EPA-660/3-74-031
MAY 1975
Ecological Research Series
Environmental Requirements of Selected
Estuarine Ciliated Protozoa
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National Environmental Research Center
Office of Research and Development
U.S. Environmental Protection Agency
Corvallis, Oregon 97330
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RESEARCH REPORTING SERIES
Research reports of the Office of Research and Development,
U.S. Environmental Protection Agency, have been grouped into
five series. These five broad categories were established to
facilitate further development and application of environmental
technology. Elimination of traditional grouping was consciously
planned to foster technology transfer and a maximum interface in
related fields. The five series are:
1. Environmental Health Effects Research
2. Environmental Protection Technology
3. Ecological Research
4. Environmental Monitoring
5. Socioeconomic Environmental Studies
This report has been assigned to the ECOLOGICAL RESEARCH STUDIES
series. This series describes research on the effects of pollution
on humans, plant and animal species, and materials. Problems are
assessed for their long- and short-term influences. Investigations
include formation, transport, and pathway studies to determine the
fate of pollutants and their effects. This work provides the technical
basis for setting standards to minimize undesirable changes in living
organisms in the aquatic, terrestrial and atmospheric environments.
EPA REVIEW NOTICE
This report has been reviewed by the Office of Research and
Development, EPA, and approved for publication. Approval does
not signify that the contents necessarily reflect the views and
policies of the Environmental Protection Agency, nor does mention
of trade names or commercial products constitute endorsement or
recommendation for use.
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EPA-660/3-74-031
MAY 1975
ENVIRONMENTAL REQUIREMENTS OF SELECTED
ESTUARINE CILIATED PROTOZOA
by
Arthur C. Horror
Department of Zoology
University of New Hampshire
Durham, New Hampshire
Grant No. 18080 FEW
Program Element 1BA022
Project Officer
Juan G. Gonzalez
National Marine Water Quality Laboratory
National Environmental Research Center
South Ferry Road
Narragansett, Rhode Island 02882
NATIONAL ENVIRONMENTAL RESEARCH CENTER
OFFICE OF RESEARCH AND DEVELOPMENT
U. S. ENVIRONMENTAL PROTECTION AGENCY
CORVALLIS, OREGON 97330
For Sili by tin National Technical Information Service
U.S. Department of Commerce, Sprincfidd. VA 22151
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ABSTRACT
This report addresses species composition and microdistribution of
ciliates (Protozoa, Ciliophora) of a tidal marsh at Adams Pt., Great
Bay, New Hampshire (1970, 1971) in relationship to temperature, salinity,
pH, concentration of oxygen, H2S, and bacteria, and occurrence of
micrometazoa. Accurate counting and precise identification allowed
measurement of tidal effects on ciliate abundance and diversity, and
the relationship of ciliates to micrometazoa and bacteria.
During 1970, we identified 79 species in 175 collections; during 1971,
83 species in 102 collections0 Although in general ciliate distribution
was not correlated with temperature, salinity, pH, or oxygen concen-
tration, some species were tolerant of anoxic environments. Ciliates
differed in distribution between the upper (Spartina patens) and the
lower (S o alterniflora) marsh. We measured responses of bacteria and
ciliates to the physical and biological changes in a patens-panne pond
caused by tidal flushing, and to the flushing of a channel in the lower
marsh by several different tidal cycles.
This report was submitted in fulfillment of Grant 18080 FEW by The Uni-
versity of New Hampshire, Durham, under the sponsorship of the Environmental
Protection Agency. Work was completed as of September 1971.
ii
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CONTENTS
Page
Abstract ii
List of Tables v
Acknowledgements vi
Sections
I Conclusions 1
II Recommendations 3
III Introduction 4
IV Materials and Methods 9
V Experimental Phase 15
VI Discussion 41
VII References 44
VIII List of Publications 49
iii
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TABLES
No. Page
1 Occurrence of several species as a function of salinity 21
2 Coleps tesselatus occurrence and abundance as a
function of oxygen concentration 22
3 Mesodinium pulex occurrence and abundance as a
function of oxygen concentration 23
4 Uronema filificum occurrence and abundance as a
function of oxygen concentration 24
5 Strombidium sulcatum occurrence and abundance as
a function of oxygen concentration 25
6 Strombidium styliferum occurrence and abundance
as a function of oxygen concentration 26
7 Strombidium latum occurrence and abundance as a
function of oxygen concentration 27
8 Faunal differences between upper and lower marsh 29
9 Locomotory types represented by species unique to
either upper marsh or lower marsh 30
10 Faunal differences between upper and lower marsh 31
11 Physical and biological parameters of a patens-panne pond 32
12 Physical and biological parameters in the lower marsh
during a full moon spring tide cycle 36
13 Physical and biological parameters in the lower marsh
during a neap tide cycle 38
14 Physical and biological parameters in the lower marsh
during a new moon spring tide cycle 39
IV
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ACKNOWLEDGEMENTS
The valuable assistance of Miss Hope Godino, Mr. Kenneth McGeary/ and
Mr. Edward Washburn, Research Assistants, is gratefully acknowledged.
Additionally, the productive input of Mr. Richard Kool, Mr. and Mrs.
Gerry Gagne, and Mrs. Edward Washburn, Laboratory Assistants, is also
greatly appreciated.
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SECTION I
CONCLUSIONS
During 1970 and 1971, 103 species of ciliates, representing 41 families
and 10 orders were identified.
No correlations were noted between ciliate occurrence or abundance and
the factor of temperature.
Several instances of negative correlations between ciliate concentration
and H2S concentration were observed.
Although some pairs of closely related species showed differences in
relationship to salinity, occurrence of the most frequently encountered
species showed no correlation with salinity.
In general the most regularly occurring species show no obvious
correlation either in occurrence or abundance with oxygen concentration.
However, numerous less regularly occurring species are tolerant of
anoxic environments.
Taxonomic procedures allowing precise species identification permitted
assessment of critical differences in microdistribution. Comparison of
ciliate faunas of the upper and lower marsh indicates instances of
intrageneric differences in distribution.
Tidal flushing of pools in the patens-panne upper marsh apparently cause
no effects upon bacterial numbers. Following flushing of such ponds by
high water, there appeared an increase in ciliate diversity on the
bottom, and a pulse in ciliate concentration. At such times, Coleps
tesselatus, that occurred more regularly in the upper marsh than in the
lower marsh, apparently was flushed through the channels of the lower
marsh and appeared in greater than usual concentration on the bottom of
the tidal channels.
Ciliates on the bottom of tidal channels of the lower marsh occurred at
greater concentrations and in greater variety at ebb tide than at flood
tide. By contrast, ciliates in surface waters of tidal channels of the
lower marsh occurred at greatest concentrations at flood tide, reflecting
an influx of individuals from the estuary.
There was no significant correlation between numbers of ciliates and
bacterial concentration in tidal channels of the lower marsh during the
course of a tidal cycle, whether it was a full moon spring tide, new
moon spring tide, or neap tide.
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Tidal marsh ciliates appear to settle into the detritus in the bottom of
the tidal channels of the lower marsh following flooding of the marsh by
a tidal cycle. In general they are not flushed out onto the mud of the
estuary below the marsh.
Contribution of tidal marsh ciliates in general to estuarine food webs
probably occurs through their association with particulate detritus.
Tidal marsh ciliates do not seem to play a significant role in the trans-
fer of organic matter from tidal marshes to the rest of the estuary
except insofar as they are carried from the marsh on particulate organic
detritus. Likewise, their concentrations in tidal marshes are not
noticeably increased by influx of protozoa from the estuary during flood
tide.
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SECTION II
RECOMMENDATIONS
Because of intrageneric differences in responses of ciliates to
environmental factors careful species identification is necessary in any
application of differences in ciliate microdistribution to assessment of
biological effects of changing water quality.
Direct counts of living protozoa augmented by permanent cytological
preparations are practical, consistent, and allow qualitative and
quantitative assessment of responses of ciliate populations to changes
in water quality.
Estimation of bacterial standing crop by Most Probable Number methods
was inadequate for assessment of the correlation between changes in
ciliate populations and changes in bacterial concentration, thus should
be replaced in future work by alternative approaches to measuring
bacterial density, such as direct observation counts.
Data such as those provided in this report can act as a baseline for
comparing protozoan diversity among marine habitats. These data indicate
a possible source of information for assessing biological response to
changing chemical and physical parameters in estuaries. Comparison of
the character of the ciliate fauna of tidal marshes subject to water of
differing quality should provide further understanding of the response
of tidal marshes to possible factors associated with pollution.
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SECTION III
INTRODUCTION
This study is designed to determine the possible role of ciliated
microorganisms (Phylum Protozoa, class Ciliophora) as water pollution
indicators. It involves investigation of factors affecting species
composition and microdistribution of populations of intertidal ciliates
of the tidal marshes of the Great Bay - Little Bay estuary, New
Hampshire, where they reach particularly high population densities.
The relationship of ciliates to the rest of the decomposer food chain
and their role in the tidal marsh and estuarine productivity will be
investigated. Measurements of relative and absolute abundance of members
of the microbenthos,.correlated with variations of physical, chemical,
and microbiological tactors, will be employed in assessing diurnal and
monthly changes in ciliate community structure. The research was carried
out using facilities at the University of New Hampshire, and the Jackson
Estuarine Laboratory of the University of New Hampshire, on Adams Pt.,
Great Bay, New Hampshire.
