of subsurface deposit feeders relative to the field
treatments. Dose related changes in functional groups did not
occur in the field treatments. A real difference was evident in
the vertical distribution of the infaunal populations between
16
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laboratory controls and field populations. By the end of the
experiment, high numbers were concentrated in the top two
centimeters of the laboratory controls. In the laboratory, most
species disappeared from the bottom-most layer (8-10 cm) by the
end of the experiment. Ax iothe 11 a mucosa contributed to most of
the observed trends in vertical distribution. Species such as
Med iomas tus and Brania were adversely affected by both lab and
field PCP treatments. This trend of relative dominance was
directed by recruitment of Axiothella in the laboratory controls
by the third week of the experiment (T3). Recruitment in the
field was not affected by PCP treatment.
Virginia—
In the spring 1985 experiment, at the York River site, good
dose equivalency between the laboratory and field treatments was
not achieved (see Figs. lOa & lOb). PCP levels were consistently
lower in the field than in the microcosm. Mean macrofaunal
abundance in the laboratory declined markedly during the first
week, but this decline was observed in undosed control treatments
and was thus not a response to PCP dosing (Fig. 12). A slight
reduction in macrofaunal abundance was observed in field dosed
treatments relative controls (Fig. 12). Species richness showed
a clear dose-specific response in the laboratory, but was
unaffected by the lower doses achieved in the field (Fig. 13).
Fall 1985 Experiments
Florida—
Dose equivalency between the laboratory and field treatments
was again achieved in the fall experiments in Florida (Figs, lla
& lib). Experimental results were similar to those during the
spring experiment with strong, dose-specific reductions in
numerical abundance and species richness in the microcosm and
slight effects in the high PCP treatment in the field (Figs. 14 &
15). Recovery was rapid in the field due to high recruitment and
slower in the laboratory where recruitment was minimal. Species
such as Med iomas tus were again adversely affected by the
laboratory PCP treatments. Recruitment of this species from
within the laboratory was low, either as a direct or indirect
result of PCP treatment. In the field recruitment was apparently
unaffected by PCP exposure, with the possible exception of some
very short-term effects on Med iomas tus. Functional feeding and
trophic organization were unaffected by PCP treatment in the
field. In the laboratory, there were proportional changes in
these relationships at high PCP concentrations which included
trophic simplification. The percent of primary carnivores tended
to be higher in the PCP-treated microcosms.
Virginia--
Comparable levels of PCP were achieved between laboratory
and field treatments during the fall 1985 experiment (Figs, lla &
lib). Macrofaunal abundance in the laboratory showed slight
declines in the high dose treatment but was unaffected by the
17
-------
lower dose (Fig. 14) In the field treatments total macrofauna
abundance did not decline in response to PCP treatment; in fact
recruitment peaks were evident earlier in the high dose treatment
than elsewhere. Species richness in the laboratory declined
sharply in the high dose treatment, but was unaffected in the
microcosm low dose treatment (Fig. 15). A similar trend was
observed in the field, with lowered species richness in the high
dose treatment. This effect in the field however was less
dramatic and recovery was fairly rapid (Fig. 15).
RECRUITMENT PATTERNS
The experiments outlined above were timed to coincide with
peak recruitment seasons in both environments since this is the
period during which the communities are expected to show the
greatest sensitivity to toxic stress. However, recruitment of
benthic invertebrates is highly variable both spatially and
temporally, raising the need to distinguish between variability
in the data resulting from recruitment variations and those
resulting from treatment effects. Though the general timing of
peak recruitment periods at each site is predictable and our
experiments spanned portions of these periods (see Fig. 2), it is
not possible in any given year to predict either the precise
timing or magnitude of recruitment for any individual species.
Differences in recruitment levels between the laboratory and the
field can lead to order-of-magnitude differences in the
abundances of individual species and total macrofaunal numbers.
The azoic sediment treatments in the seawater table at the
Florida and Virginia sites have revealed that recruitment of
macrobenthic invertebrates through the seawater systems is
minimal. Recruitment events in the field during the course of an
experiment may lead therefore to substantial differences betwen
laboratory and abundances. For instance, at the York River site
in the spring 1982 experiment recruitment of Streblospio
benedicti . E t e o n e h e t e r op od a and immature Capitellidae resulted
in large differences between laboratory and field abundances
throughout the experiment. In the fall 1982 experiment at the
same site low levels of recruitment in the field by £ . bened ic t i
and Tub ificoides spp. caused only moderate divergence between
laboratory and field abundances. During the spring 1984
experiment laboratory and field abundances were similar until the
fourth week when a large recruitment event by S.. benedicti led to
three-fold differences in total abundance. Similar temporal
differences in recruitment were observed at the Florida site.
While recruitment of macrofauna into microcosms through the
seawater system was negligible, recruitment from within the
microcosms was occasionally substantial. Species which reproduce
asexually, have demersal eggs or brood their young have the
capability to reproduce and recruit from within the microcosms.
When a species recruited from within the laboratory it suffered
less mortality from epibenthic and demersal predators and from
sediment disturbance than in field, resulting in large
18
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differences between laboratory and field abundances. This
appears to have occurred in the spring 1983 experiments in
Florida during which Axiothella muc o s a recruited via demersal
eggs and increased dramatically in the laboratory. Also, in the
spring 1985 experiment in Virginia the asexually reproducing
faranaia littorali q attained higher densities in the laboratory
than in the field.
In recognition of the interpretationa 1 difficulties which
arise as a result of these recruitment differences we have taken
two approaches towards drawing inferences from these data.
First, as outlined above, guild designations include a
reproductive component; this groups together species which at
least have the potential to display similar recruitment
differences between the laboratory and field. Second, the
emphasis we place on similarity of temporal patterns of abundance
rather than absolute magnitudes of abundance reduces the problems
associated with varying levels of recruitment.
RESPONSE VARIABLES
An important part of addressing our objectives was to
determine which (if any) characteristics of benthic
macroinvertebrate communities were modelled well in the
laboratory and could therefore be used to predict responses of
natural communities. The greatest detail is of course obtained
by examining the population responses of individual species, and
in earlier reports we have devoted considerable attention to the
dynamics of at least the dominant species. Some species-specific
patterns have emerged from this effort [e.g. Streblospio
bened ic t i response in the laboratory and field are similar when
experiments are conducted during times of no recruitment; or
Ax iothella muc o s a may undergo population explosions in the
laboratory during its recruitment times]; these individual
patterns may be pieced together in an effort to make generalized
predictions. Yet the number of species is large and the variety
of response patterns observed is great. No doubt many general
patterns remain obscured by our inability to extract them from
such variable data.
At the other extreme of response variables we have
investigated the use of community- 1eve1 indices to describe
patterns in the field and microcosms. Total numbers of
macrofauna, species richness, species diversity and evenness
parameters have been reported for all treatments in each test in
earlier reports. Some generalizations are possible. Figures 3-8
show mean total macrofaunal abundance and species richness values
in control treatments for the six concurrent experiments
conducted between 1982 and 1985 in both estuaries. In both the
Apalachicola Bay and York River experiments mean total abundance
of macrofauna in the microcosms was consistently a poor model of
field abundances. Two problems occur which lead to this lack of
concurrence. (1) Some animals recruit from within the microcosm
where in the absence of epibenthic and demersal predators they
experience large population increases which are not seen in the
19
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field. This occurred in Florida in the spring 1982 test (Fig.
3), the fall 1982 test (Fig. 4) and the fall 1985 test (Fig. 8).
In Virginia this situation was observed at the beginning of the
tests in spring 1983 (Fig. 5) and spring 1985 (Fig. 7). (2) In
other tests recruitment into field sites by species which lack
the ability to recruit from within the microcosms resulted in
increases in field abundances which were not tracked by the
laboratory assemblages (Florida: spring 1983, Fig. 5; fall 1983,
Fig. 6; spring 1985, Fig. 7; Virginia: fall 1982, Fig. 4; fall
1983, Fig. 6; fall 1985, Fig. 8). These problems make total
macrofaunal abundance a poor statistic for tracking natural
communities with laboratory models and a poor indicator of
response to a toxin (Figs. 12 & 14).
Species richness values in laboratory and field controls
were more often similar (Figs. 3-8). In most tests species
richness in the laboratory controls was not significantly
different from the field controls or the pattern of change was
similar. Good examples of this latter phenomenon can be seen in
the Apalachicola Bay data from spring 1983 to spring 1985 (Figs.
5-7). In a few instances there were exceptions to this patterns
of concurrence; in spring 1982 species richness at week 5 had
diverged between the York River site and the microcosms (Fig. 3)
and field recruitment during the spring and fall of 1983 in the
York River led to changes in species richness which were not
reflected in the laboratory. In general, however, we find
species richness to be a fairly conservative community descriptor
which shows few laboratory artifacts. In addition species
richness showed dose-specific responses to PCP treatment (Figs.
13 & 15).
Between these two extremes of species-specific and
community- 1 eve 1 responses, we have investigated a number of
approaches to summarizing individual species data without
obscuring much of the relevant within community response.
Categorization of species into higher taxonomic groupings is the
most straightforward approach and it has the advantage, if
successful, of alleviating the need for detailed species- leve 1
taxonomy in impact assessment. However, we find that very often
individual species within a given taxon do not show similar
patterns of concurrence between the laboratory and field. For
instance, the pattern of abundance of Streblospio b ened ic t i (a
spionid polychaete) in the fall 1982 experiments in the York
River was more similar to that of Scoloplos spp. (an orbiniid
polychaete) than it was to the confamilial Polydora 1 i g n i . a
pattern largely set by recruitment events occurring only in the
field. In later experiments P. 1 i g n i has been observed to
recruit into the microcosms.
4 posteriori methods of grouping species have been attempted
using cluster techniques (Diaz et al. 1984). These techniques
can identify species groups which have similar abundances in the
laboratory and field and groups which do not. Groups of the
latter type can then be ignored when attempting to assess toxic
impacts. A disadvantage of this approach is that it is entirely
a. posteriori and requires substantial experimentation for every
20
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test. Moreover, we are posing questions regarding temporal
patterns, not absolute abundances, so methods which group species
by abundances are inappropriate. A more desirable approach would
be to identify species groups which show similar patterns in the
laboratory and field, and to do so & priori based upon their
ecologies. Responses to stress should then be observed only in
those groups found to be good laboratory models.
The guild approach to classifying species outlined earlier
in this report is our attempt at such an a priori categorization.
Figure 9 shows some composite values of abundances through all
tests for nine of the numerically dominant guilds in the
Apalachicola Bay and York River systems. We caution that since
these figures are composites from all tests that they should not
be taken as actual time courses of abundances, they merely serve
as a convenient way to summarize a lot of data. The patterns in
these figures discussed below are also evident in each of the
individual tests. These plots show only abundances in field and
laboratory controls and their intent is to identify those guilds
for which laboratory assemblages are good models of the field.
The nine guilds represented in Figure 9 are those which
comprise the five most abundant in each of the tests in Florida
and Virginia (Table 5); they therefore include the majority of
individuals collected. Of the nine guilds shown we interpret
five of them as generally showing concurrence between laboratory
and field abundance patterns (Table 6). Mobile burrowing
pedators/omnivores with limited dispersal (Fig. 9, p. 56)
generally showed good agreement between the microcosm and field
in Virginia, but were present in only very low numbers in
Florida. Mobile epifauna which were detritivorous/ omnivorous
scavengers with limited dispersal were again more abundant in
Virginia but appear to be adequately modelled by both of our
laboratory systems (Fig. 9, p. 57). For this guild the absolute
abundances between the laboratory and field often differed, but
the patterns were similar. Mobile burrowing, detrivivorous /
omnivorous, deposit-feeders with wide dispersal were always among
the dominant guilds at each site (Table 5) and generally were
well modelled in the laboratory through the first 5 weeks (Fig.