The necessity of a fuller understanding of the biological complexity of
the estuaries of the northeastern coast of the United States is becoming
greater with the increasing stress put on these systems by our increasing
population and its technology1. As a site of a significant if not major
portion of the productivity of estuaries, tidal marshes in particular
continue to be important foci for basic research2'•*. Evidence from many
directions implicates bacterial decomposition of detritus derived from
tidal marshes as one of the basic processes of energy transformation
involved in the food chains of estuaries^"9.
In addition, primary productivity by microscopic saltmarsh algae provides
a significant fraction of the carbon available for estuarine ecosys-
tems °~12. During the past 10 years, the approximately 50 publications
on taxonomy, distribution, and ecology of free-living benthic ciliated
protozoa continues to demonstrate that ciliates are cosmopolitan, their
local occurrence depending on suitable microhabitats. A large number of
these papers report investigations of the psammolittoral zone, with
emphasis on physical factors (e.g. that of Dragesco 3). There have been
few investigations of ciliates of tidal marshes, compared with the
psammolittoral zone1'* '^. Despite the known high productivity of tidal
marshes, and the potential these coastal wetlands have for supporting
a rich and diverse microfauna, relatively few protozoologists have
investigated the ditches, pools, and algal mats of coastal marshes.
Compared with the ciliate fauna of other marine habitats, such as the
psammolittoral, little is known of the extent or variety of the ciliate
population inhabiting coastal wetlands.
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By contrast, numerous protozoologists have come to the conclusion that
in fresh water habitats, ciliates occur in a wide variety of habitats
and may have value in assessing water quality 1°~20.
While conducting doctoral research on ecological relationships among
benthic marine ciliates at the Florida State University21 it became
increasingly clear to me that patterns of distribution of marine ciliates
were correlated with availability of suitable microhabitats22- Some of
this dependence was related to food and feeding; studies of Cohnilembus
indicated morphological specializations associated with feeding and
locomotory behavior2-'. After moving to the University of New Hampshire,
my interests in morphological specialization of ciliates and their
relationships to ecology continued2^. Studies of distribution of
ciliates2^ further indicated that congenors occupied different niches
and often different microhabitats.
Effects of levels of viral and bacterial pollution upon shellfish pro-
duction and productivity in the Great Bay - Little Bay estuarine complex
are continuing areas of investigation at the University of New Hampshire.
However, the ciliated protozoa, members of the next higher trophic
level to bacteria in the decomposer food chain, had been investigated
here only preliminarily. Thus I began investigation of the morphology
and ecology of tidal marsh ciliates^'2"7. A series of monthly samples
between July, 1964 and June, 1965 showed seasonal differences in popu-
lation structure, replacement of species, and indications of differences
in ciliate fauna compared with intertidal mud or sand. These studies
O Q O O
and others^°~JJ suggested that ciliates play an important role in the
ecology of the benthos and show responses to factors that may allow them
to be useful as indicator organisms. The ciliated protozoa in addition
seem particularly adaptable to investigation of ecological principles at
work3*"38.
THEORETICAL APPROACHES
This study has been planned to include taxonomic procedures to assure
precise species identification and an approach to ciliate community
ecology to allow assessment of seasonal turnover and succession. Criti-
cal differences in microdistribution may be significant in relationship
to spatial differences in environmental factors associated with pollu-
tion. This is expected to be a relatively unique and significant aspect
of the research and is necessary prior to evaluation of particular
species as indicator organisms.
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Faunistic Study
Despite our information generally on the biology of Great Bay39, and the
wealth of information on the microbiology of the Bay ' , the ecological
role of ciliates here has been little studied. Published works from
elsewhere in the world suggest that one should expect to find rich and
diverse ciliate faunas in appropriate microhabitats in Great Bay .
The following faunistic study enables one to identify possible competi-
tive situations, to assess the relationship of locomotory type with
microhabitat preference, allows assessment of possible correlation of
microdistribution with various physical and chemical factors, and finally
allows enumeration of species available for further culturing and experi-
mental purposes.
Quantitative Sampling
Development of accurate and reproducible counting methods for tidal marsh
ciliates should allow assessment of effects of tidal flushing upon
ciliate abundance and diversity, tide cycle effects upon contribution of
ciliates to the estuary proper, and the relationship of ciliates to
micrometazoa and bacteria.
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ADAMS
'° PI
71 °° 10 miles
1 mile
Figure 1. Great Bay, New Hampshire
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'iff' f\
'''/"
Sporting alternifiora
Sporting pgtens
S. glterniflorg
(dwarf)
sedges
water
10m
Figure 2. Tidal marsh
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SECTION IV
MATERIALS AND METHODS
SITE
Description of Sampling Area
Adams Point tidal marsh (Latitude 43°5'22" N.; Longitude 70O5'15" W.)
comprises about 300 m2 of intertidal wetlands at the north end of Great
Bay, New Hampshire (see Pigs. 1,2). Major features of the marsh, illu-
strated in Fig. 2, include extensive panne ponds surrounded by Spartina
patens, dwarf Spartina alterniflora, and sedges (Scirpus sp.), as well
as lesser numbers of other typical tidal marsh plants. This portion of
the marsh, referred to below as "upper marsh" comprises the northeastern-
most 100 m2 of the study area. The upper marsh contrasts sharply with
the remainder of the area, referred to below as the "lower marsh". Here
the dominant vegetation is tall Spartina alterniflora accompanied by
small numbers of various broad-leaved plants including Atriplex sp.,
Salicornia sp., and Limonium sp. A small number of extensive panne
ponds, here elongated in an east-west direction/ are confined to the
northern part of the lower marsh. In the remainder of the lower marsh,
there are numerous narrow tidal channels 1 m deep draining to the south-
west.
Although the maze-like configuration of these ditches including many
blind channels and isolated sections of ditch may change from year to
year due to ice action, the general drainage patterns remain consistent.
One important drainage pattern includes a series of more or less parallel
ditches and widened pools along the south edge of the sampling area in
the lower marsh.
Although the algal flora of the marsh was not examined extensively from
a taxonomic point of view, there are extensive seasonal developments of
filamentous green algae (Cladophora sp.) and bluegreen algae around the
margins of the panne ponds. Pelt-like mats of Vaucheria occur along
edges of ditches in the lower marsh.
By mid-February, the entire sampling area in Fig. 2 is covered by 15-
45 cm of ice. In the coldest winters, ice extends to the bottom of the
deepest pannes in the upper marsh.
At spring ice-out in March, the vegetative cover of the lower marsh is
scoured free, the maze of channels is altered, and entire blocks of peat
become dislodged from the underlying till and come to rest out on the
tidal flat.
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The average tidal amplitude is approx. 2.5 m but varies considerably
from predicted values due to the extensive tidal flats and the shallow-
ness of the estuary. At times of spring high water the water level
reaches the road edge at the top of Fig. 2.
Quadrat System
To assist in development of Fig. 2, and to allow precise location of all
collections, the system of quadrats, each 10 m square was surveyed across
the entire area of Fig. 2. The corner of each square was marked by a
wooden stake. Stakes were identifiable by painted numbers and letters.
FIELD METHODS
Permanent Station -Platforms
Two elevated wooden platforms, each 1 x 2 m were constructed in sites
chosen for repeated sampling, to avoid unnecessary trampling of the tidal
marsh peat (black squares in Fig. 2). One of these, in the upper marsh,
extended to the southwest border of a panne pond, facilitating collec-
tions from the edge of the pond and the floating algal mats there. The
second platform, in the lower marsh, extended to the edge of a narrow
tidal creek, allowing repeated collections from the water running in
that creek without disturbing sediments on either side. The platforms
were large enough to accommodate three technicians and our assemblage of
field equipment (see description below).
Equipment
pH - Beckman portable field pH meter, model 1009, equipped with an
electrode sufficiently long to allow measurement of vertical gradients
in pH at all sampling locations.
Temperature - A 0.5 m rod, drilled at 5 cm intervals, accommodated mini-
ature thermistor probes connected through a 10-point switch box to an
Atkins Temperature Indicator (Model 3HOI-C10). Such an apparatus allowed
nearly simultaneous measurement of vertical temperature gradients at any
station.
Oxygen - Following the method of Burke^, we drew a water sample into the
barrel of a 10 ml syringe that had been greased with vacuum grease to
prevent the plunger from slipping. Appropriate volumes of manganous
sulfate, alkaline iodide solution, and ortho phosphoric acid were then
drawn into the syringe in sequence in the field. We conducted the rest
of the oxygen determination later in the laboratory.
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H2S - During the summer of 1970, H2S concentration was estimated using
the Hach sulfide test kit, Model H3-1.
Salinity - We collected water samples in 50 ml screw cap bottles for
later laboratory determination of salinity.
Field Box - We designed and constructed a plywood weather-proof field
box built to accommodate the pH meter, the thermistor and switchbox and
probes, glassware for water samples, equipment and supplies for field
oxygen and I^S determination, and recording equipment.
Sampling for Ciliated Protozoa
Tubes - Collection tubes were 10 cm lengths of polyvinyl chloride non-
toxic tubing, 2.5 cm I.D., that were filled and stoppered beneath the
water surface. Ordinarily, each such tube, of an average volume of
18-20 ml when stoppered, provided 4-5 replicates for quantitative
analysis of ciliate populations (see Laboratory Methods section) . The
proximity of the marsh station to the Jackson Estuarine Laboratory (less
than 0.5 km) permitted examination of tube contents within 1 hr after
collection.
Syringes - We collected water from the interstices of mats of filamentous
green algae by drawing water from the mats into a 10 ml syringe.
Although not accurate quantitatively, due to the narrow aperture of the
syringe, such technique allowed sampling from microhabitats not available
to tube sampling.
Noland Bottles - By fitting a 250 ml wide-mouth bottle with a 2-hole
rubber stopper, each hole fitted with a V-shaped glass tube, and one of
these fitted with a 0.5 m length of rubber tubing, we were able to aspi-
rate into the bottle small volume samples from specific spots on the
bottoms of panne ponds .
Bacteriological sampling - We drew 1 ml samples of water into sterile
1.0 ml tuberculin syringes that were capped and transported to the lab-
oratory for dilution and inoculating of test cultures.
Time factor
General sampling - Except when conducting specific sequences of collec-
tions related to tidal factors (see description below) , samples were
collected at various times during daylight hours. Because of the proxi-
mity of the sampled area to the Jackson Estuarine Laboratory we were able
to examine collections for ciliates 30 min - 1 hr after collection.
Comparison of such collections with collections examined microscopically
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in the field immediately after sampling convinced us that there was no
significant loss - or multiplication - of protozoa during the interval
of transportation to the laboratory.
Relationship to tidal regime - We designed the timing of one series of
collections to allow determination of effects of full moon spring tides
and the resulting flooding of a pool in the upper marsh upon ciliate and
bacterial populations. In these instances, time of day of each collec-
tion was constant (11:00 a.m.) and the particular day of each collection
was chosen to allow collections both before and after initial flooding
of the pool.
Relationship to lunar regime - We designed three series of collections
to determine effects of full moon spring tides, neap tide, and new moon
spring tide upon diurnal changes in ciliate and bacterial populations in
a tidal channel in the lower marsh. We made three series of collections
on each of three days in which full moon spring, neap, and new moon
spring tides were predicted, and sampled the same station every three
hours over a 24 hr period.
Additional comments
We recorded all field data originally on mimeographed data sheets or on
IBM portable dictaphone tape for later transfer and storage on Keysort
punchcards (12.8 x 20.3 cm) .
LABORATORY METHODS
Chemical
We determined salinity in the laboratory by one of three methods. We
used a Hach chloride test kit, but found this less dependable than the
standard Knudsen titration method. During 1971, all our measurements
were made with an American Optical Company refractometer designed to
allow measurement directly of salinity in parts per thousand. With an
initial calibration to a water of known salinity as measured by the
Knudsen method, and occasional recalibration, this instrument allowed
determination of salinity on as little as 1 drop of water with an
accuracy of * 1 part per thousand.
Ciliates and Metazoa
In order to identify and count ciliates in quantitative samples, we
brought the original collection tubes to the Jackson Estuarine Laboratory
along with a 125 ml bottle filled with water from the same water mass as
that from which the collection tube was filled. After filtration, this
additional water served as diluent (see technique described below).
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We assembled in a vertical series a glass funnel to hold a square of
1 mm mesh Nitex gauze and 4 Buchner funnels fitted with Nitex gauze
circles of 505 urn, 253 jmr 153 urn, and 64 yai mesh size respectively,
held firmly in place with Tygon C-rings. These were spaced to allow use
of an eyedropper conveniently and so as to drain into a 150 ml beaker.
We began filtration by pouring the sample into the 1 mm Nitex gauze so
that any material that was held by the gauze was as close to the bottom
of the cone as possible. Pouring was at a sufficiently slow rate so as
to avoid overflow of the Buchner funnels. A volume of filtered water
from the habitat, of an amount 4 times that of the original collection,
was used to flush the filters to insure maximum passage of ciliates
through the filter series into the beaker below. Routine inspection of
filters indicated greater than 90% effectiveness of passage of ciliates
through the series, while allowing retention on the filters of most of
the organic debris and larger microorganisms that otherwise would inter-
fere with the counting process.
After filtration finished (about 10 min), we agitated the contents of
the beaker until any solid matter was homogeniously distributed and then
while still stirring vigorously, we poured 22.5 ml aliquots into 4-5
square plastic Petri dishes (Falcon No. 1012) inscribed with a grid of
36 squares, each 12 mm on edge. We prepared each Petri dish as a
counting chamber by marking on the lower side 5 squares (a central square
and 4 at the corners.)
Upon placing 22.5 ml of the filtrate into the counting chamber, we placed
the counting chamber on the stage of a dissecting microscope and allowed
it to stand a few minutes until any remaining debris settled. Proceeding
first with the central square and the remaining ones in a clockwise
direction, we counted all organisms observed and identified them as far
as possible. The data were recorded verbally on an IBM dictaphone and
later transcribed onto punchcards.
To corroborate identifications made during the analysis described above,
we maintained cultures of all collections and many isolated species of
ciliates. We employed various media, including rice grains, split peas,
and proteose peptone. Carnivorous forms were grown using Oxyrrhis marina
(a dinoflagellate) or small Uronematid ciliates as a food source. We
maintained all cultures in a room held at 20°C. We observed living
individuals with a dissecting microscope and bright-field microscopes,
corroborating cortical details by the nigrosinmercuric chloride formalin
method48,49 and the Chatton-Lwoff silver impregnation technic as modified
C /\
by Corliss130, and nuclear morphology by an iron hematoxylin method and a
modified Fuelgen nucleal reaction. We used a camera lucida, a calibrated
ocular micrometer, and a Nikon automatic photomicroscope employing high
contrast copy film to record observations of morphology. Because of the
importance of species identification to the fulfillment of this project,
13
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it became necessary to make fundamental taxonomic revisions, develop
family diagnoses, and construct binary keys to aid in identification of
ciliates to family, and then to species.
Permanent preparations containing type material of new species discovered
in this project have been deposited in the U.S. National Museum .
Bacteriological Methods
We estimated concentration of bacteria in the 1 ml samples described
above by a procedure involving dilution of the sample in a nutrient
liquid medium extended so that most of the tubes containing the highest
dilution remained sterile after incubation-^; specifically employing the
basal medium and extinction dilution method adopted for seawater
bacterial populations by Jannasch and
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SECTION V
EXPERIMENTAL PHASE
AUTECOLOGY
During 1970 and 1971, I identified 103 species of ciliates, representing
41 families in 10 orders. Nearly a third of these were poorly known and
four were new to science. These have been discussed in detail else-
whereSl. During the summer of 1970, 79 species were identified from 175
collections. During the summer of 1971, 82 species were identified from
102 collections. Species identified in summer collections represent
approximately 90% of the total number of species recorded from the
Adams Pt. tidal marsh. The most commonly encountered species in 1971,
Strombidium sulcatum, was identified in 73 of 102 collections. At the
other extreme, 20 of the species recorded during 1971 were collected only
once.
The following list employs the classification scheme used by Borror,
1973^4. The numbers following each species denote the number of records
during the summers of 1970 and 1971 respectively. Numbers indicated
with an asterisk are instances of species recorded other than during
those summers.
Members of some genera (Cyclidium, Euplotes, Pleuronema, and Vorticella)
were not identified to species during quantitative sampling.