9, p. 58). Detritivorous/omnivorous, mobile tube-builders which
feed at the sed ime n t-water interface and have limited dispersal
also showed good general agreement between laboratory and field
populations (Fig. 9, p. 59). Recruitment peaks for this guild
were not always of equal intensity between the laboratory and
field but similar patterns were evident. Detritivorous/
omnivorous, mobile burrowers which feed at the interface and have
limited dispersal had similar abundance patterns in the
laboratory and field (Fig = 9, p. 60).
Four other common guilds [(1) d e tritivorous / omnivorous,
mobile burrowing deposit-feeders with limited dispersal, (2)
detri t ivo rous/omnivorous, mobile tube-builders which feed at the
sediment-water interface and have wide dispersal, (3) mobile
burrowing, herbivorous suspension-feeders with wide dispersal,
and (4) mobile-burrowing predators/carnivores with wide
dispersal; Fig. 9, pp. 61-64] did not show good concurrence
21
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between the laboratory and field. The general pattern among
these four guilds was that the guild with limited dispersal
sometimes underwent population blooms in the laboratory, while
the guilds with wide dispersal had recruitment peaks in the field
which were not reflected in the laboratory. These problems with
these guilds did not occur in every experiment, but were present
frequently enough to limit their utility as laboratory models of
field populations.
We argue that only those components of macrobenthic
communities which are modelled well in the laboratory should be
used to assess toxic impact. Based upon the forgoing
consideration of response variables, species richness and the
numerical abundance of the guilds listed in Table 6 appear to be
the most appropriate components in our systems. In the following
section we therefore emphasize these components in our discussion
of predicting field impact from microcosm tests. This is a
conservative approach and we note that among those guilds we have
termed as inadequately modelled in the laboratory are some which
responded well in some tests but not in others. For instance, in
the fall 1982 test in Florida mobi1e-burrowi ng predators/
carnivores with wide dispersal showed good agreement between
numbers in the laboratory and field controls throughout the
experiment, but divergence between microcosm and field patterns
in other tests (Fig. 9, p. 64) caused us to reject this group as
a good laboratory model. In practice it may be that our
procedure of identifying guilds a. priori is best used to flag
species groups which are suspect in their concordance between
laboratory and field; the response of these guilds in undosed
treatments could be examined a, posteriori to make decisions
concerning their utility in predicting impacts of toxic stress.
A limitation to this approach as we employ it here is the
lack of truly objective criteria for assessing differences in
response patterns. As we pointed out above the issue here is how
well temporal patterns of abundance in the laboratory model those
in the field. (e.g., As one declines does the other decline?)
This question is not amenable to answering with ANOVA or
clustering techniques. Both of these techniques are dependent
upon actual abundances rather than temporal patterns.
Specialized non-parametric pattern analysis techniques may prove
useful in the future for providing objective criteria.
PREDICTING RESPONSE TO TOXIC STRESS
Based upon the arguments made above we examined the response
of species richness and the numerical abundances of the guilds
listed in Table 6 to address the question, can the response of
natural communities to a toxic stress be predicted from the
response in the laboratory? The response of species richness to
PCP dosing is shown in Figs. 13 & 15. The responses of the
guilds listed in Table 6 are shown in Figs. 16 & 17. Table 7
summarizes the concordance between the laboratory and field
observations. Since dose equivalency between the microcosms and
field was not achieved in the Virginia spring 1985 experiment,
22
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the observations from that test are omitted. From the
information in Table 7 it is clear that the microcosm results
provided reliable predictions of the response of the natural
communities for those components listed. Moreover these results
show that the response in a microcosm experiment at one location
is frequently a good indicator of response at the other location.
This result, however, is tempered by the fact that differences in
recruitment times between locations may lead to discrepancies in
response s.
The results of Table 7 are promising. In all but one case
(for which sufficient numbers were present) the response to PGP
treatment in the laboratory served as a good indicator of the
field response. Our approach is a conservative one; by including
only those components of the community we know to be well
modelled in the laboratory, we virtually assure that the
responses observed are related to the PCP treatment.
The findings of this study suggest that properly conducted
multi-species tests with estuarine benthos may yield valuable
information regarding responses of natural communities to an
iduces stress, provided that sufficient knowledge of the ecology
or the orgaisms is available and incorporated into evaluating the
results.
23
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REFERENCES
Cairns, J., jr. 1983. Are single species toxicity tests alone
adequate for estimating hazard? Hydrobiologia 100:47-57.
Cairns, J., Jr. (ed.) 1985 Multispecies Toxicity Testing.
Pergamon Press, New York, 253 pp.
Cairns, J., Jr. 1986a. The myth of the most sensitive species.
BioScience 36:670-672.
Cairns, J., Jr. 1986b. What is meant by validation of predictions
based on laboratory toxicity tests? Hydrobiologia 137:271-
278.
Diaz, R. J. 1984. Short term dynamics of the dominant annelids in
a polyhaline temperate estuary. Hydrobiologia 115:153-158.
Diaz, R. J., M. W. Luckenbach, F. Batchelor and S. Morgan. 1986.
Response of macrobenthic laboratory and field communities to
PCP. Unpublished report. U.S. EPA, Gulf Breeze, Florida.
Diaz, R. J., S. C. Thornton, and M. H. Roberts, Jr. 1984. Field
validation of a laboratory derived aquatic test system.
Unpublished report. U.S. EPA, Gulf Breeze, Florida.
Dugan, P. J. and R. J. Livingston. 1982. Long-term variation in
macroinvertebrate assemblages in Alalachee Bay, FLorida.
Estuar. Coast. Shelf Sci. 14:391-403.
Fauchald, K. and P. A. Jumars. 1979. The diet of worms: a study
of polychaete feeding guilds. Oceanogr. Mar. Biol. Ann. Rev-
17:193-284.
Kimball, K. D. and S. A. Levin. 1985. Limitations of laboratory
bioassays: the need for ecosystern-leve1 testing. BioScience
35:165-171 .
Levin, L. A. 1984. Multiple patterns of development in
Streblospio bened ic t i Webster (Spionidae) from three coasts
of North America. Biol. Bull. 166:494-508.
Livingston, R. J., R. J • Diaz and D. C. White. 1985a. Field
validation of 1 ab o r a t o r y-d e r ived tnu 11 i spec ie s aquatic test
systems. EPA/600/4-85/039, U.S. EPA, Environmental Research
Laboratory, Gulf Breeze, FL.
Livingston, R. J., C. C. Koenig and L. E. Wolfe. 1985b.
Preliminary analysis of pentach 1 orop he no 1 in field and
laboratory samples taken during the spring experiment
(1985). Unpublished report. U.S. EPA, Gulf Breeze, FL.
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Livingston, R. J. and L. E. Wolfe. 1985c. Experimental analysis
of the relative impact of sampling disturbance on field
treat-ments and laboratory microcosms (Spring, 1985).
Unpublished report. U.S. EPA, Gulf Breeze, FL.
Livingston, R. J., L. E. Wolfe, C. C. Koenig and G. Ray. 1985d.
Analysis of the 1 aboratory-fie Id response to POP (Spring,
1985). Unpublished report, U.S. EPA, Gulf Breeze, FL.
Livingston, R. J., L. E. Wolfe, C. C. Koenig and G. Ray. 1986.
Analysis of the 1 aboratory-fieId response to PCP (Fall,
1985). Unpublished report, U.S. EPA, Gulf Breeze, FL.
Lu, M. Z. 1982. Organic compound levels in a sediment core from
the Elizabeth River of Virginia. MS Thesis, College of
William and Mary, Wi11iamsburg, VA pp. 157.
Virnstein, R. W. 1977. The importance of predation by crabs and
fishes on benthic infauna in Chesapeake Bay. Ecology 58:
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BioScience 35:330.
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TABLE 1. GENERALIZED PROTOCOL FOR LABORATORY MICROCOSM/FIELD
VALIDATION STUDIES
f\
I. Laboratory microcosms (0.1-1.0 m )
A. Physical/chemical data
1. temperature ( C)
2. salinity (% )
3. dissolved oxygen (ppm)
4. pH
5. sediment % organics
6. sediment grain size
7. sediment temperature, salinity, Eh
B. Infaunal macroinvertebrates (500- and 250- sieves)
1. repetitive cores (3 replicates, 1-3 treatments)
2. vertical distribution (2-cm intervals)
3. azoic sediment samples (500- and 250- sieves)
C. Microbes
1. repetitive cores (3 replicates, 1-3 treatments) (Florida
only)
II. Field
A. Treatments (3 replicates)
1. unscreened platforms
2. screened platforms (exclusion cages)
3. screened platforms (predator-inclusion cages)
4. weekly core samples (no platform)
5. additional treatments (specific for individual experiments)
B. Physical/chemical data (same as I.A.)
C. Infaunal macroinvertebrates (same as I.E.)
D. Microbes (same as I.C.)
III. Variables analyzed
A. Infaunal macroinvertebrates, epibenthic organisms
1. numerical abundance (total and dominant species)
2. ash-free dry weight biomass (total and dominant species)
3. species richness
4. species diversity and evenness indices
5. functional group associations
6. numerical response of guilds
B. Microbes
1. total biomass
2. bacteria
3. photosynthetic microbes
4. microeukaryotes
5. bacterial ecotype
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TABLE 2. SAMPLING SCHEDULES FOR THE COMBINED (FSU-VIMS)
EXPERIMENTAL PROGRAM (1981-1985)
I. Weekly samples
A. FSU
1. oligohaline stations (11/24/81-11/17/83)
2. polyhaline station (11/25/81-3/15/84)
B. VIMS
1. polyhaline marine lab station (10/13/79-12/18/83)
II. Microbiological data
A. FSU
1. oligohaline stations (fall 1982; spring 1983)
2. polyhaline stations (spring 1982)
B. VIMS
1. marine lab station (spring 1982)
III. Combined (field-laboratory) experiments
A. Spring 1982
1. Florida
2. Virginia
B. Fall 1982
1. Florida
2. Virginia
C. Spring 1983
1. Florida
2. Virginia
D. Fall 1983
1. Florida
2. Virginia
3. Treatments included:
a. Field controls
b. Field predator exclusion cages
c. Field predator inclusion cages
d. Microcosm controls
e. Field and lab treatments dosed with PCP
E. Spring 1984
1. Virginia only
F. Spring 1985
1. Florida (station ML)
2. Virginia
3. Treatments included:
a. field controls
b. microcosm controls
c. replicate lab and field treatments dosed with PCP
d. azoic sediments
G. Fall 1985
1. Florida (station ML)
2. Virginia
3. Treatments as in F.3.
-------
Table 3a - I'unrUonal Group Assignment? for Taxa Collect od in Florida
TAXON
TROPHIC MODE/
LEVEL
DISPERSAL MOBILITY
MODE MODE
GAttwius
ADELODRILUS SP.
AREN1COLA CRISTATA
AR1CIDEA FRAG 1L IS
AR1CIDEA JEFFERSII
ARICIDEA PHIL8INAE
ARICIDEA SP.
ARICIDEA SP. A
ARICIDEA TAILOR!
ARICIOEA HASS1
CTENODRILUS SERRAIUS
DASTBRANCHUS SP.
ENCHITRAEUS ALB1DUS
EMCHTTRAEUS SP.
HAEMONAIS HALDVOGELI
HAPLOSCOLOFIOS FOLIOSUS
HAPLOSCOLOPLOS FRA61LIS
HAPLOSCaOPLOS ROKJS1US
IMMAT TUBIFICID K/0 CAP SEIAE
L1MNOPRILOIDES SP.
LIMNODRILOIDES HINOXLMANNI
MONOCULOtCS SP. (Cf NtEI)
MONOP1LEPHORUS IRRORATUS
MONOPTLEPHORUS FARVUS
MONOFtLEFHORUS SP.
NAINERIS SEIOSA
NA1S COMMUNIS
fMIS aiNGUlS
OLI60CHAETA
ORBIN1A RISERI
FAR6MAIS LIIORALIS
FARAONIS FULGENS
PHALLOPRILUS MEPIOFORUS
FHALLOPRILUS MONOSFERttATHECUS
PHALLOPRILUS SF.
SCOLOftOS RtlfRfl
SfllTHSONIWILUS MflftlNUS
SIfLflRlfl LflCUSIRIS
TUPIFE* LIIORflLiS
TUPIFICOIDES BEWDENI
TUP1FIC01DES 6ABRIELLAE
TIJBIFICOIKS HEIETOWEIUS
TUPIFICOIOES PSEUDOGflSIER
TUPIFICOIDES SP.
TUPIFICOIDES SHIRENCOMI
CAECUM SP.
nUCOSfl
SARSI
M/HPAHIDftE
DEPOSIT FEEDER -«RBIVORE
-LIMITED DISPERS-BURftWinOBlLEI
DEPOSIT FEEDER -Otll/DETRIT
DEPOSIT FEEDER -OWI/DETRIT
DEFtJSIT FEEDER -«t(l/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OWl/DETRIT
DEPOSIT FEEDER -OHNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -«t(l/DETRIT
DEFflSIT FEEDER -OWI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEFflSIT FEEDER -OWI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEFtKIT FEEDER -OHMI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OHMI/DETRIT
DEPOSIT FEEDER -OMNI/PETRIT
DEFDSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEF-OSIT FEEDER -OnNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OWI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMMI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
KFtJSIT FEEDER -OMNI/DETRIT
DEFDSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
PEFOSIT FEEDER -OMK1/DETRIT
DEFtKIT FEEDER -OMNI/DETRIT
KPOSIT FEEDER -OMNI/KTRIT
[lEFOSIT FEEKR -OMNI/DETRIT
[lEFOSIT FEEKR -OMNI/DETRIT
DEFtKIT FEEDER -OMNI/PETRIT
[lEFQSIT FEEDER -OMNl/ICTRIT
PEFDSIT FEEttR -OMNI/PETRIT
KFDSIT FEEDER -OMMI/PETRIT
DEFtKIT FEEKR -OMNI/DETRIT
DEFtKIT FEEDER -OMNI/DETRIT
-LIMITED DISPERS-BUKROH(MOBILE)
-LIMITED DISPERS-BURROMIMOBILEt
-LIMITED OISPERS-RJRROM(MOBILE)
-IIMITED DISPERS-BURRW (MOBILE)
-LIMITED DISPERS-BURROW(MOBILE)
-LIMITED DISftRS-BWiROM(HOBlLE)
-LIMITED DISFERS-BURRWIHOBILEt
-LIMITED DISPERS-BURROH(MOBILE)
-LIMITED DISFERS-HJRROKIMOBILEI
-LIMITED DISPERS-BURDOK(MOBILE)
-LIMITED DlSfERS-BURROMIMOBILE)
-LIMITED DISPERS-BURRWIHOBILEI
-LIMITED DISftRS-BUh«W(MOBILE)
-LIMITED DISPERS-BURRWIMOBILEI
-LIMITED DISFERS-BURRW(MOBILE)
-LIMITED DISFERS-BURRW (MOBILE)
-LIMITED DISFERS-PtJRROM(MOBILE)
-LIMITED DISFERS-BURROM(MOBILE)
-LIMITED DISFERS-B"JRROK(MOBILE)
-LIMITED D1SFERS-BURROW (MOBILE)
-LIMITED DISFERS-WRRW(MOBILE)
-LIMITED DISPERS-BURROHIMOBILE)
-LIMITED D1SFEPS-BURWWIMOBILE)
-LIMITED DISPERS-BURROK(MOBILE)
-LIMITED DISFERS-BURROM(MOBILE)
-LIMITED DISPERS-BURROHIMOBILE)
-LIMITED DISFERS-BtKROHIMOPILE)
-LIMITED D I SPERS-BURRW (MOBILE)
-LIMITED DISPERS-PURROHIMOPILE)
-LIMITED PISPERS-BURROMIMOBILE)
-LIMITED DISFWS-HJWOH(MOBILE)
-LIMITED DISPERS-PliRROM(MOBILE)
-LIMITED DISFERS-fUM^iMOPILE)
-LIMITED DISPERS-PUWW(MOBILE)
-LIMITED DISFEftS-PURF-OWIMOBILEI
-LIMITED DISFEfS-piftRWIMOPILEI
-LIMITED DISfEW-WjSfiOHlMOBlLEI
-LIMITED DISFERS-BISROMIMOPILE)
-LIMITED PISFEftS-P.RROMIMOBILEI
-LIMITED BISFEKS-BIJKPOMIMOBILE)
-LIMITED PISFEf-3-PiJSROMIMOBILE)
-LIMITED DISFERS-BUMW(MOBILE)
-LIMITED DISFERS-PI)WW(MOBILE)
-LIMITED DISPERS-BURRW(MOBILE)
DEFtKIT FEEDER -OMNI/DETRIT -LIMITED DISFERS-EFIFAUIMOPILEI
DEPOSIT FEEKR -OttNI/rtTRIT
KFOSIT FEEltR -OMNI/PETRIT
DEPOSIT FEEDER -OMNI/KIRIT
KMKI1 FtfWR -l)l»i|/Lt If-II
-LIMITED PISfEKS-TllPEISEBSILEI
LIM1TEP ['ISmS-TUFEISESSIlEI
-LIMITED DISFf'r.-TWEiSESSILE)
?8
-------
Tablp 3a
TAXON
CAPITEUA CAP IT AT A
CAPITELLA JQNESI
ARMANDIA A6ILIS
COSSURA SQTERI
HOLOTHUROID1A
MEDIOMASTUS AH8ISETA
NOTOMASTUS HEHIPDDUS
NOTOMASTUS LATERICEUS
PARANAITIS SPECIOSA
SIPUNCULA
OPH1UR01DEA
Cl SIENA GOULD I
OHEMA FUSIFORMIS
CHAETOZOtC SP.
CIRRATULIDAE
THARTX SP.
TROPHIC MODE/
LEVEL
KFDSIT FEEDER -OMNI/DETRIT
ttPOSIT FEEDER -OW1/DETRIT
DEFOSII FEEDER -OMNI/DETRIT
DEFDSIT FEEDER -OMWI/OETR1T
KPOSIT FEEDER -OMNI/DETRIT
DEPOSIT FEEDER -CTWI/DETRIT
DEPOSIT FEEDER -OMN1/KTRIT
D€POS1T FEEDER -OMNl/DETRIT
DEPOSIT FEEDER -OHI/DETR1T
DEPOSIT FEEDER -OWI/DETRIT
DEPOSIT FEEDER -OMW/DETRIT
DEPOSIT FEEDER -OMNI/DETRIT
DEFDSIT FEEDER -OMW/DETRIT
INTERFACE FEED -OWI/DETRIT
INTERFACE FEED -OHI/DETRIT
INTERFACE FEED -Omi/DETRIT
DISPERSAL MOBILITY
MODE MODE
-VARIABLE DlSPER-PURflOMinOSILEI
-VARIAH.E DISPER-BURftOM (MOBILE!
-HIDE DISPERSAL -BURROW (M081 LEI
-HIDE DISFERSAL - BURROW IfCeiLEI
-«IDE DISPERSAL -BURROMIM08ILE)
-MIDC DISPERSAL -PURROMtnOBILE)
-MIDE DISPERSAL -WJRROWIMOBILEI
-WIDE DISPERSAL -PURROM(nOBILE)
-MIPE DISFtRSAL -PURROMinOBIbE)
-HIDE DISPERSAL -BURROHINttlLE)
-HIDE DISPERSAL -EPlFAUinOBILE)
-HIDE DISPERSAL -TUBE(MOBILE)
-HIDE DISF-ERSAL -TUBE (SESSILE)
-LIMPED DISPERS-PUKROHIMOBILEI
-LIMITED DISFtRS-PLKROHinOBILEI
-LIMITED DISFERS-BURROH(MOBILE)
ER1CHSOTCLLA (CF FIL1FORHIS)
LEMBOS SMITHI
LEMBOSSP. I
AMPELISCA VADORUH
AHFELISCA VERR1UI
AMPHARETIDAE
CARAZZiaLA HOBSONAE
COPOFHIUn LOUISIANltl
COROPHIUH TUBEROJLATUM
ERICHTHONIUS BRASILIENSIS
HOPSONIA aORIDA
MELINNA MAOJLATA
INTERFACE FEED -OWI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INfERFACE FEED -OMNI/DETRIT
-LIMITED DISFtRS-EFIFAU(HOBILE)
-LIMITED DISFERS-EPIFAU(MOeiLE>
-LIMITED DISPERS-EPIFAU(HOBILE)
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED -OMNI/PETR1T
INTERFACE FEED -OMNI/DETRIT
-LIMITED DISfERS-TUK (MOBILE)
-LIMITED DISPERS-TUBEIMOB1LE)
-LIMITED BISTERS- TUBE (MOBILE)
-LIMITED DISPERS-TUBEIMOBILE)
-LIMITED BISPERS-TUBEIMOBILEI
-LIMITED DISPERS-TUBE(MOBILE)
-LIMITED DISFERS-TUBE (MOBILE I
-LIMITED PISPERS-TUBE (MOBILE)
-LIMITED DISFERS-TUK(MOBILE)
CIRRIFORfllA TENTACULATA
FOLTPORA LIGNI
rartflRfl SOCIALIS
FOL>[flRA SP.
SFIO PETTIBONEAE
STREBLOSPIO BENEDICT!
PflRNEA TRUNCATA
TELLINA TEXANA
Af OPRIONOSPIO PTGttAEA
LOIMIA MEDUSA
MftGELONA PETT1PONEAE
MIIWSPIO FERKINSI
FftRflrRIONOSFlO PIWWTA
JOUSTONI
INTERFACE FEED -OMNI/DETRIT -VARIABLE DISPER-RURROW(MOBILE)
INTERFACE FEED
INTERFACE FEED
INTERFACE FEEP
INTERFACE FEED
INIERFACE FEED
-OMNI/PETRIT
-OHNI/DETRIT
-OMNI/DETRIT
-OMNI/PETRIT
-OMNI/DETRIT
INTERFACE FEED -OMNI/T.CTR1T
INTERFACE FEED -OMNI/DETRIT
INTERFACE FEED
INTERFACE FEED
INTERFACE FEED
INTERFACE FEED
INIERFACE FEED
IIIIERFACE FEED
INTERFACE FF.EP
-OMNI/DETRIT
-OMNI/PETRIT
-OMNI/DETRIT
-OTHI/PETRIT
-OMNI/DETRIT
-OMNI/PETPIT
-UNIII/PFTRIT
-VARIARE DISFER-TUBEIMOBILE)
-VARIABLE DISFER-TUK (MOBILE)
-VARIABLE DISFER-IUBElMOBILE)
-VARIAFAE DISfER-TUKIMOPlLE)
-VARIABLE DISFER-TUBEIMOBILEI
-WIPE PI9FERSAL -BUfiROWiroPILE)
-WIDE DISFERSAL -BURROHIMOBILE)
SlAWVMA
-WIPE PISFtRSAL
-WIPE PISFtRSAL
-WIPE DISPERSAL
-WIPE DISFERSAL
-WIPE PISFERSAL
-Wlft PISFFRSAL
-wilt PISffr.r-nL
-Kill IHSfFhV.I
-TUBE(MOBILE)
-TUKIMOBILEI
-TUBE(MPBILE)
-TUPE(MOBILE)
-IUPF(MOPILE)
-TIIPEIMOBILEI
-lH[f iMfiplifi
-Hi|f mul'ILE)
-------
3a
TAXON
scaaEPis IEXANA
SPIOPHANES BOMBU
PHOROWDA
ARABELLA SP.
AUTOL1TUS SP.
DORVILLEA SP.
EHLERSIA SP.