Systematic list
Order Gymnostomatida
Amphileptidae
Cryptopharynx setigerus 1 2
Litonotus cygnus 2 1
Litonotus sp. 11 10
Loxophyllum chaetonotum 31 13
Loxophyllum setigerum 3 3
Chlamydodontidae
Chilodonella sp. 10
Chlamydodon lynchelliformis 9 2
Chlamydodon obliquus 6 0
Chlamydodon triquetris 3 2
Colepidae
Coleps tesselatus 49 42
Didiniidae
Askenasia stellaris 1 0
Mesodinium pulex 23 17
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Dysteriidae
Dysteria marina 4 5
Dysteria monostyla 3 5
Trochilla slgmoides 1*
recta 3 0
Enchelyidae
Chaenia sp. 7 4
Enchelyodon trepida 0 2
Holophrya coronata 1 1
Lacrymaria marina 17 8
Placus salinus 4 8
Metacystidae
Metacystis striata 4 4
Nassulidae
Nassula labiata 6 6
Nassula sp. 0 2
Prorodontidae
Prorodon marinus 29 29
Spathidiidae
Spathldixim sp. 7 4
S tephanopogonidae
Stephanopogon apogon 2 1
Trachelocercidae
Trachelonema oligostriata 2 17
Order Trichostomatida
Colpodidae
Colpoda cucullus 1*
Geleiidae
Geleia orbis 0 1
Geleia simplex 14 6
P lagiopylidae
Plagiopyla nasuta 10 2
Sonderia cyclostomata 0 1
Sonderia sinuata 10 12
Trimyemidae
Trimyema pleurispirale 0 1
Order Suctorida
Acinetidae
Acineta sp. 1*
Order Hymenostomatida
Cinetochilidae
Cinetochilnm margaritacevtm 0 1
Cohnilembidae
Cohnilembus verminias 3 4
16
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Frontoniidae
Frontonia fusca 0 3
Frontonia marina 40 38
Frontonia aicrostoma 2 0
Ophryoglenidae
Gphryoglena flava 2 0
Parameciidae
Paramecium calkinsi 7 1
Philasteridae
Paranophrys magna 1 3
Paraiironema acutum 2 5
Porpostoma notatum 4 4
P leuronematidae
Cyclidixmn marinum 27 24
Cyclidium plouneotiri 27 24
Pleuronema coronatum 17 4
Pleuronema small! 17 4
Pseudocohnllembidae
hargisi 1 0
Paralembus marinus 1 1
Pseudocohnilembias longisetus 1*
Tetrahymenidae
Paratetrahymena was si 1*
Uronematidae
Uronema filificum 25 33
Uronema marinum 8 5
Uronema sp. 0 1
Uropedalium pyriforme 7 9
Order Peritrichida
Vaginicolidae
Cothurnia simplex 1*
Vaginicola sp. 11
Vorticellidae
Vorticella nebulifera 12 5
Vorticella striata 12 5
Zoothamnium sp. 10
xmid. swarmer 0 2
Order Heterotrichida
Condylos tomatidae
Condylostoma arenarium 25 13
Metopidae
Metopus contortxis 2 8
Peritromidae
Peritromus faurei 28 15
17
-------
Spirostomatidae
Gruberia lanceolata 32 23
Parablepharisma bacteriophorum 4 7
Parablepharisma chlaTaydophorum 1 6
Protocruzia depressa 2 1
Spirostomum sp. 0 1
Order Odontostoraatida
Mylestomatidae
Mylestoma bipartitum 0 3
Order Oligotrichida
Halteriidae
Stroiribidium kahli 7 1
Strombiditrm la turn 16 4
Strombidixan purpureum 2 1
Stromibidi-am styliferum 12 12
Stroiribidium sulcatum 32 73
Stroiribidium viride 0 2
Strobilidiidae
Strobilidium caudatxam 0 2
Order Tintinnida
unid. sp. 33
Order Hypotrichida
Euplotidae
Aspidisca aculeata 21 12
Aspidisca baltica 0 1
Aspidisca sp. 1 1
Diophrys oligothrix 6 4
Diophrys scutum 12 1
Euplotes bisulcatus 0 1
Euplotes charon 1 2
Euplotes crassus 1*
Euplotes harpa 2 2
Euplotes quinquecarinatus 0 1
Euplotes trisulcatus 0 2
Uronychia transfuga 12 1
Holostichidae
Holosticha diademata 16 8
Paraholosticha polychaeta 1*
Trichotaxis pulchra 1*
Oxytrichidae
Gastrostyla pulchra 1*
Histriculus similis 1 2
Oxytricha halophila 0 3
18
-------
Psammomitra sp. 10
Tachysoma saltans 1*
Trachelostyla pediculiformis 11 7
Spirofilidae
Stichotricha sp. 10
Urostrongylum sp. 10
Correlation of Occurrences with Physical and Chemical Factors
Temperature - Since the overwhelming majority of collections were made
during the warmer months, over a relatively narrow temperature range, I
have attempted no correlations between ciliate occurrence or abundance
and temperature. Although a vertical temperature gradient was often
recorded in panne ponds (e.g. see Table 11), there was little evidence
that concomitant vertical differences in ciliate distribution were the
result of such temperature differences.
H2S concentration - Of 175 collections during the summer of 1970, H2S
concentration was measured in 48 instances. Most species occurred over
a wide range of I^S concentration (0-5 ppm) . In a few instances
(Gruberia lanceolata and Uronema filificum), the species were recorded
more regularly in areas of high sulfide levels (3-5 ppm). In a larger
number of instances (Coleps tesselatus, Loxophyllum chaetonotum,
Mesodinium pulex, Peritromus faurei, and Strombidium latum), there were
apparent negative correlations - the species usually recorded in samples
with relatively low (0-2 ppm) H_S concentration.
Hydrogen Ion Concentration - Although pH values in the marsh varied
considerably (6.3 - 9.5), analysis of pH values associated with occur-
rences of the most frequently occurring species (those occurring in at
least 10 samples each summer) reveal no significant correlation between
pH and ciliate occurrence. Such findings are not unexpected .
Salinity - Although rain water can collect in the panne ponds and tempo-
rarily reduce salinity in the tidal marsh, there are no permanent fresh
water streams draining into the sample area, hence salinities remain at
estuarine levels. During the summers of 1970 and 1971, most recorded
salinities were between 22 and 33 parts per thousand (e.g. see Tables 11-
14). No summer samples were taken with salinities below 16 parts per
thousand. Occurrence of the most frequently encountered species (e.g.
Coleps tesselatus and Mesodinium pulex) show no correlation with salinity
(see Table 1).
A frequently encountered hymenostome, Uronema filificum, generally
occurred more frequently, and reached higher concentrations in higher
salinities (greater than 28 parts per thousand) than a closely related
species, Uropedalium pyriforme (see Table 1). In 1970, they showed no
19
-------
overlap in distribution with relationship to salinity. In 1971, although
there was some overlap, Uropedalium pyriforme occurred with greater
regularity in waters of lower salinity than those inhabited by Uronema
filificum.
In a few instances (Prorodon marinus, Peritromus faurei, and Strombidium
laturn, styliferum and sulcatum) the species were recorded more frequently
in higher salinities (28 - 34 parts per thousand) than in waters of lower
salinity (see Table 1). Strombidium sulcatum, for which the most data
were available, seemed to show the strongest positive correlation with
salinity. The small number of samples at salinities below 20 parts per
thousand precluded any negative correlations between ciliate occurrence
and salinity.
Oxygen concentration - In general, the more regularly occurring species
showed no obvious correlation in either occurrence or in abundance with
02 concentration. In 1970 and 1971, Coleps tesselatus, Frontonia marina,
Gruberia lanceolata and Peritromus faurei occurred over a wide range in
02 concentration and reached high numbers in samples saturated with 02
as well as in samples devoid of 02. Table 2 illustrates a typical situ-
ation of a ciliate apparently highly tolerant of variations in oxygen
availability. Mesodinium pulex and Prorodon marinus distribution in 1970
showed a slight positive correlation to 02 concentration, but showed no
obvious correlation in 1971 (Table 3). In the case of Uronema filificum
(Table 4), both the 1970 and the 1971 data indicate that this ciliate
occurs more frequently, and reaches higher numbers the lower the oxygen
concentration, reaching its greatest abundance at 0-1 ppm. Less regu-
larly occurring species that similarly are tolerant of anoxic environ-
ments include Cohnilembus verminus, Histriculus similis, Paraplepharisma
spp., Metopus contortus, Mylestoma bipartitum, Sonderia sinuata,
Trachelonema oligostriata, and Uropedalium pyriforme. 1970 data indicate
also that Stephanopogon apogon and Parauronema acutum similarly occur
most frequently at low 02 concentrations.
Among the many species of the Oligotrich genus Strombidium, three of the
more regularly occurring species (S. laturn, styliferum, and sulcatum)
showed differences in tolerance to 02 concentration. S. sulcatum
apparently is tolerant to the entire range of 02 concentration, reaching
high numbers throughout the range (Table 5).
Strombidium styliferum is capable of reaching high numbers at low 02
concentrations, but at a lower frequency of occurrence (Table 6).
Strombidium latum is more regular at higher 02 concentrations, but not
at the highest levels (Table 7).
20
-------
to
H
TABLE 1. OCCURRENCE OF SEVERAL SPECIES AS A FUNCTION OF SALINITY
Salinity,
o/oo
>34
28-33
22-27
>34
28-33
22-27
a. number c
b. number c
Mesodinium pulex
1970
#a
3
4
6
1
RPb
20
18
33
13
1971
#a
0
11
3
0
RPb
0
19
15
0
Coleps tesselatus
1970 1971
#a RPb
12 80
13 59
5 31
4 50
Strombidium sulcatum
1970 .1971
#a
9
11
4
0
RPb
60
50
25
0
*a
7
44
15
2
RPb
100
75
75
2
*a
4
24
7
1
RPb
57
41
47
50
Uronema filif.
1970
#a
7
9
0
0
RPb
47
41
0
0
Strombidium styliferum
1970 1971
*a
4
3
1
0
RPb
27
14
7
0
#a
1
6
0
2
RPb
14
10
0
100
1971
#a
0
22
9
0
RPb
0
37
45
0
Uropedalium pyr.
1970 1971
#a
0
5
2
0
RPb #a RPb
000
23 3 5
13 5 25
000
Strombidium latum
1970 1971
#a
5
5
1
0
RPb
33
23
7
0
#a
1
1
2
0
RPb
13
2
10
0
f occurrences at that salinity.
>f occurrences expressed as a percentage of total samples at that oxygen concentration
-------
TABLE 2. COLEPS TESSEIATUS OCCURRENCE AND ABUNDANCE
AS A FUNCTION OF OXYGEN CONCENTRATION
Oxygen ,
ppm
>12
10-11
8-9
6-7
4-5
2-3
0-1
*a
6
0
5
3
4
2
6
^b
27
0
100
38
67
29
40
1970
cone . ,
cells/ 10 ml
40,60,20,
420,80,20
0
40,60,40,
180,880
40,120,180
20,60,80,100
60,200
20,40,60,
80,80,100
1971
#a
8
2
2
3
1
6
8
RPb
90
100
50
75
50
67
25
cone. ,
cells/10 ml
234,188,76
20,19,x,x
8,21
20,207
37,4, x
28
107,80,4,
32,6,18
230,400,80,
20,46,160,x,x
a. number of occurrences at that oxygen concentration
b. number of occurrences expressed as a percentage of total samples at
that oxygen concentration
22
-------
TABLE 3. MESODINIUM PULEX OCCURRENCE AND ABUNDANCE
AS A FUNCTION OP OXYGEN CONCENTRATION
Oxygen/
ppm
>12
10-11
8-9
6-7
4-5
2-3
0-1
1970
#a
6
0
1
1
1
1
3
RPb
35
0
20
13
17
14
20
cone . ,
cells/10 ml
1,260,60,
100,80,40
0
40
20
40
40
60,40,20
1971
#a
0
1
2
1
0
2
4
RPb
0
50
50
25
0
22
13
cone. ,
cells/10 ml
0
3
40,9
8
0
9,18
18,l,x,x
a. number of occurrences at that oxygen concentration
b. number of occurrences expressed as a percentage of total samples at
that oxygen concentration
23
-------
TABLE 4. URONEMA FILIFICUM OCCURRENCE AND ABUNDANCE
AS A FUNCTION'OP OXYGEN CONCENTRATION.