LUHWINERIS LATREILLI
HARFWSA SANGUINEA
HICROPnmLUS SP.
OPHIODROMUS ABSCURA
PETTIBONEIA SP.
PSEUTJOSTLLIDES CWACOENS1S
SCHISTOMERINGOS RUDOLPH!
STLL1S CORNUTA
CAFRELLA PENANTIS
CAFflElUOAE
LUCONACIA INCERTA
HELONGENA CORONA
TANAIDACEA
AMERICONUPHIS MA6NA
DIOPATRA CUPREA
ONUPHIS ERENITA OCULATA
POLLIHICES DUPLICATUS
AMFHINOME ROSTRATA
GRUKULEPIS SP.
HARMOIHOE SP.
IEPIDON01US SUBLEVIS
FflLTNOIPAE
S1GALIONIDAE
AGLAOFHAMUS VERR1LLI
ANCIS1ROSTLL1S HAfilMANrtE
WICISIROSILLIS FAP1LLOSA
CflPIRA INCERfA
ETEOIIE HE1EROPODA
ETEOHE LAC1EA
EUMIPA SWK5UINEA
GLTCERA AMERICANA
GUCINPE SOLIIARIA
GON1ADIDAE
GlPflS BREVIFALFA
GIFTIS VIITAIA
NEFHITS PUCERA
NEFWTS INCISA
HEFHTIS FICIA
FABAHESIWJE LUIEOLA
rflR/VIPflLIA AMERIC/VIA
nuilOtlltE AREIW
Mmuiiix:innE
TROPHIC MODE/
LEVEL
INTERFACE FEED -OMNI/DEtRlI
INTERFACE FEED -OIHl/DETRIT
INTERFACE FEED -OtU/DETRIT
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR ^WNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
FFtDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PREDATOR -CARNIVORE
PttDATOR -CARNIVORE
FREPATOR -CARNIVORE
FKEDATOR -CARNIVORE
FREDA10R -CARNIVORE
FREMTOR -CARNIVORE
FREt'ATOR -CARNIVORE
FttPATOR -CARNIVORE
FREPA10R -CARNIVORE
FFEWIOR -CARNIVORE
FREPAIOR -CARNIVORE
FfEPAlOR -CARNIVORE
F-REPA10R -CARNIVORE
FREPATOR -CARNIVORE
FREPAIOR -CARNIVORE
FREPATOR -CARNIVORE
FREPATOR -CARNIVORE
FREWtOR -CARNIVORE
FREDATOR -CARNIVORE
FREPATOR -CARNIVORE
FREWIIOR -CARNIVORE
r&ECAIOR -CARNIVORE
FWW10R -CARNI'flJftE
DISPERSAL MOBILITY
MODE MODE
-HIDE DISPERSAL -TUPE (MOBILE!
-NIDE DISPERSAL -TUBEinOBILEI
HIIDE DISPERSAL -TUEE(SESSILE)
-LIMITED DISPERS-BURROH(MOBILE)
-LIMITED DISPERS-BURRO«(f10BILEI
-LIMITED DISPERS-BURROM(MOBILE)
-LIMITED DISPERS-BURROM(MOBILE)
-LIMITED DISPERS-BURROM (MOBILE)
-LIMITED DISPERS-BUV)OH(N)BILE)
-LIMITED DISFtRS-BURRON(MOBILE)
-LIMITED DISPERS-BURROH(MOBILE)
-LIMITED DISFERS-BURROM(MOBILE)
-LIMITED DISFERS-R)RROM(MOBILE)
-LIMITED DISPERS-BURROMIM08ILEI
-LIMITED DISFERS-BURROMIMOBILEI
-UNITED DISPERS-EPIFAUIMOBILEI
-LIMITED DISFERS-EFIFAUIMOBILE)
-LIMITED DISFERS-EPIFAUIMOBILEI
-LIMITED DISFERS-EPIFAU(MOBILE)
-LIMITED DISFERS-EPIFAUIMOBILE)
-LIMITED DISPERS-TUBE(SESSILE)
-LIMITED DISFERS-TUK(SESSILE)
-LIMITED D1SPERS-TUEE (SESSILE)
-LIMITED DISF-ERS-UWCHN
-VARIABLE DISF-ER-BURRWifCPILE)
-VARIABLE DISFER-BURMNtHOBILE)
-VARIABLE DISFER-BURROM(MOPILE)
-VARIABLE PISfER-FjW?ON (MOBILE)
-VARIABLE PISFER-BURROH(MOPILE)
-VARIABLE BISFER-BUKROW(MOB1LEI
-WIPE DISF'ERSAL -RIRROMIMOPILE)
-HK€ PISFERSAL -BUfRtW(MOPILE)
-HIDE PI?ERSAL -BURROW! MOBILE I
-NIDE PISFERSAL -K*flOWIflOBILE)
-MIK DISFERSAL -PUF.'F,-OW(nOPILE)
-Mire PISFER-SAL -PUF-DOMIMOPILEI
-M1PE PISFERSAL -PUF,W«(MOPILE)
-MIC? PISFEftSAL -PIJF.ROHIMOBILE)
-MIK PISFERSAL -PURRONIMOPILEI
-Mire PISFERSAL -PUKROW(MOPILE)
-Mire PISFERSAL -PURROHI MOBILE)
-MU€ PISFERSAL -MJRROMIMOPILEI
-MIK DISFERSAL -PURftOMIMOPILEI
-MU€ DISFERSAL -PURROM(MQPILE)
-Mire PISPERSAL -PUFM)W(MOPILE)
-Mire DISFERSAL -WJRROMinOPllE)
-HIPE PISFfF-SAL -EnJS^OMIMOPlLEI
-MIPE PISFERSAL -EWOUimpHE)
-Milt ri5IFF-"fH. -FDf.PUUIUJMlEI
-------
'ront'cH
TAXON
FHtLLOOOCIDAE SP. 2
SIGAflBRA BASSI
SIGAHBRA TENTAOJLATA
OJTVCEA
LAEOMEREIS CU.VERI
NERE1DAE SP.
NEREIS FALSA
NEREIS HICROIttt
MEBSTERNEREIS TRIDENTATA
NEREIS SUCCINEA
PTCN060NIDA
ANTHJRIPAE
APANTHURA tlAGNlFICA
BATEA CATHARINENSIS
BRANIA OAVATA
BRANIA HELLFLEETEMSIS
CASSIDINIDEA OVALIS
CASSIDINIDEA SP.
CIATHJRA PaiTA
EJOGONE D1SPAR
LISTRiaifl 8ARNARDI
nONOCUODES SP. 2
HIMM RETNOLDSl
OOONTOSTLLIS ENLOFA
SPHAEROSJLLIS TATLORI
SYNCHELIOlUn AHERICANUH
XENANTHURA BREVITELSON
EDOIEA SP. (CF HONIOSA)
ELASHOPUS LEV IS
GRANTIOIERaLA BONNIEROIPES
LEMPOS SP.
LEUCOIHOE SFINICAfiFA
LTSIflNOf^lS ALW>
HELIIA flfPENDICafllfl
HELITA ELttJGAIA
MEIITA SF.
MICRODEUTOFUS HfltCOCKI
niCRODEUIOFliS mERSI
NASSARIUS V1BEX
AMPITHOE SP.
CERflFUS SP. (CF lUBaflRIS)
CtMADUSA COMFTA
HICROFPOTOF1JS RAHEI1
FHOT1S MACRWWNUS
CHIRONOHIME
FtAlIllERf IS Wit Ml I
TROPHIC MODE/
LEVEL
FREDAIOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENFjER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENFJER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENFJER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCflVENGER
SCWNGER
-CARNIVORE
-CARNIVORE
-CARNIVORE
•CARNIVORE
-OHI/DETRIT
-WN1/DETRIT
-flmi/ttTRIT
-Omi/DETRIT
-
-ilHITED DISPERS-BURROH(MOBILE)
-LIHITED DISPERS-BURROHIMOBILEI
-LIMITED DISPERS-BURROMIMOBILE)
-LIMITED DISFERS-BURROHIHOBILEI
-LIHITED DISFERS-BURROHlHOBILEt
-LIMITED DISFERS-BURROH(HOBILE)
H.IM1TED DISFERS-BURROHIMOBILE)
-LIHITED DISFERS-BURROHIMOBILEI
-LIMITED DISPERS-BURROH(MOBILE>
-LIMITED DISPERS-BURROHIMOBILE)
-LIMITED DlSFERS-EPIFAUIMOfilLEt
-LIMITED DISfERS-EPIFAUIHMILEI
-LIMITED PISfERS-EPlFAUIHPPlLEI
-LIMITED DISFERS-EFIFAU(HOPILE)
-LIMITED DISF'ERS-EFIFAU(MOBILE)
-LIMITED DISFERS-EPIFAUIHOB1LEI
-LIMITED PISfERS-EFIFftlKMGBILEI
-LIMITED [ilSfERS-EPIFAUIHOeiLEI
-LIMITED DISFERS-EFIFAU(HOPILE)
-LIMITED DISFERS-EPIFAU(HOBILE)
-LIMITED DISFERS-EPIFAUIMOBILE)
-LIMITED DISfERS-EPIFAU(MOBILE)
-LIMITED DlSfERS-TUBE(HOBILE)
-LIMITED DISFERS-TUBE(MOBILE)
-LIMITED DISF-ERS-TUBE (MOBILE)
-LIMITED PISFERS- TDK (MOBILE)
-LiniTED PISF£RS-TUBE(MOBILE>
-IHflHOWJ -W»tflHN
-Wilt DISltRSftL -BUWWIWEMLEI
-------
'font
TAXON
CANTHARUS CANCELLARIA
UROSALPINK TAMFAENSIS
OL1VELLA SP.
PRUNUH APICINUM
PtRAmDEUA SP.
fipsEurtssp.
CREPIDULA SP.
OOC DUNERl
MEGALOItIA PIGMENTUM
SABELLIDAE
ABBA AEQUALIS
ANADARA SP.
ANOnALOCARDIA AUBERIANMA
CH10NE CANCEILATA
DENTAL lUn LAOUEATUH
DOSINIA ELEGANS
EMS IS SP.
GLOTTIDIA PTRANIDATA
MACTRA FRAGILIS
MULINIA LATERAL I 5
POLIMESOOA CAROL1NIANA
RANGIA CUNEATA
SEMELE pRaiFicA
TAGaUSSP.
TROPHIC MODE/
LEVEL
SCRAPER -CARNIVORE
SCRAPER -CARNIVORE
SCRAPER -OMNI/DETRIT
SCRAPER -OMNl/PETRIT
SCRAPER -OMNI/DETRIT
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEE1HCRBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-WRBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
DISPERSAL MOBILITY
MODE MODE
-LIMITED DISfERS-EPIFAUIMOBILEI
-LIMITED DISPERS-EPIFAUIMOBILE)
-LIMITED DISFERS-EPIFAUIMOBILE)
-LIMITED BISFERS-EPIFAU(MOBILE)
-LIMITED DISFERS-EPIFAUIMOBILEI
-LIMITED DISPERS-EPIFAU(HOBILE)
-LIMITED DISPERS-EPIFAU(SESSILE)
-LIMITED DISFERS-TUKIMOBILE)
-LIMITED DISFERS-TUBE (SESSILE)
-LIMITED DISFERS-TUBE (SESSILE)
-IW€ DISPERSAL -PURROMIMOBILEI
-HIDE DISPERSAL -PURROHI MOBILE I
-«IPE DISfERSAL -BURROMI MOBILE)
-MIR DISPERSAL -PURRW(MOBILE)
-HIDE DISF-ERSAL -BURROM(MOBILE)
-HIDE DISFCTSAL -BURROM(NOBILE)
-HIDE DISfERSAL -BURROH(MOBILE)
-HIDE DISPERSAL -BURROH(MOBILE)
-HIDE DISFERSAL -BORROH(MOBILE)
-Hire DISPERSAL -BURROH((WILE)
-Hire DISPERSAL -BURROH (MOBILE I
-Hire DISFERSAL -BURROH(MOBILE)
-Hire DISFERSAL -BURROH (MOBILE)
-Hire DISPERSAL -BURROH(MOBILE)
BRACHIDONTES SP.