Oxygen,
ppm
>12
10-11
8-9
6-7
4-5
2-3
0-1
1970
#a
2
0
0
2
3
0
7
RPb
12
0
0
25
50
0
47
cone . ,
cells/10 ml
20,60
0
0
40,20
80,140,40
0
20,20,20,80
40,740,100
1971
#a
0
1
0
1
0
4
18
'
RPb
0
50
0
25
0
44
56
cone . ,
cells/10 ml
0
8
0
4
0
20,18,18,8
15,200,5,5,9,
36,4,400,18,
12,18,18,1600
x,x,x,x
a.
b.
number of occurrences at that oxygen concentration
number of occurrences expressed as a percentage of total samples at
that oxygen concentration
24
-------
TABLE 5. STROMBIDIUM SULCATUM OCCURRENCE AND ABUNDANCE
AS A FUNCTION OF OXYGEN CONCENTRATION.
Oxygen,
ppm
>14
12-13
10-11
8-9
6-7
4-5
2-3
0-1
1970
~#a
4
0
0
1
3
3
2
8
^b
24
0
0
20
38
50
29
53
cone . ,
cells/10 ml
40,300,400,
1500
0
0
60
20,40,80
60,60,160
100,180
1500,100,60,
220,20,180,
20,7900
1971
*a
3
2
2
3
2
2
9
27
^b
38
75
100
75
50
100
100
84
cone . ,
cells/10 ml
40, 565, x
20, x
2,35
420,468, x
52,1765
24,28
39,540,5,356,
10,12,920,71,
576
35,5,21,1360,
240,16,18,19,
14,600,800,
1250,1037,561,
336,40,7000,
60,8,9,4,9,15,
x,x,x
a. number of occurrences at that oxygen concentration
fa. number of occurrences expressed as a percentage of total samples at
that oxygen concentration
25
-------
TABLE 6. STROMBIDIUM STYLIFERUM OCCURRENCE AND ABUNDANCE
AS A FUNCTION OF OXYGEN CONCENTRATION.
Oxygen
ppm
>14
13-14
12-13
10-11
8-9
6-7
4-5
2-3
0-1
1970
*a
4
0
1
0
2
0
0
1
2
^b
24
0
33
0
40
0
0
14
13
cone . ,
cells/10 ml
1,40,60,260
0
20
0
20,20
0
0
40
40,60
1971
*a
3
1
1
0
0
0
0
1
3
^b
38
100
50
0
0
0
0
11
9
cone . ,
cells/10 ml
17,40,x
20
X
0
0
0
0
18
1,54,80
a. number of occurrences at that oxygen concentration
b. number of occurrences expressed as a percentage of total samples at
that oxygen concentration
26
-------
TABLE 7. STROMBIDIUM LATUM OCCURRENCE AND ABUNDANCE
AS A FUNCTION OF OXYGEN CONCENTRATION.
Oxygen,
ppm
"714
13-14
12-13
10-11
8-9
6-7
4-5
0-1
#a
1
0
2
0
2
2
2
2
1970
^b
6
0
67
0
40
25
33
13
cone . ,
cells/10 ml
180
0
140,800
0
20,40
20,140
20,80
20,700
1971
#a
0
1
1
1
0
0
0
2
^b
0
100
50
50
0
0
0
6
cone . ,
cells/10 ml
0
20
4
4
0
0
0
1,80
a. number of occurrences at that oxygen concentration
b. number of occurrences expressed as a percentage of total samples at
that oxygen concentration
27
-------
SYNECOLOGY
Faunal Differences Between Upper and Lower Marsh
As a result of an initial survey of the sampling area (Fig. 2), and
establishment of a map coordinate system for the grid of 10 m square
quadrats in the marsh, it was possible to determine microdistributional
differences for all species identified. Some similarities and differ-
ences of possible ecological significance were noted between ciliate
faunas of upper marsh and lower marsh (both terms defined in Materials
and Methods). Table 8 allows comparison of the degree of uniqueness of
ciliate faunas of these two zones based on nearly equal numbers of col-
lections. Table 9 allows comparison of locomotory types represented by
species unique to either the upper marsh or the lower marsh. For
instance, Strombidium purpureum and S^ viride are planktonic; that is,
they swim and feed independent of any substratum and often occur high in
the water column away from the bottom. They were identified only from
samples collected in the upper marsh. By contrast, Strombidium kahli,
Trimyema pleurispirale, tintinnids, and Peritrich swarmers, also plank-
tonic, were identified only in samples collected in the lower marsh.
In a number of instances (indicated by an asterisk following the name in
Table 9), there were cases of two or more members of the same genus
apparently separated microgeographically within the marsh. Although the
more benthic Strombidium sulcatum and S. latum were ubiquitous, three
other species were separated microgeographically. Two similar species
of Euplotes (see Table 9) were also apparently microgeographically sepa-
rated. These two species are so similar morphologically that greater
magnification than that used in quantitative examination of samples is
necessary for species identification. The microdistributional difference
between the two species of Dysteria is particularly intriguing inasmuch
as each was identified in five samples, one from the upper marsh and one
from the lower marsh. The morphological differences, as well as docu-
mented uniformity within a species is fairly well understood
Table 10 is a summary of a different type of analysis of faunal differ-
ences between the upper and lower marsh. In this instance, only those
species that occurred relatively frequently and in both halves of the
sampling area were analyzed.
Physical and Biological Parameters of a patens-panne Pond
Table 11 summarizes data compiled from collections made at approx. 11 a.m.
on July 5, 9, 12, and 16, 1971. On each day, I made nearly simultaneous
collections at the water surface, at the surface of the substratum, and
at midwater level near the edge of a panne pond in the upper marsh (right-
hand black square in Fig. 2). The first two collections, on July 5 and
28
-------
TABLE 8. PAUNAL DIFFERENCES BETWEEN UPPER AND LOWER MARSH.
Comparison of endemism of upper marsh (patens-panne zone) and lower marsh
(intertidal alterniflora zone).
Zones
upper marsh
lower marsh
total
# coll.
45
54
102
# species
recorded
82
83
99
unique species/
times recorded
1
10
11
2
5
1
3
0
2
4
0
3
5
1
1
endemism, %
19.5
21.7
29
-------
TABLE 9. LOCOMOTORY TYPES REPRESENTED BY SPECIES UNIQUE
TO EITHER UPPER MARSH OR LOWER MARSH
Locomotory
type
Species
Upper marsh
Lower marsh
sessile
thigmotactic
intermediate
planktonic
Cryptopharynx setigerus
Chlamydodon Lynchelliformis
Cinetochilum margaritaceum
Dysteria marina*
Euplotes quinquecarinatus *
Aspidisca baltica
Protocruzia depressa
Paralembus spp.
Paramecium calkinsi
SpriostomuHi sp.
Uronychia transfuga
Geleia orbis
Enchelyodon trepida
Plagiopyla nasuta
Strombidium purpureum*
Strombidixrai viride*
Vaginicola sp.
Litonotus cygnus
Stephanopogon apogon
Oxytricha halophila
Dysteria monostyla*
Euplotes bisulcatus*
Diophrys scutum
Holophrya coronata
Sonderia cyclostomata
Uronema sp.
Chaenia sp.
Cohnilembus verminus
Porpostoma notatum
Strombidium kahli*
Trimyema pleurispirale
tintinnids
peritrich swarmers
a member of at least two species in a genus separated
microgeographically
30
-------
TABLE 10. FAUNAE, DIFFERENCES BETWEEN UPPER AND LOWER MARSH.
Occurrences of regularly encountered species in upper marsh (patens-panne
zone) and lower marsh (intertidal altemiflora zone) .
Species
Strombidium sulcatum
Coleps tesselatus
Frontonia marina
Uronema filificum
Prorodon marinus
Gruberia lanceolata
Mesodinium pulex
Trachelonema oligostriata
Peritromus faurei
Condylostoma arenarium
Total number of
collections
Occui
Upper
marsh
29
27
12
7
12
7
9
4
8
8
45
rrences
Lower
marsh
44 '
15
26
26
17
16
8
13
7
5
57
Total
73
42
38
33
29
23
17
17
15
13
102
Chi- square
value
0.486
6.979*
2.360
6.920*
0.090
1.592
0.537
2.922
0.530
1.653
* significant at 0.01 level of probability (D.F. = 1)
31
-------
TABLE 11. PHYSICAL AND BIOLOGICAL PARAMETERS OF A PATENS-PANNE POND.
Effect of flooding by full noon spring high waters upon physical factors,
bacterial concentration, ciliates and micrometazoa.