HTDROIDES UNCINATA
LTONSIA HTALINA
SERFULIDAE SF.
TRACHTCARDlUn EGMONTIAftUM
SFIONIDAE POST-LARVA
LEPIDACTTLUS SP.
MACOMA BALTH1CA
MACOMA MITCHELLI
MACOMA TENTA
CHAETOPTER1PAE
SP10CHAETOFTERUS COSTORW
GASTROPOD SP. 2
ISOPODA
BIVALVE POST-LARVA 1
BIVALVE SP. 2
PIVftLVE SP. 3
BIVALVE SP. 5
SP. S
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-HERBIVORE
SUSPENSION FEED-OMNI/KTRIT
SUSPENSION FEED-OHNI/reiRIT
SUSPENSION FEEP-Omi/PETRIT
SUSPENSION FEED-OHNI/KTRIT
SUSPENSION FEED-OTWI/KTR1T
SUSPENSION FEEO-OMHI/PEIRIT
SUSPENSION FEEP-WVil/reiRIT
-Hire DISFERSAL -EPIFAU(SESSILE)
-Hire DISFERSAL -EPIFAU(SESSILE)
-Hire DISF-ERSAL -EFIFAU(SESSILE)
-HIS DISPERSAL -EPIFAU(SESSILE)
-H|[i£ DISFERSAL -EFIFAU(SESSILE)
-INVALID TO ftS3I-TUBE(SESSlLE)
LIMKED DISFERS-PUF,«OH(MOBILEI
-WIPE PISFEP,SflL -PURROH(MOPILE)
-Wltf PISFERSflL -PUS1» HOHN
-UM NCWH -(f» NOUN
-UHI (Kiel -l»n t«Mi
-lit.1' 111 MI -\it m vi
-ll'll MI"J>I -H'l> Ui M'i
-------
Tab!-- ^a 'Vont'dl
TROPHIC MODE/ DISPERSAL MOBILITY
TAXON LEVEL MODE MODE
IMSECT IffNtf. UNKNOWN MJNKNOWN -UNKNOWN -UNKNOWN
BRANCWOSTOMA CARIMEUN UNkWOHN -UNKNOWN -HIOC DISFtRSflL -WHROM(MOBILE)
NUCULWW flCUIfl UMKNOMN ^JNWO(N HdDE DISftRSflL -BlWWMinOBIlEI
FatCH(€TE (UNIDeNT.I UNKNOWN -UNKNOWN -*IDE OlSftRSAL -BUW)OW(WBILE)
-------
Table 3h - Functional Group Assignments for Taxa Collected in
Virginia
TAXON
PORIFERA
HYDROZOA
ANTHOZOA
TUR8ELLAR1A
POLYCLADIA
RHYNCHOCOELA
TUBULANUS PELLUC1DUS
CAR I NO* I ME
CEREBRATULUS LACTEUS
CEREBRATULUS LUR1DUS
MICRURA
MICRURA LE1DYI
MICRURA RUBRA
MICRURA SP. 1
AMPHIPORUS
AMPHIPORUS SP 1
TETRASTEMMA VERMIOJLUS
ANNELIDA -IPOLYCHAETA)
POLYNOIDAE
LEPIDPHETRIR COMMENSALIS
HARMOTHOE EXTENUATA
LEPJDONOTUS SUBLEVIS
PHYU.ODOCIME
ETEONE
ETEOtC SP A
ETEONE LACTEA
ETEONE HETEROPODA
EUH1DA SAN6UINEA
PARANfllTIS SPEC10SA
PHYaODOCE
PHYLLODOCE ARENAE
EULAL1A SANGU1NEA
MESIONIDAE
AMPHIOUDOS
6YPTIS VITTATA
6YPTIS BREVIPALPA
PARflHESIONE LUTEOLA
N1CROPHTHALMUS SCZELKOWII
SIGAMBRA TENTAOJLATA
SYLLIDAE
BRAN1A CLAVATA
NEREIDAE
NEREIS SUCC1NEA
NEPHTYIDAE
NEPHTYS PICTfi
GLYCERIDAE
GLYCERA
6LYCERA DIBRANCH1ATA
GLYCERA AMERICANA
GON1AD1DAE
6LYC1NDE SOL1TARIA
CAf'ITaLlDAE
CAPITELLA CAPITATA
CAP1TELLH JONES I
HETEROMASTUS FILlFOfffllS
NOTOHASTUS HEH1PODUS
MEDIOMASTUS AHBlSETfl
MALDANIDAE
CLYNENELLA
aYHENELLfl TOROUftTP
CLYNENELLH zoNflLis
STERNASf'IWt
WRAONIS & A
Sf'IONlDflE
POLYOOIW LI6N1
PflRAPRlONOSPIO
PflRftPRIONOSPIO P1NNATA
SO)LEC«.Ef'lDES VIR1DIS
TftUfWIL NOW-.
SUSl-tNSlON
SUSPENSION
SUSPENSION
PREDATOR
PRrD«TOf<
PREDATOR
PREDATOH
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOK
PREDATOR
PREDATOR
UNKNOWN
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
DEPOSIT-FEEDER
DEPOSIT -FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DtKlSlT-FEEDER
DEPOSIT-FEEDER
nepos IT-FEEDER
litf'US IT-FEEDER
DEPOSIT -FEEDER
INTERFACE
INTFRFflCt
lNlERf«C£
INTERFACE
INTERFACE
MOBILITY MODE
£P I FAUNAL -SESSILE
EPIFAUNAL -SESSILE
EPIFAUNAL -SESSILE
BURROUER-flOBlLE
BURROMEH-Kl&lLb
BURROMER-H06ILE
BURROUER-MOBlLt:
BORROWER-MOBILE
BURROMER-MOBILE
BURROUER-HOBILE
BURROUER-HOBILE
bURROWER-HOBlLE
BURROUER-HOBILE
BURROWER-HOB1LE
BURROV€R-HOBILE
BORROWER-MOBILE
BURROUER-MOblLE
ONKNOUN
BORROk'ER-WBlLE
BORROWER-MOBILE
BURROWER-MOBILE
BURROUER-WBILE
BURROUER-MOB1LE
BURROHER-MOBILE
BURROWER-MOBILE
BURROWER-WB1LE
BORROWER-MOBILE
BURROWER-MOBILE
BORROWER-MOBILE
BURROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BURROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BURROUER-MOB1LE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BURROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
BORROWER-MOBILE
TUBE BUILDER-SESSILE
TUBE BOlLDER-SESSlLE
TUBE BUILDER-SESSILE
TUBt BOlLDER-SESSK.E
Bi ifiROWt* -MOBILE
BURROWE^-MOHLE
TUBt BUlLDfcR-«06
TUBE BUILDER-MOB
TUBE BUlLDEfi-MUt
TUBE BOlLD£f(-MOb
TUBE BUlLDER-nOfa
DISPtRSAL MODfc
LIMITED DISPERSAL
LIMITED DISPERSAL
WIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
UNKNOWN
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
HIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
HIDE DISPERSAL
WIDE DISPERSAL
HIDE DISPERSAL
WIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
WIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMPED DISPEhSAi
LlrlTEn DISPERSE
HIL€ Dlbt-tR'^rtL
wl[€ DISPERSAL
WlPfc [ilSrtfcSAL
HluF DISPEKSHL
wlDE DISPERSAL
HIDE DISPERSAL
wltt DISPERSA^
TROPHIC LEVEL
HtRBlVORE
HERBIVORE
HERBIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
UNKNOWN
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMN1VORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
DETRlTlVORE/OWiIVORE
DETRITIVORE/OMNIVOW:
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNji-OHE
DETRITIVORE/OMNIVORE
DETRlTlvORE/OMNlvOht
uETRITIVORE/OMNlVOKt
DETftlllvOftE/OwIvOfe
DETRITIvORE/OMNlVURE
DETRITlVORE/OMMVGRt
DETRlTIVORE/OMNlvOKf
DETR1TIVORE/OMNIVOH!-
DETRITIVORE/OMNIVORE
DtTRIV.VOxt/OMNlvOrtt
-------
Table 3b (cont'd)
TAXON
SCOLELEP1S SOUANATA
SCOLELEPIS TEXANH
SPIO SETQSA
SPIOPHANES BOHBYX
STREBLOSP10 BENEDICTI
D1SP10 UNC1NATA
SPIOCHAETOPTERUS OCULATUS
D10PATRA
D10PATRA CUPREA
ARABEUIDAE
PSEUDEURYTHQE PAUC1 BRANCH 1 AT
ORB1NIIDAE
SCOLOPLOS
ORBINIA ORNATA
SCOLOPLOS FRA6ILIS
SCOLOPLOS ROBUSTUS
SCOLOPLOS RUBRA
CIRRATUL1DAE
CIRRATULUS
THARYX
THARYX SP A
CISTENA BOULDII
AMPHARETIDAE
ASABELL1DES OOJLRTP
NELINNA HAQJLATA
TEREBELLIDAE
AHPHITRITE ORNATA
LOIHIA MEDUSA
PISTA PALKATA
SABRLIDAE
POTAH1LLA fCaeCTfl
SABELLA MICROPHTHALM
HYDROIDES OIANTHUS
TUBIFICIDAE
TUBIFICOIOES SP. 1
PARANA1S LITTORAL1S
6ASTROPOOA
RISS01DAE
SAYELLA
CREPIDULA FORN1CATA
CREP1DULA CONVEXA
UROSALP1NX CINEREA
NASSARIUS V1BEX
ILYANftSSA OBSOLEIA
NAN6ELIA PLICOSA
PROPEBELA PYGHAEA
ACTEON PUNCTOSTRIATUS
CYLICHNA ALBA
HAMINOEA SOLlTARIP
ACTEOCINA CflNALl COPTS
RETUSA OBTUSA
ODOSTOHIA
OOOSTOM1A BJSUTURALIS
TURBONILLfl INTERRUPTA
A€OL1DI1DA£
PELECYPODA
YOLD1A L1HATULA
ANADARA TRANSVERSE
MYTILUS EDULIS
GEUKENSIA DEH1SSA
LUC1NA NULTIL1NEATA
ALiGENfi ELEVATP
VENERIWtf
HtRCENrtRIfl MtRCENHfilh
bt«fW GEWW
1ELL1N1IWE
TELLlNfl M61L1S
WtCOUft
HACOHfl BflLTHICfl
HACOHA TENIP
ip&aus
TROP'HIL «OD£
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
SUSPENSION
PREDATOR
PREDATOR
PREDATOR
PREDATOR
DEPOSIT-FEEDER
DEPOSlT-fEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
INTERFACE
INTERFACE
INTERFACE
INTERFACE
DEPOSIT-FEEDER
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
SCAVENGER
SCAVENGER
SCAVENGER
PREDATOR
PREDATOR
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
UNKNOWN
INTERFACE
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
iNTEhF&Ct
iMfEM-ACt
INTERFACE
INTEfcFttCE
INTERFACE
SUSttNSION
MOBILITY nan
TU6E &UILD£R-«Ob
TUBE &UILD£R-«OB
TUBE BUILDER-MOB
TUBE BUILDER-HOB
TUBE BU1LDER-WB
TUBE BUILDERH06
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
BURROUER-HOBILE
BURROUER-HOBILE
BURROUER-WBILE
BURROHER-WBILE
WJRROWER-BO&ILE
BURROMER-WEilLE
BURROWER-WOblLE
BURROMER-nOBlLE
BURROUERHOBILE
BURROUER-MOBILE
BURROUER-HOBILE
BURROUER-HOBILE
TUBE BUILDER-HOB
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
TUBE BUILDER-SESSILE
BURROMER-HOBILE
BURROUER-HOBILE
BURROUER-ftOBlLE
EPIFAUNAL-HOBILE
EPIFAUNAL-HOBILE
£PIFAUNflL-*06IL£
EPIFAUNAL-SESSILE
EPIFAUNAL-SESSILE
EPIFAUNAL-HOBILE
EPIFAUNAL-HOBILE
EPIFAUNAL -MOBILE
EPIFAUNAL-HOBILE
EPIFAUNfiL-HOBlLE
EPIFAUNAL-HOBILE
EPIFAUNAL-HOBILE
EPlFAUNAL-n&lLE
EPIFAUNA.-H06ILE
EPIFAUNAL-WfrlLE
EPIFAUNAL-HOBILE
EPIFAUNAL-HOBILE
EP I FAUNft. -MOBILE
EPIFAUNAL-HOBILE
UNKNOWN
&URROHER-HO&1LE
BURROUER-HOBILE
EPIFAUNAL-SESSILE
EPIFAUNAL-SESSILE
BURROUER-HOB1LE
BURROUEft-HO&lLE
PUftftOWER-HOtlLE
MJRHOWER-H06ILE
f-UftROUER-HOblLE
BUftROwtR-HOEilLE
bU^ROWtR-HOBlLE
WJRROHEH-HOblLE
MfiftOWfR-HOblLE
BURRtWEh-HOblLE
KJ&ROWER-SESS1U
UISffRSAL MODt
WIDE OlSftftSAL
HIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
HIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
HIDE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
UNKNOWN
UNKNOWN
UNKNUWN
UNKNOWN
VflRIA&LE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VARIABLE DISPERSAL
VfiRIPBLE DISPERSAL
VHftlfl&LE DISPERSAL
HIDE DISPERSAL
UNKNOWN
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
HIDE DISPERSAL
UNKNOWN
Hit* DISPERSAL
LIMITED DISPERSAL
HIDE L'ISPERSAL
HIDE DISPERSAL
wit* DIS«f»SHL
HIDE DISPERSAL
wlDE DISl-'ERSAL
Hlbt DlbtiRSAL
tftCVr'C LEVEL
DETRITIVORE/OMNIVORE
DETHlTIVUHE/OMNlVOfcE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETR1T1VORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
DETRITIVCRE/OHNIVORE
DETRITIVOR£/OHNIVOR£
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETR1TIVOR£/OHNIVORE
DETRITIVORE/OHNlVOftE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OHNIVORE
HIRBIVORE
HERBIVORE
HERBIVORE
HERBIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
UNKNOWN
DETRITIVORE/OHNIVORE
DETR1TIVORE/OMN1VORE
CARNIVORE
CARNIVORE
CARNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVOftE
DETRITIVORE/OMNIVORE
DETRITIVOftf/OHNlVORE
DETRlTIVOftE/OMNlVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNlVOftE
DETRITIVORE/OHNIVORE
DETRITIVORE/OHNlVOftE
DETRITIVORE/OHNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OHNIVORE
UNKNOWN
DETRITIVORE/OHNIVORE
HERBIVORE
HERBIVORE
HERBIVORE
HERBIVORE
HEP&IVORE
HEftblVORE
HERblvORt
HERblvORE
DETRlTlVOWt/UHNwDRr
D£TRITIVOR£/OMN]VO(
-------
Table 3b (cont'd)
IHXON
TAGELUS PLEBE1US
ENSIS DIRECTUS
SPISULA SOLIDISSIMP
MULINIA LATERAL1S
MYA ARENARIA
CYRTOPLEURA COSTATft
LYONS1A HYALINA
LIMULUS POLYPHEMUS
ACARINA
PYCN060NIDA
CRUSTACEA
CIRRIPEDIA
BALANUS IMPROVISUS
PERICAR1DA MYSIDACEA MYSIDA
NEOMYS1S AMERICANA
MYSIDOPSIS BIGELOWI
CUMACEA
CYCLASPIS VARIANS
LEUCON AMERICANUS
OXYUROSTYLIS SMITHl
PERACARIDA ISOPODA
ERICHSONELLA
I DOTE A BALTICA
EDOTEA TRIL06A
CYATHURA BURBANCX1
PTILANTHURA TENU1S
PERACARIDA AMPHIPODA
AMPELISCA
AMPELISCA ABDITA
AMPELISCA VADORUM
AMPELISCA VERR1LLI
AMPITHOE VALIDA
CYMADUSA COMPTA
CERAPUS TUBULARIS
COROPHIUN
COROPHIUH ACHERUSIOJM
COROPHIUH TUBERCULATUH
ERICHTHONIUS BRASILIENSIS
ERICHTHONIUS RUBRICORNIS
UNCIOLA
UNCIOLA SERRATA
GAMMARIDAE
6AMMARIDAE SP 1
ELASHQPUS LEVIS
6AMMARUS
GAMMARUS MUCRONATUS
MELITA NITIDA
1DUNELLA
LISTR1ELLA BARNARD!
LISTRIELLA CLYMENELLAE
MONOCULODES EDWARDS I
Fl£USTIDAE
STENOTHOIDAE
PARAMETOPELLA CYPRIS
STENOTHOE
STENOTHOE MINUTA
CAPRELLIDAE
AEG1NINA LONG 1 CORN IS
CAPRELLA PENANTIS
PARACAPRELLA TENUIS
PALAEMONETES
PALAEMONETES PUGIO
CRANGON SEPTEMSPINOSA
UPOGEB1A AFFINIS
PORTUNlDAt
CW.LINECTES SAPIDUS
XflNTHlDAE
PlNNOTHERlDflE
PINNIXA
PlNNlXP SAYANP
PHOTON IS
I HOC* II mtlii
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SUSPENSION
SCAVENGER
PREDATOR
PREDATOR
UNKNOWN
SUSPENSION
SUSPENSION
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
UNKNOWN
INTERFACE
INTERFACE
INTERFACE
INTERFACE
UNKNOWN
SCAVENGER
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
INTERFACE
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
UNKNOWN
UNKNOWN
SCAVENGER
SCAVENGER
SCAVENGER
PREDATOR
PREDATOR
PREDATOR
PREDATOR
SCAVENGER
SCAVENGER
SCAVENGER
SCAVENGER
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PREDATOR
PCEDwTG-
PREDATOR
PREDATOR
PREDATOR
PREDATOR
INTERFACE
MOBILITY HODt
BURROWER-SESSILE
BURROWER-nOblLE
BURROWER-MOBILE
BURROWER-MOBILE
BURROWER-MOBILE
BURROWER-MOBILE
EPIFAUNAL -SESSILE
EPIFAUNAL-MOfilLE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
UNKNOWN
EPIFAUNAL -SESSILE
EPIFAUNAL -SESSILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-flOBlLE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
UNKNOWN
TUBE BUILDER-MOB
TUBE BUILDER-HOB
TUBE BUILDER-MOB
TUBE BUILDER-MOB
UNKNOWN
EPIFAUNAL-MOBILE
TUBE BUILDER-MOB
TUBE BUILDER-MOB
TUBE BUILDER-MOB
TUBE BUILDER-MOB
TUBE BUILDER-MOB
TUBE BUILDER-MOB
TUBE BUILDER-MOB
TUBE BUILDER-HOB
BURROWER-MOBILE
BURROWER-MOBILE
BURROWER-MOBILE
BURROWER-MOBILE
BURROWER-MOBILE
BURROWER-HOBILE
UNKNOWN
UNKNOWN
TUBE BUlLDER-SESSlLt
EPIFAUNAL-MOBILE
EPIFAUNAL -MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL -MOBILE
EPIFAUNAL -MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL -MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBlLE
EPIFAuNAL-MOBJLE
EPIFAUNAL -MOBiLE
EPIFAUNAL -MOBILE
EP I FCtuKAL -MOBILE
EPIFAUNAL-MOBILE
EPIFAUNAL-MOBILE
TUBE BUILDER-SESSILE
DISPERSAL MODE
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
UNKNOWN
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
LIMITED DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
wI&E DISPERSAL
WIDE DISPERSE.
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
fhUPnlC LfcVEL
HERBIVORE
HERBIVORE
HERBIVORE
HERBIVORE
HERBIVORE
HERBIVORE
HERBIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRIT1VORE/OHN1VORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
OETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETR1TIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DORITIVORE/OMN1VORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETHlTIVORt/OHNlVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNlVOftE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETHITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETR1TIVORE/OMNIVORE
DETRITIVORE/OHNIVORE
DETRITIVORE/OMNIVORE
DETRITIvORE/OMNlvORE
CARNIVORE
UiRNlvORE
C&RNIVORE
CARNIVORE
CARNIVORE
CARNIVORE
DETRHlVORE/OMNlvORi
-------
Table 3b (cont'd)
TROPHIC hUDE
HOfrlLlTY MODE
DISPERSAL KODt
TRUPnJC LEVEL
ECTOPROCTA
HOLOTHUROIDEA
LEPTOSYNAPTA TENU1S
HEHICHORDATA
ENTEROPNEUSTA
SACCOGLOSSUS
SACC06LOSSUS KOMALEWSKI1
UROCHORDATA
MOL6UL1DAE
MOL6ULA MANW1TTENS1S
SUSPENSION
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
DEPOSIT-FEEDER
SUSPENSION
SUSPENSION
SUSPENSION
EPIFAUNAL -SESSILE
EP I FAUNA. -MOBILE
EP1FAUNAL -MOBILE
BURROWER-SESSILE
BURROUER-SESSILE
BURROWER-SESSILE
BURROUER-SESSILE
EPIFAUNAL-SESSILE
EPIFAUNAL-SESSILE
EP1FAUNAL -SESSILE
LIMITED DlSI^RSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
WIDE DISPERSAL
DETRITIVORE/OMN1VORE
DETRITIVORE/OHNIVURE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMN1VORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
DETRITIVORE/OMNIVORE
HERBIVORE
HERBIVORE
HERBIVORE
-------
TABLE 4. SPECIES COMPOSITION OF DOMINANT GUILDS
Interface feeders, detriv/omniv, mobile tube builders, wide dispersal
Virginia Florida
Dispio uncinata Apoprionospio pygmaea
Loimia medusa Loimia medusa
Paraprionospio pinnata Magelona pettiboneae
Pista palmata Minuspio perkinsi
Polydora ligni Paraprionospio pinnata
Scolecolepides viridis Poecilochaetus johnstoni
Scolelepis (2 sp.) Prionospio heterobranchia
Spio setosa Scolelepis (2 sp.)
Spiophanes bombyx Spiophanes bombyx
Streblospio benedicti
Deposit feeders, detriv/omniv, mobile burrowers, limited dispersal
Virginia Florida
Elasmopus levis Adelodrilus sp.
Gammarus mucronatus Arenicola cristata
Melita nitida Aricidea (7 sp.)
Orbinia ornata Ctenodrilus serratus
Orbiniidae Dasybranchus sp.
Paranais littoralis Enchytraeus (2 sp.)
Scoloplos (3 sp.) Haemonais waldvogeli
Tubificidae Haploscoloplos (3 sp.)
Tubificiodes sp. Immature tubificid w/o cap setae
Limnodriloides (2 sp.)
Monoculoides sp.
Monopylephorus (3 sp.)
Naineris setosa
Nais (2 sp.)
Oiigochaets
Orbinia riseri
Paranais littoralis
Paraonis fulgens
Phallodrilus (3 sp.)
Scoloplos rubra
Scithsondrilus marinus
Stylaria lacustris
Tubifex littoralis
Tubificiodes (6 sp.)
-------
TABLE 4 (cont'd)
Deposit feeders, detriv/omniv, mobile burrower, wide dispersal
Florida
Armandia agilis
Cossura soyeri
Holothuroidea
Mediomastus ambiseta
Notomastus (2 sp.)