Date,
1971
7/5
7/5
7/5
7/9
7/9
7/9
7/12
7/12
7/12
7/16
7/16
7/16
time
1030
1030
1030
1100
1100
1100
1055
1055
1055
1030
1030
1030
Depth
top
mid
bottom
top
mid
bottom
top
mid
bottom
top
mid
bottom
Temp., °C.
Air
29
29
29
26
26
26
26
26
26
28
28
28
H2°
29
27
26
26
26
27
25
24
24
29
30
29
°2'
ppm
sat
1.5
2.6
sat
2.1
0.0
sat
7.4
6.3
sat
10.7
3.6
pH
9.3
9.1
7.0
8.5
7.3
7.0
9.4
7.7
7.2
8.4
8.0
8.2
S, o/oo
-
-
-
35
34
37
30
29
30
33
33
33
32
-------
TABLE 11, cont'd. PHYSICAL AND BIOLOGICAL PARAMETERS OF A PATENS-PANNE
POND.
Date,
1971
7/5
7/5
7/5
7/9
7/9
7/9
7/12
7/12
7/12
7/16
7/16
7/16
Depth
top
mid
bottom
top
mid
bottom
top
mid
bottom
top
mid
bottom
Bacteria
x 10 cells
per ml
1.1
1.5
2.4
10.0
4.9
1.3
10.0
1.3
2.4
0.49
2.9
2.4
Organisms ,
number/ ml
Ciliates
56.0
7.8
8.1
-
70.7
34.0
110.7
18.7
21.1
54.4
7.1
87.2
01
0)
Dinoflagellat
80
10
68
-
-
150
-
216
204
1200
310
445
Nematodes
8
-
-
10
-
-
9
-
-
24
1
7
Copepods
--
-
1.8
8.0
0.8
-
-
-
-
-
—
—
Nauplii
-
0.4
1.4
-
1.2
4.0
-
-
-
-
-
-
Gastrotrichs
-
-
0.45
-
-
-
1.7
-
-
-
-
-
Rotifers
6
-
-—
16
-
-
-
-
-
8
1
-
Alleocoeles
-
-
-
-
-
-
6.8
-
-
4.8
-
-
Veligers
-
-
-
-
0.4
-
-
-
-
-
-
-
0)
(0
•o
-
-
-
-
-
-
-
-
-
1.6
-
—
33
-------
9, were taken toward the end of a 2-week period dominated by a neap tide
cycle, during which no water exchange occurred between the panne pond and
the ebb and flow of the tide. In other words, the station was a tidal
pool isolated from the effects of tidal.flushing over a period in excess
of a week. The samples of July 12 were collected following a session of
full moon spring tides whose amplitude (+ 2.47 m) exceeded the height of
the pool level (+ 2.41 m), thus allowing influx of tidal water from the
estuary, and a resulting mixing of the pool water. Direct microscopic
counts of ciliates, dinoflagellates, and micrometazoa were then compared
with estimates of bacterial abundance and summarized in Table 11.
Physical and Chemical Factors - Except for a brief increase in depth at
time of actual flushing at high tide, water depth at the station remained
constant throughout the series of samples. We made all collections at
the same time of day. There was relatively little change in air and
water temperature' during the period. Although salinity data are incom-
plete, salinity apparently fell during flushing, suggesting that evapor-
ation during the period prior to flushing had led to higher salinities
than those in the estuary. Hydrogen ion concentration generally
decreased with depth in the pool and was most variable in mid-water. It
was relatively constant at the top and bottom stations, and did not show
correlation with flushing of the spring tides. Oxygen was at saturation
or above throughout the series, due to photosynthetic activity of the
heavy mats of Cladophora near the water surface. At middle levels and
at the bottom, 02 concentration was relatively low before flushing and
rose after flushing by the spring tide.
Biological Factors - Numbers of species of ciliates were highest in the
surface mats of algae and on the surface of the substratum, correlated
with increased niche diversity. Ciliate numbers were lowest in mid-water
where Strombidium sulc_atum dominated. There appeared to be an increase
in diversity on the bottom following flushing, associated with a rise in
Q£ concentration there, with dinoflagellates and bacteria about constant.
Concentrations of ciliates were highest at the surface and on the bottom.
A pulse in concentration in midwater occurred following the 02 increase
associated with flushing.
Dinoflagellate numbers increased slightly after flushing. Their abun-
dance apparently was not related to their position in the water column
except one pulse in the Cladophora mat. There were as many dinoflagel-
lates at mid-water as on the bottom.
Numbers of copepods and copepod nauplii diminished upon flushing. There
appeared to be no effect of flushing on rotifers and nematodes; they
occurred primarily within interstices of the Cladophora mat. One veliger
was seen in mid-water before flushing. Flatworms were not observed
before flushing, but were seen on the bottom afterwards.
34
-------
There appeared to be no effects of tidal flushing upon bacterial numbers
as measured by Most Probable Number counts in our dilution series. The
only exception was a pulse in bacterial numbers in the Cladophora. mat.
Numbers of bacteria remained fairly constant at mid-water and on the
bottom.
Physical and Biological Parameters in the Lower Marsh During a Tide
Effects of a Full Moon Spring Tide - Table 12f summarizes data from
analysis of a series of samples collected every three hours on 6 July
1971 at the time of full moon spring tides. Samples from the water
surface and the substratum-water interface of a tidal channel (at a
station indicated by the black square in the lower marsh in Fig. 2) were
examined microscopically for ciliates, dinoflagellates, and micrometazoa,
and correlated with bacterial abundance as determined by Most Probable
Number analysis (see Materials and Methods section). The collection
series allowed comparison of numbers of organisms in the water surging
through the tidal channel on the ebb and the flood. Simultaneous with
collections at the lower marsh station, we also analyzed a sample from
the extreme lower end of the tidal marsh beyond the extent of the
Spartina peat (southwest beyond the area illustrated in Fig. 2). We
made no collections at high water since at that time water was not con-
fined simply to the tidal ditches, but covered the platform entirely.
Physical and Chemical Factors - During the two full moon spring tidal
cycles of July 6, water temperature was uniformly 3 - 9°C lower in the
forenoon, particularly on the ebb or at low water. 02 cone, remained
uniformly generally nearly zero, rising to about 8 ppm in the evening
flood tide. Salinity ranged between 25-29 parts per thousand, with
little regular variation. The pH at the water-substratum interface was
generally approx. 0.2 units lower than at the water surface. The water
coursing through the tidal channel on the ebb tide usually was about
0.5 - 1.0 units lower than the water at flood tide. Intermediate values
were recorded at low tide.
Biological Factors - Ciliates in the tidal channel in the lower marsh
were considerably more numerous, and usually represented by more species
along the substratum-water interface than at the water surface. Ciliate
numbers at the water surface usually were below that reached by popula-
tions in the gatens-panne pond. In general, there were higher numbers
of species, at greater concentrations, on the ebb tide than on the flood.
Surface mud of the channel station at the lower edge of the tidal marsh
contained considerably fewer species of ciliates and sometimes none at
all.
Dinoflagellates were not present in significant numbers in any of the
collections. Micrometazoa essentially were restricted to the substratum-
water interface.
35
-------
TABLE 12. PHYSICAL AND BIOLOGICAL PARAMETERS IN THE LOWER MARSH DURING
A TIDE. Effect of a full moon spring tide cycle upon physical factors,
bacterial concentration, ciliates, and micrometazoa.
Tide
flood
flood
flood
flood
flood
ebb
ebb
ebb
ebb
ebb
low
low
low
low
low
Depth
top
top
bottom
bottom
channel
top
top
bottom
bottom
channel
top
top
bottom
bottom
channel
Time
1000
2200
1000
2200
1030
0400
1540
0400
1540
1540
0640
1900
0640
1900
0640
H20
Temp. ,
°C.
22
25
22
26
-
20
28
21
27
-
18
27
18
26
-
°2
ppm
0
7.5
0
8.5
-
0
0
0
2.1
-
1.0
1.0
1.0
0
-
PH
7.3
7.7
7.2
7.3
-
6.7
6.8
6.1
6.9
-
7.3
6.9
7.1
6.8
-
S
o/oo
26
27
-
27
28
29
26
-
27
28
25
26
27
26
27
01
0)
-H
0
0)
CO
0)
.p
a
•rl
-H
o
3
1
8
12
2
12
7
14
18
0
7
4
8
15
10
Organisms/ml
r-
o
H
X
^
td
•rl
M
O
(d
n
—
-
0.4<
2.2
-
-
-
1.1
1.7
-
-
-
-
-
—
(0
0)
4J
id
•H
i-l
•H
0
2.2
0.5
7.2
88.0
10.0
1.5
6.2
5.6
106
-
2.3
3.2
15.8
384
1.2
<>i
•4J
a
i—4
H
0)
id
H
O
-S
Q
1
-
-
*•!
12
1
-
1
—
-
-
-
-
—
-
10
0)
•d
o
1
-
-
60
-
-
1
-
-
2
-
—
-
1
-
-
id
•H
id
o
H
-------
There was no significant correlation between numbers of ciliates and
bacterial concentration.