Paranaitis speciosa
Siphuncula
Cistena gouldii
Heteromastus filiformis
Mediomastus ambiseta
Notomastus hemipodus
Paraonis sp.
Sternaspidae
Interface feeder, detriv/oraniv, mobile burrower; limited dispersal
Virginia Florida
Cirratulus sp.
Tharyx (2 sp.)
Chaetozone sp,
Cirratulidae
Tharyx sp.
Predator, carnivore, mobile burrower, wide dispersal
Virginia Florida
Amphiduros sp.
Arabellidae
Eteone (3 sp.)
Eulalia sanguinea
Eumida sanguinea
Glycera (2 sp.)
Glycinde solitaria
Gyptis (2 sp.)
Microphthalamus sczelkowii
Nephtys picta
Nephtyiidae
Parahesione luteola
Paranaitis speciosa
Phyllodoce arenae
Pseudeurythoe paucibranchiata
Sigambra tentaculata
Aglaophamus verrilli
Ancistrsyllis (2 sp.)
Cabira incerta
Eteone (2 sp.)
Eumida sanguinea
Glycera americana
Glycinde solitaria
Goniadidae
Gyptis (2 sp.)
Nephtys (3 sp.)
Parahesione luteola
Parandalia americana
Phyllodocidae (3 sp.)
Sigambra (2 sp.)
-------
TABLE 4 (cont'd)
Interface feeder, detriv/omniv, mobile tube builder, limited dispersal
Virginia Florida
Ampilesca (3 sp.)
Cerapus tubularis
Corophium (2 sp.)
Erichthonius (2 sp.)
Unciola serrata
Ampilesca (2 sp.)
Ampharetidae
Carazziella hobsonae
Corophium (2 sp.)
Erichthonius brasiliensis
Hobsonia florida
Mellina maculata
Predator, carnivore, mobile burrower, limited dispersal
Virginia Florida
Amphiporus bioculatus
Carinomidae
Cerebratulus (2 sp.)
Micrura (3 sp.)
Polycladia sp.
Rhyncocoela sp.
Tetrastemma vermiculus
Tubulanus pellucidus
Turbellaria
Arabella sp.
Autolytus sp.
Dorvillea sp.
Ehlersia sp.
Lumberneris latreilli
Marphysa sanguinea
Microphthalamus sp.
Ophiodromus abscura
Pettibonea sp.
Pseudosyllides curacoensis
Schistomeringos rudolphi
Syllis cornuta
Scavenger, detriv/omniv, mobile epifaunal, limited dispersal
Virginia
Aeginina longicornis
Caprella penantis
Caprellidae
Cyathura burbanki
Cymadusa compta
Edotea triloba
Erichsonella
Idotea baltica
Florida
Edotea sp.
Elasmopus levis
Grandidierella bonnieroides
Lembos sp.
Leucothoe spinicarpa
Lysianopsis alba
Melita (3 sp.)
Microdeutopus (2 sp.)
Nassarius vibex
40
-------
TABLE 5 - PERCENT OF TOTAL INDIVIDUALS IN THE TOP 5 GUILDS IN EACH TEST
SPRING 1982
Florida
Guild % of Total
Depos feeder, detriv/omniv, mobile burrower, wide dispersal 46.8
Depos feeder, detriv/omniv, mobile burrower, limited disper 10.6
Interface feeder, detriv/omniv, mobile tube bldr, wide disp 10.3
Scavenger, detriv/omniv, mobile burrower, limited dispersal 6.6
Interface feeder, detriv/omniv, mobl burrower, limited disp 5.0
Virginia
Guild
Interface feeder, detriv/omniv, mobl tube bldr, wide disper 34.6
Depos feeder, detriv/omniv, mobile burrower, limited disper 30.8
Depos feeder, detriv/omniv, mobile burrower, wide dispersal 15.8
Interface feeder, detriv/omniv, mobl burrower, limited disp 11.3
Predator; carnivore, mobile burrower, wide dispersal 4.2
41
-------
Table 5. (cont'd)
FALL 1982
Florida
Guild
Depos feeder, detriv/omniv, mobile burrower, limited disper
Depos feeder, detriv/omniv, mobile burrower, wide dispersal
Interface feeder, detriv/omniv, mobl tube bldr; varibl disp
Predator, carnivore, mobile burrower, wide dispersal
Predator; carnivore, mobile epifaunal, wide dispersal
% of Total
72.5
14.8
8.8
2.6
0.2
Virginia
Guild % of Total
Interface feeder^ detriv/omniv, mobl tube bldr, wide disper 28.9
Depos feeder, detriv/omniv, mobile burrower, limited disper 23.2
Depos feeder, detriv/omniv, mobile burrower, wide dispersal 17.2
Predator; carnivore, mobile burrower, wide dispersal 6.4
Interface feeder, detriv/oraniv, sess tube bldr, limited disp 4.3
SPRING 1983
Florida
Guild
Scavenger, detriv/omniv, mobile tube bldr, limited dispersal
Depos feeder, detriv/omniv, mobile burrower, wide dispersal
Interface feeder, detriv/omniv, mobl tube bldr, limited disp
Interface feeder, detriv/omniv, mobl tube bldr, variabl disp
Scavenger, detriv/omniv, mobile epifaunal, limited dispersal
% of Total
26.6
19.5
16.8
14.4
9.2
Virg inia
Guild % of Total
Interface feeder, detriv/omniv, mobl tube bldr, wide disper 40.1
Depos feeder, detriv/omniv, mobile burrower, limited disper 26.1
Depos feeder, detriv/omniv, mobile burrower, wide dispersal 17.7
Predator, carnivore, mobile burrower, wide dispersal 5.2
Interface feeder, detriv/omniv, sess tube bldr, wide disper 3.4
-------
Table 5. (cont'd)
FALL 1983
Florida
Guild % of Total
Depos feeder, detriv/omniv, mobile burrower, limited disper 27.6
Depos feeder, detriv/omniv, mobile burrower, wide dispersal 25.8
Scavenger, detriv/omniv, mobile burrower, limited dispersal 10.2
Interface feeder^ detriv/omniv, mobl tube bldr, wide disper 9.5
Interface feeder, detriv/omniv, mobl tube bldr, limited disp 6.4
irginia
Guild % of Total
Predator, carnivore, mobile burrower, wide dispersal 14.9
Scavenger, detriv/omniv, mobile epifaunal, limited dispersal 6.9
Interface feeder, detriv/omniv, mobl tube bldr, wide disper 6.8
Depos feeder, detriv/omniv, mobile burrower, limited disper 6.2
Predator, carnivore, mobile epifaunal, wide dispersal 5.8
SPRING 1985
Florida
Guild % of Total
Deposit feeder, detriv/omniv, mobile burrower, limited disp 28.9
Deposit feeder, detriv/omniv, mobile burrower, wide dispersal 21.0
Scavenger, detriv/omniv, mobile burrower, limited dispersal 17.9
Depos feeder, detriv/omniv, mobile tube bldr, wide dispersal 11.0
Interface feeder, detriv/omniv, mobile burrower; limited disp 4.0
Virginia
Guild % of Total
Predator, carnivore, mobile burrower, wide dispersal 11.0
Interface, detriv/omniv, sess tube bldr, wide dispersal 9.3
Depos feeder, detriv/omniv, mobile burrower, limited disper 8.5
Interface feeder, detriv/omniv, mobl tube bldr, limited disp 6.9
Predator, carnivore, mobile epifaunal, wide dispersal 5.2
-------
Table 5. (cont'd)
FALL 1985
Florida
Guild % of Total
Deposit feeder, detriv/omniv, mobile burrower, limited disp 43.8
Deposit feeder, detriv/omniv, mobile burrower, wide dispersal 21.8
Interface feeder, detriv/omniv, mobl tube bldr, wide dipersal 9.6
Scavenger, detriv/omniv, mobile burrower, limited dispersal 9.2
Interface feeder, detriv/omniv, mobl tube bldr, limited disper 4.4
Virginia
Guild % of Total
Deposit feeder, detriv/omniv, mobile burrower, limited disp 36.6
Interface feeder, detriv/omniv, mobl tube bldr, wide disper 31.0
Deposit feeder, detriv/omniv, mobile burrower, wide dispersal 10.3
Predator, carnivore, mobile burrower, limited dispersal 4.4
Predator, carnivore, mobile burrower, wide dispersal 3.4
44
-------
Table 6. Guilds which showed good agreement between
temporal trends in the lab and field.
Interface-feeder, detritivore/ omnivore, mobile burrower, limited dispersal,
Interface-feeder, detritivore/omnivore, mobile tube-builder, limited
dispersal.
Deposit-feeder, detritivore/omnivore, mobile burrower, wide dispersal.
Predator, carnivore, mobile burrower, limited dispersal.
Scavenger, detritivore/omnivore, mobile epifauna, limited dispersal.
-------
Table 7. Evaluation of concordance between laboratory and field results PCP-dose experiments.
Virginia Spring 1985 test is omitted since dose-equivalency was not achieved.
Community
Component
Species
Richness
Guild:
INDOBMLD1
INDOTMLD2
DFDOMBWD3
PRCVBMLD"
SCDOEMLD5
Spring
1985
Florida
Response In Response In
Laboratory Field
Reduction by
low and
high doses
No effect
of dose
Slight
reduction at
high dose
Reduction by
low and
high doses
Too few to
predict
Too few to
predict
Reduction by
high dose
No effect
of dose
Slight
reduction at
high dose
Reduction by
low and
high doses
Too few to
predict
Too few to
predict
Fall
Florida
Response In Response In
Laboratory Field
Reduction by
high dose
Recruitment
depressed by
high dose
Reduction by
low and
high doses
Reduction by
high dose
No effect
of dose
Too few to
predict
Reduction by
high dose
Recruitment
depressed by
high dose
Reduction by
high dose
Reduction by
high dose
No effect
of dose
Too few to
predict
1985
Virginia
Response In Response In
Laboratory Field
Reduction by
high dose
Too few to
predict
Recruitment
depressed by
high dose
No effect
of dose
Slight
depression of
recruitment
Too few to
predict
Reduction by
high dose
Too few to
predict
Recruitment
depressed by
high dose
No effect
of dose
No effect
of dose
Too few to
predict
1 Interface-feeder, detritivore/omnivore, mobile burrower, limited dispersal.
2Interface-feeder, detritivore/omnivore, mobile tube-builder, limited dispersal.
'Deposit-feeder, detritivore/omnivore, mobile burrower, wide dispersal.
MPredator, carnivore, mobile burrower, limited dispersal.
5Scavenger, detritivore/omnivore, mobile epifauna, limited dispersal.
-------
-pi
-,!
CAPE SAN BLAS
FIGURE i-a Apalachicola Bay, Florida Study Site
-------
00
FIGURE 1-b York River, Virginia Study Site
-------
FIGURE 2 - MEAN ABUNDANCE IN WEEKLY SAMPLES FROM 1981 - 1986
Vertical lines indicate test dates.
2
U
o
o
oc.
u
a.
kj
o
<
o
z
D
m
z
UJ
2
APALACHICOLA BAY, PL
150
100«
50
\
YORK RIVER, VA
2
u
o
o
UJ
Q-
UJ
U
m
300
200
100
49
-------
FIGURE 3 - TOTAL MACROFAUNA AND SPECIES RICHNESS
SPR NG 1982
LAB AND FIELD CONTROLS
Ul
o
APALACHICOLA BAY, FL
YORK RIVER, VA
•na »tKct MTUTioN rnritt
TUC JtNCt NTTUTIOM CWItXl)
APALACHICOLA BAY, FL
TUT JINCt MTUTIOH (Kt»J)
YORK RIVER, VA
TIUI JINCl MTUTION (»ttlt5)
FIELD
LAB
-------
FIGURE 4 - TOTAL MACROFAUNA AND SPECIES RICHNESS
FALL 1 982
LAB AND FIELD CONTROLS
APALACHICOLA BAY, FL
rut >wct MTUTIOM rwttn)
YORK RIVER, VA
\
•nut jiNct MTUTioH (wit«)>
APALACHICOLA BAY, FL
TUC JlWCt MTUT10N (Tttlfl)
YORK RIVER, VA
TIUC JINCt MTliTIOM f»tt»J)
FIELD
LAB
-------
FIGURE 5 - TOTAL MACROFAUNA AND SPECIES RICHNESS
SPR NG 1983
LAB AND FIELD CONTROLS
LH
ro
APALACHICOLA BAY, FL
YORK RIVER. VA
TTJt JIHCt MTUT10N (WtCkl)
TIUI tOtCl MTlATiaM (VttKl)
APALACHICOLA BAY, FL
TXIC JIMCT MTUT10N (VCCKI)
YORK RIVER, VA
nut iiNct tmiTioM
FIELD
LAB
-------
FIGURE 6 - TOTAL MACROFAUNA AND SPECIES RICHNESS
FALL 1 983
LAB AND FIELD CONTROLS
un
U-J
APALACHICOLA BAY, FL
YORK RIVER, VA
TUt JIHCt MTUTIOM (•ECK3)
TUt 1WCI MTUTION OttKJ)
APALACHICOLA BAY, PL
TUC 3INCC MTUTION (»CCK]>
YORK RIVER, VA
TUC IMCE MTUTION (WCEK5)
FIELD
LAB
-------
FIGURE 7 - TOTAL MACROFAUNA AND SPECIES RICHNESS
SPRING 1 985
LAB AND FJELD CONTROLS
APALACHICOLA BAY, FL
TUC JWCt MTUTION (WRK»
YORK RIVER, VA
TUt SINCE MTUTION (*tCK]>
APALACHICOLA BAY, FL
TWE JKCC MtlATlOH CVtEK»
YORK RIVER, VA
TUt XMCI MTUTION f«EEK5)
FIELD
LAB
-------
FIGURE 8 - TOTAL MACROFAUNA AND SPECIES RICHNESS
FALL 1 985
LAB AND FIELD CONTROLS
un
Ln
APALACHICOLA BAY, FL
YORK RIVER, VA
TUC 1MCE tnUTIOM (VCEKI)
TUt JIMCE MTUTION (VC[K»
APALACHICOLA BAY, FL
TUC JINCt MTUTION OttXJ)
YORK RIVER, VA
TUt IINCC MTUTIOH (IfttKJ)
FIELD
LAB
-------
FIGURE 9
LAB-FIELD CONTROL COMPARISONS
TESTS 1 -7
GUILD: PREDATOR, CARVIVORE, MOBILE BORROWER, LIMITED DISPERSAL
APALACHICOLA BAY, PL
YORK RIVER, VA
01
a
in
u
o
o
a
UJ
a.
u
o
o
z
5
UJ
2
01 3 3 * 5 7 • »
a
in
U
o
o
OL
Ul
a.
a
z
3
100
•0
01 1 3 * 9 t I
TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPAR SONS
TESTS 1 -7
GUILD: SCAVENGER, DETRIV/OMNIV, MOBILE EPIFAUNA, LIMITED DISPERSAL
APALACHICOLA BAY, FL
YORK RIVER, VA
O
o
o
oc
UJ
0.
UJ
u
3
UJ
01 1343(711
too.
O
in
o
o
oc
bJ
0.
u
u
o
z
0.00 1.00 2.00 3.00 4.00 3.00 t.OO
TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 CCont'd)
LAB-FIELD CONTROL COMPARISONS
TESTS 1 -7
GUILD:DEPOSIT-FEEDER, DETRIV/OMNIV, MOBILE BURROWER, WIDE DISPERSAL
APALACHICOLA BAY, FL
YORK RIVER, VA
un
CD
U
o
o
UJ
a.
ui
u
o
z
I
UI
a
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0123
TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPARISONS
TESTS 1 -7
GUILD: INTERFACE-FEEDER, DETRIV/OMNIV, MOBILE TUBE-BLD, LIMITED DISPERS
APALACHICOLA BAY, FL
YORK RIVER, VA
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPARISONS
TESTS 1-7
GUILD: INTERFACE FEEDER, DETRIV/OMNIV, MOBILE BURROWER, LIMITED DISPERS
APALACHICOLA BAY, FL
YORK RIVER, VA
en
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TIME SINCE INITIATION (WEEKS)
FIELD
TAB""
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPARISONS
TESTS 1 -7
GUILD: SUSPENSION-FEEDER, HERBIVORE, MOBILE BURROWER, WIDE DISPERSAL
APALACHICOLA BAY, FL
YORK RIVER, VA
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPARISONS
TESTS 1 -7
GUILD: DEPOSIT-FEEDER, DETRIV/OMNIV, MOBILE BURROWER, LIMITED DISPERS
APALACHICOLA BAY, FL
YORK RIVER, VA
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPAR SONS
TESTS 1 -7
GUILD: INTERFACE EEEDER, DETRIV/OMNIV, MOBILE TUBE-BLD, WIDE DISPERSAL
APALACHICOLA BAY, FL
YORK RIVER, VA
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 9 (Cont'd)
LAB-FIELD CONTROL COMPARISONS
TESTS 1 -7
GUILD: PREDATOR, CARNIVORE, MOBILE BURROWER, WIDE DISPERSAL
APALACHICOLA BAY, FL
YORK RIVER, VA
en
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB ~
-------
FIGURE 1 OA
LOW DOSE
SPRING 1985
- LAB AND FIELD PCP LEVELS
APALACHICOLA BAY, FL
YORK RIVER, VA
en
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 10B - LAB AND FIELD PCP LEVELS
H GH DOSE
SPRING 1 985
APALACHICOLA BAY, FL
YORK RIVER, VA
en
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TIWE SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 1 1 A
LOW DOSE
FALL 1 985
- LAB AND FIELD PCP LEVELS
APALACHICOLA BAY, FL
YORK RIVER, VA
en
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1.5
TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FIGURE 1 1 B
H GH DOSE
FALL 1 985
- LAB AND FIELD PGP LEVELS
APALACHICOLA BAY, FL
YORK RIVER, VA
CD
00
2
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TIME SINCE INITIATION (WEEKS)
TIME SINCE INITIATION (WEEKS)
FIELD
LAB
-------
FGURE 12 - RESPONSE TO PCP
TOTAL MACROFAUNA-
SPRING 1 985
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
en
in
TUC JINCC INITIATION nrtcxi)
TIUC JINCl INITIATION fWEEKJ)
APALACHICOLA BAY, FL - LAB
TIUC SINCE IVIITUTION OtfCCKl)
YORK RIVER, VA - LAB
\
\
TlUt SINCE INITIATION (YCCK3)
CONTROL
LOW DOSc
HIGH DOSL
-------
F CURE 13 - RESPONSE TO PCP
SPEC ES RICHNESS •
SPRING 1985
O
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
TIUE SINCE INITIATION CVCCKJ)
TUC SINCE INITIATION (WCCICS)
APALACHICOLA BAY, FL - LAB
f •
TIUC SINCE INITIATION (WEEKJ)
YORK RIVER, VA - LAB
TlUt 1MCI INITIATION. CHEEKS)
CONTROL
LOW DOSE
HIGH DOSE
-------
FGURE 14 - RESPONSE TO PCP
TOTAL MACROFAUNA •
FALL 1 985
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
8
a
Z
3
5
TIUC jwcc wmAnoH
TlUt 1IHCC MTWHOM
APALACHICOLA BAY, FL - LAB
TlUt JlNCt CWTUTIOM (WICKJ)
YORK RIVER, VA - LAB
TTUt JINCC MTDAT10N (WCCKJ)
CONTROL
LOW DOSE
HIGH DOSE
-------
F CURE 15 - RESPONSE TO PCP
SPEC ES R CHNESS •
FALL 1 985
ro
APALACHICOLA BAY, PL - FIELD
YORK RIVER, VA - FIELD
J«Ct IWTUT1ON CVCtKI)
•nut SINCE nrruTioN
APALACHICOLA BAY, FL - LAB
nut suet imriATioN rwrcicj)
YORK RIVER, VA - LAB
nut since MTwnoM CWEIKS)
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 16 - RESPONSE TO PCP
SPR NG 1985
GUILD: PREDATOR, CARNIVORE, MOBILE BURROWER, LIMITED DISPERSAL
APALACHICOLA BAY, FL - FIELD
•nm max HIUTUB ncaca
YORK RIVER, VA - FIELD
I I
' ..1
I '
I -J
1
'V
APALACHICOLA BAY, FL - LAB
3 J: . '"••..
nui tftct MTunoa
YORK RIVER, VA - LAB
~1
-
1
T»M MM MTUTK>H <
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 16 - RESPONSE TO PCP
SPRING 1 985
GUILD: SCAVENGER, DETRITIV/OMNIV, MOBILE EPIFAUNA, LIMITED DISPERSAL
APALACHICOLA BAY, FL - FIELD
YORK RIVER. VA - FIELD
APALACHICOLA BAY, FL - LAB
:i
YORK RIVER, VA - LAB
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 16 - RESPONSE TO PCP
SPRING 1 985
GUILD: DEPOSIT-FEEDER, DETRIV/OMNIV, MOBILE BORROWER, WIDE DISPERS
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
TM WOE HTUTIOB attxn
APALACHICOLA BAY, FL - LAB
YORK RIVER, VA - LAB
TU tote* MTUTWII
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 16 - RESPONSE TO PCP
SPRING 1985
GUILD: INTERFACE-FEEDER, DETRIV/OMNIV, MOBILE TUBE-BLD, LIMITED DISPERS
CTi
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
IWCC MnMlOl OKZKD
(mown nczxu
APALACHICOLA BAY, FL - LAB
YORK RIVER, VA - LAB
CONTROL
TOW DOSE
HIGH DOSE
-------
FIGURE 16 - RESPONSE TO PCP
SPR NG 1985
GUILD: INTERFACE-FEEDER, DETRIV/OMNIV, MOBILE BORROWER, LIMITED DISPERS
APALACHICOLA BAY, FL - HELD
YORK RIVER, VA - FIELD
~l
TUB 1MOE MTTVttlOM
•n* ana wrvmoK nccu>
APALACHICOLA BAY, FL - LAB
1
I-"
\
T»M mic* •rrvk-no* atoa)
YORK RIVER, VA - LAB
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 17 - RESPONSE TO PCP
FALL 1 985
GUILD: DEPOSIT-FEEDER, DETRIV/OMNIV, MOBILE BORROWER, WIDE DISPERS
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
CO
APALACHICOLA BAY, FL - LAB
YORK RIVER, VA - LAB
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 17 - RESPONSE TO PCP
FALL 1 985
GUILD: INTERFACE-FEEDER, DETRIV/OMNIV, MOBILE BORROWER, LIMITED DISPERS
OJ
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
I -i
\ u\
-r
APALACHICOLA BAY, FL - LAB
YORK RIVER, VA - LAB
I "
\ »
CONTROL
LOW DOSE
HIGH DOSE
-------
FIGURE 17 - RESPONSE TO PCP
FALL 1 985
GUILD: PREDATOR, CARNIVORE, MOBILE BORROWER, LIMITED DISPERSAL
CO
o
APALACHICOLA BAY, FL - FIELD
YORK RIVER, VA - FIELD
APALACHICOLA BAY, FL - LAB
YORK RIVER, VA - LAB
CONTROL
LOW DOSE
HIGH'DOSE
-------
FIGURE 1 7
FALL 1 985
RESPONSE TO PCP
GUILD: INTERFACE-FEEDER, DETRIV/OMNIV, MOBILE TUBE-BLD, LIMITED DISPERS
CD
APALACHICOLA BAY, FL - FIELD
r
YORK RIVER, VA - FIELD
APALACHICOLA BAY, FL - LAB
YORK RIVER, VA - LA8
CONTROL
LOW DOSE
HIGH DOSE
------- |