Effects of a Neap Tide - We collected a series of samples every three
hours during 29 July 1971 in a manner similar to that described for the
above series. They were designed to determine the effect of neap tidal
cycles upon relative abundance of ciliates, dinoflagellates, micrometa-
zoa, and bacteria in a lower marsh tidal channel on the ebb and on the
flood. At each sampling time, we collected a sample from the water sur-
face and from the water—substratum interface at the station on the lower
marsh indicated by a black rectangle in Fig. 2, and simultaneously from
a station 90 m SW from that station, at a point just beyond the extent
of the Spartina peat. As before, no collections were made at high water,
since at that tine water extends beyond the top of the tidal channel,
completely inundating the sampling platform. Physical and biological
data are summarized in Table 13.
Physical and Chemical Factors - 02 was uniformly nearly absent. Salinity
was nearly constant, at 28 parts per thousand. There was little vertical
difference, and little understandable diurnal difference in pH. Water
temperature was uniformly much lower in the forenoon, particularly at
the time of the morning flood tide.
Biological Factors - Ciliates were present in greater variety (5-8
species) at the bottom than at the surface (1-4 species), and at greater
concentrations at the bottom on the ebb tide than on the flood tide. As
was the case during a springtide cycle, there again were relatively few
ciliates at the channel station. There was no significant correlation
between ciliate concentration and bacterial concentration. Bacteria
were low and constant in numbers, about an order of magnitude lower than
levels estimated for samples collected during the full moon spring tide
cycles (July 6 data discussed above). Relatively few micrometazoa were
observed, either at the water surface or at the substratum-water inter-
face.
Effect of a New Moon Spring Tide - We collected a series of samples every
3 hrs during a 24-hr period on August 17 and 18, 1971, to determine the
effects of new moon spring tidal cycles on the ebb and flow of ciliates,
bacteria, dinoflagellates, and micrometazoa in the tidal marsh ditch
described above. Since analysis of samples from the water surface in
the previous two experiments indicated low and insignificant populations
of ciliates and micrometazoa, we sampled only the substratum interface
in this series. As before, we also sampled at the substratum surface at
a point 90 m SW of the lower marsh station. Physical and biological data
are summarized in Table 14.
37
-------
TABLE 13. PHYSICAL AND BIOLOGICAL PARAMETERS IN THE LOWER MARSH DURING
A TIDE. Effect of a neap tide cycle upon physical factors, bacterial
concentration, ciliates, and micrometazoa.
Tide
flood
flood
flood
flood
flood
ebb
ebb
ebb
ebb
low
low
low
low
low
Depth
top
top
bottom
bottom
channe]
top
top
bottom
bottom
top
top
bottom
bottom
channel
Time
0330
1550
0330
1550
1550
0940
2100
0940
2100
0035
1255
0035
1255
1255
HoO
"2
Temp. ,
°C.
16.8
28
19.6
27
-
21
24
-
24.6
18.5
26
21
25
-
°2
ppm
0
4.3
0
0
-
1.0
0
-
0
1
0
-
2.4
-
pH
-
6.8
-
7.3
-
8.6
6.8
-
6.7
7.6
7.4
7.4
8.0
-
S
o/oo
28
28
28
28
28
28
30
28
28
28
28
28
28
28
*
01
0)
•H
O
0)
ft
01
G)
I)
(o
•H
H
•rj
U
4
1
5
8
2
1
4
8
7
3
2
7
5
5
Organisms/ml
^
o
H
X
^
Q
•p
jrt
i
&
-
-
i
16
-
-
-
3.4
-
-
-
-
1.6
-
•H
"M
H
Qi
3
%
-
-
-
-
-
1
-
1.7
-
-
-
-
-
-
(d
H
M
(d
o
a)
u
-
-
-
-
-
-
-
-
-
1
-
-
-
-
01
fi
o
•H
M
4J
O
-P
01
id
0
-
-
-
-
-
-
-
-
-
-
-
-
1.6
-
38
-------
TABLE 14. PHYSICAL AND BIOLOGICAL PARAMETERS, LOWER MARSH, DURING A TIDE.
Effect of a new moon spring tide cycle upon physical factors, bacterial concentration, ciliates,
and micrometazoa. Samples taken from bottom.
w * Tide
flood
flood
channel
ebb
ebb
low
low
channel
Time
0930
2040
0930
0330
1430
0630
1740
0630
H20
Temp. ,
-
22
•H
20
26
18.5
24
_
o2,
ppm
-
0.5
— —
0
3.0
1.1
0
2.1
PH
-
6.2
_
6.3
6.1
6.5
6.1
6.5 ,
s,
o/oo
28.5
28.5
30
29.5
29.5
28
29
29.5
1
*
CO
(U
•H
o
0)
0)
-P
•H
•H
•H
U
3
6
1
7
10
7
11
6
Organisms /ml
r^
o
rH
X
(0
•H
-------
Physical and Chemical Factors - 0- cone, was uniformly low, as in the
previous series. Again salinity was constant at 28.5 - 30 parts per
thousand. Hydrogen ion cone, was relatively stable (6.1 - 6.5).
Water temperature fluctuated considerably during the day rising from an
early morning low of 18.5°C at 6:30 a.m. to a high of 26°C at 2:30 p.m.,
and then gradually decreasing during the evening.
Biological Factors - Ciliates were present in greater variety and in
greater concentrations at ebb tide and at low tide than when the channel
was subjected to an inflooding of water from the estuary. Estimates of
bacterial abundance from samples collected from this series indicate
slightly higher concentrations on the ebb tide than on the flood tide,
but relatively little correlation between ciliate concentration and
bacterial concentration. As before, there were relatively few micro-
ma tazoa, in relatively low numbers.
40
-------
SECTION IV
DISCUSSION
Because of preliminary indications from previous work that closely
related and morphologically similar species may differ significantly in
microdistribution27 a particularly careful and extensive assessment had
to be made of the taxonomic position of ciliates encountered. To esta-
blish a firm basis for discussion of both autecological and synecological
roles of the tidal marsh ciliates, extensive research was conducted under
this grant on the ciliate taxonomy^ to the extent of the necessity of
major revision of one order **. With this background of understanding,
it was then possible to describe, in more detail than is possible here,
the anatomy, behavior, autecology, and systematic position of many of
the tidal marsh ciliates encountered in this study, including 29 species
of relatively poorly known forms, including 4 new species^l. Thus these
data may serve as a baseline for judging protozoan diversity and commu-
nity structure in marine habitats of differing water quality.
Where ciliate genera are represented in the tidal marsh by several
species there were often differences in their relationship to physical
and chemical factors. These are most notable within the genus
Strombidium. Such a phenomenon is to be expected, and parallels
previously published data in intrageneric differences in saprobial
valance among genera of fresh water ciliates . Whether or not these
situations are interpreted as examples of competitive exclusion, they
corrobrate previously published conclusions27 and further justify care-
ful identification procedures used in ecological studies of protozoa.
The ciliate fauna displays a wide array of locomotory types, sensitivity
to various chemical and physical factors, and morphological adaptations.
With the exception of a very few species known heretofore primarily only
from fresh water (e.g. Colpoda cucullus, Ophryoglena flava, and
Spriostomum sp.), the tidal marsh ciliate fauna is essentially a marine
assemblage. In general, the locomotory types represented by the species
unique to either the upper marsh or the lower marsh (Table 9) were
similar; that is, both microhabitats supported a similar number of thig-
motactic and planktonic species, while supporting as well a large number
of forms of intermediate locomotory type.
Table 10 indicates that of the 10 most regularly encountered species in
the upper and lower marsh, only two (Coleps tesselatus and Uronema
filificum) showed statistically significant differences in microdistri-
bution. Coleps tesselatus occurred more frequently than expected in the
upper marsh while Uronema filificum occurred more frequently in samples
from the lower marsh. It is possible that the unusual capacity of U.
filificum to become temporarily sessile by means of temporary attachment
41
-------
to the substratum by excretions of its mucocysts may allow members of
the species to maintain their position in that habitat and avoid being
swept away by tidal currents. Coleps tesselatus on the other hand con-
gregates along substrata to feed upon bacteria attached to the substratum
and are thus perhaps more likely to occur in the quieter pools of the
upper marsh. The remaining species in Table 10 are either strongly
thigmotactic (e.g. Trachelonema oligostriata) or are free-swimming,
feeding independent of the substratum (e.g. S_. sulcatum). Judging from
the similar number of species recorded in each habitat and the similarity
of the percent of endemism in the two areas, the diversities of ciliates
in the upper and lower marsh appear similar. In general these points
are supported by data collected in both the summer of 1970 and the summer
of 1971.
Despite elaborate experimental design to allow correlation of ciliate
and bacterial concentrations in a patens-panne pond during a period of
flushing by a spring tide cycle, little obvious relation was evident.
Although there was an increase in diversity of ciliates along the bottom
following the flushing of the pond there appeared to be no obvious link
with bacterial concentration. In retrospect, it seems likely that bac-
terial standing crop is such a dynamic phenomenon that one would be
required to sample the population more frequently - if not continuously -
to be able to document the actual relationships of the change of the
water mass of the pool following flushing, changes in bacterial popu-
lations, and changes in ciliate populations. Although direct microscopic
methods for enumeration of bacteria were considered to have sufficient
complications that they were not employed in this study, it may be that
their advantage in revealing a more exact count of bacteria in a sample
might render them useful in future research in this area. Although the
Most Probable Number method employed here would allow gross comparisons
with other work, direct observation counts may be necessary to evaluate
such dynamic changes as may occur during a tidal change.
A number of generalities emerge from measurement of physical and biologi-
cal parameters in the lower marsh during a full moon spring tide, a neap
tide, and a new moon spring tide (Tables 12-14). In general, ciliates
were present in greater variety and in larger numbers at ebb tide and low
tide than when the channel was subjected to an inflooding of water from
the estuary, regardless of the tidal amplitude.
Ciliate populations high in the water column or near the water surface
in the tidal channel in the lower marsh generally either were absent or
at relatively low concentrations. However, greater numbers and variety
were evident during flood tide than during ebb or low tide. Diversity
was higher during a spring tide cycle than during a neap tide cycle.
Of all the ciliates occurring in both the upper and lower marsh, only
Coleps tesselatus appeared to occur at a statistically significant higher
42
-------
level in the upper marsh and therefore might be looked upon as an upper
marsh form. Counts made in the lower marsh across tidal cycles further
corroborate this point. The overall frequency of C_. tesselatus in the
collections summarized in Tables 12-14 was very close to the total
percent from all combined lower marsh stations (about 25%). Yet the
percentage seen at low tide (38.5%) is higher, and the percentage at
ebb (20%) and flood (15.4%) is lower than expected. It is possible that
at spring high tide, C_. tesselatus is flushed from the panne pools down
through the channels of the lower marsh and then lost to the estuary
with relatively little recruitment to the marsh during the subsequent
flooding.
Concomitant sampling from surface mud at the lower edge of the tidal
marsh below the channel station consistently showed fewer species of
ciliates than the tidal channel station, and sometimes none at all.
This finding, along with vertical differences in concentration of
ciliates mentioned above, make it appear plausible that ciliates flushed
by tidal currents from the upper parts of the marsh settle out on detri-
tus at the bottom of channels rather than remaining higher in the water
column and being swept from the marsh in the plankton. Thus it appears
most likely that tidal marsh ciliates enter estuarine food webs beyond
the marsh by their association with particulate detritus, as has been
postulated previously". Ciliates observed high in the water column in
tidal channels during flood tide generally represent planktonic forms
(e.g. Strobilidium sp., and tintinnids) not ordinarily found in associ-
ation with substrata higher in the marsh. Ciliate numbers in surface
samples in tidal channels during a tide were generally higher during
flood tide than during ebb tide reflecting the likely settling out of
marsh ciliates during the ebb and an influx of planktonic ciliates from
the estuary during flood tide.
Although any attempt to quantify field populations of microorganisms is
likely to be fraught with difficulty, and often involves tedious methods,
the vertical filter series and subsequent counting method was a practical
approach, and yielded consistent results, unlike other methods for
sampling ciliates of marine habitats58,59 ^e method is suitable for
habitats with considerable organic matter, and avoids loss of relatively
thigmotactic forms. Corroboration of direct counts of living ciliates
by rapid staining methods such as the nigrosin-mercuric chloride-formalin
method allows the necessary viewing of cytoplasmic detail and permanent
record to document identification.
43
-------
SECTION VII
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Danzig Bay I. The protozoa in some biotopes of the seacoast.J Polsk.
Arch. Hydrobiol. 1£(23):39-109, 1962.
43. Czapik, A. Mikrofauna slonawego jeziora Ptasi Raj. (Microfauna of
brackish lake Ptasi Raj/) Polsk. Arch. Hydrobiol. 10(23):371-378,
1962.
44. Dietz, G. Beitrag zur Kenntnis der Ciliatenfauna einiger Brack-
wassertumpel (etangs) der Franzosischen Mittelmeerkuste. [contri-
bution to the knowledge of the ciliate fauna of some brackish ponds
of the French Mediterranian coast.3 Vie et Milieu 15_: 47-93, 1964.
45. Lackey, J. B. Bottom sampling and environmental niches. Limnol.
Oceanogr. £(3):271-279, 1961.
46. Reuter, J. Einige faunistische und okologische Beobachtungen uber
Felsentumpel-Ziliaten. £some faunistic and ecological observations
on rockpool ciliates^ Acta Zool. Fennica 99_:l-42, 1961.
47. Burke, J. D. Determination of oxygen in water using a 10-ml.
syringe. J. Mitchell Soc. p. 145-147, November 1962.
48. Borror, A. C. Nigrosin-HgCl2-Formalin; a stain-fixative for
ciliates (Protozoa, Ciliophora) . Stain Technol. 43_(5) : 293-294,
1968.
49. Borror, A. C. Application of the stain-fixative nigrosin-HgCl--
formalin to fragile or contractile ciliates. Trans. Amer.
Microscop. Soc. 88(3):454-458, 1969.
47
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50. Corliss, J. 0. Silver impregnation of ciliated protozoa by the
Chatton-Lwoff technic. Stain Technol. ^8_: 97-100, 1953.
51. Borror, A. C. Tidal marsh ciliates (Protozoa): morphology,
ecology, systematics. Acta Protozool. 1£(2):29-71, 1972.
52, Porter, J. R. Bacterial chemistry and physiology. New York,
John Wiley and Sons, Inc., 1946. 1073 p.
53. Jannasch, H. W. and G. E. Jones. Bacterial populations in seawater
as determined -by different methods of enumeration. Limnol. Oceanogr.
4_(1): 128-139, 159.
54. Borror, A. C. Marine flora and fauna of the northeastern United
States. Protozoa: Ciliophora. NOAA Tech. Rept. NMFS Circ-378,
September 1973. 62 p.
55. Borror, A. C. Revision of the order Hypotrichida (Ciliophora,
Protozoa). J. Protozool. JL9_(1) :l-23, 1972.
56. Bick, H. and S. Kunze. Eine Zusammenstellung von autdkologischen
und saprobiologischen Befunden an Sttsswasserciliaten. [A review
of autecological arid saprobiological data on freshwater ciliates.]
Int. Rev. ges. Hydrobiol. _56_(3) : 337-384, 1971.
57. Fenchel, T. Studies on the decomposition of organic detritus
derived from the turtle grass Thalassia testudinum. Limnol.
Oceanogr. 15_(1) :14-20, 1970.
58. Fjeld, P. On some psammobiotic ciliates from DrjJbak (Norway).
Nytt Mag. Zool. _3:5~65> 1955.
59. Uhlig, G. Untersuchungen zur Extraktion der vagilen Mikrofauna aus
marinen Sedimenten. [Investigation on extraction of vagile micro-
fauna from marine sediments.] Verh. Deutsch. Zool. Ges. 10:151-157,
1965.
48
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SECTION VIII
LIST OF PUBLICATIONS
Borror, A. C. Revision of the order Hypotrichida (Ciliophora, Protozoa)
J. Protozool. 19/1):l-23, 1972.
Borror, A. C. Tidal marsh ciliates (Protozoa): morphology, ecology,
systematic^. Acta Protozool. 10^(2) :29-71, 1972.
Borror, A. C. Marine flora and fauna of the northeastern United States.
Protozoa: Ciliophora. NOAA Tech. Report NMFS Circ-378, 62 pp. 1973.
49
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SELECTED WATER
RESOURCES ABSTRACTS
INPUT TRANSACTION FORM
J,
3, Accession No,
w
4. Title
Environmental requirements of selected estuarine
ciliated protozoa
S. Report £«?«
6.
8. Perfofvu$M- OtgaHiisati&a
7. Author(s)
Arthur C. Borror
10. Project No.
18080 FBW
9. Organization
Department of Zoology and the Jackson Estuarine Lab.
University of New Hampshire
Durham, New Hampshire 0382^
11, Contract/ Grant No.
13. Type of Report and
Period Covered
12. Sponsoring Of gardKtion^ Off lee Of Res. & tfonltprlnf
15. Supplementary Notes
16. Abstract Measurements of temperature, pH, oxygen concentration, HpS concentration,
salinity, bacterial concentration, occurrence of micrometazoa, and distribution and
abundance of ciliated protozoa were recorded during the summers of 1970 and 1971 in a
tidal marsh at Adams Pt,, Durham, New Hampshire,
Numerous differences in ciliate distribution occurred between the upper (Spartina
patens) marsh and the lower (Spartina alterniflora.) marsh. Physical and biological
parameters of a patens~panne pond were measured during a 2-week period involving the
initial flushing of the pool by a session of full moon spring tides. Effects of a full
moon spring tide cycle, a neap tide cycle, and a new moon fpring tide cycle for one
station in the lower marsh were evaluated.
During 1970 and 1971, 103 species of ciliates, representing 4l families and 10
orders were identified, including b new species. Several instances of correlations
between ciliate abundance and oxygen concentration, ^S concentration, and-salinity
were observed.
Contribution of tidal marsh ciliates in general to estuarine food webs
probably occurs through their association with particulate detritus.
This report was submitted in fulfillment of Project Number 18080 FEW by Arthur
C. Borror under the sponsorship of the Environmental Protection Agency. Work was
completed as of September, 1971*
17a. Descriptors
estuarine, ciliate, protozoa, tidal marsh, distribution, ecology
Ub. Identifiers
17c. COWRR Field & Group
18. A variability
19. ' Security €fassf'
(Report)
20. Security Class,
(Pigs)
2f. Mo, Of
Pages
22. Price
Send To:
WATER RESOURCES SCIENTIFIC INFORMATION CENTER
U.S. DEPARTMENT OF THE INTERIOR
WASHINGTON, D. C. 2O24O
Abstractor
\ Institution
WRSIC 102 (REV. JUNE 1971)
U. S. GOVERNMENT PRINTING OFFICE: 1975-698-401/132 REGION 10
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