BULLETIN

OF MARINE SCIENCE, 29(4): 459-464, 1979

EPIPHYTIC DIATOMS OF THREE SEAGRASS SPECIES IN

MISSISSIPPI SOUND
Michael

J. Sullivan

ABSTRACT
Epiphytic diatoms were collected on 20 July 1977from the leaves of three seagrass species
(Halodule beaudettei, Cymodoceafiliforme, and Thalassia les(udinum) in Mississippi Sound,
U.S.A. Examination of cleaned material under the light and scanning electron microscope
revealed a total of 45 diatom taxa epiphytic on the three seagrasses. The four most abundant
diatoms were Fragilaria hyalina, Maslogloia pusilla, Ucmophora cr. debilis, and Opephora
pacifica. Except for the rare taxa, the taxonomic composition was identical regardless of the
seagrass species examined. Based on counts of 1,000 valves, values of community diversity
statistics (H', S, and R') were virtually identical for the three epiphytic diatom samples. Use
of a selected similarity index (SIMI) revealed that any two samples shared between 82 and
88% of the maximum similarity possible. Therefore, the available taxonomic and structural
information indicated that the above three seagrasses supported a single, nearly homogeneous epiphytic diatom community at the time of sampling.

Seagrasses are highly productive marine angiosperms which produce annual

crops of leaves from perennial rhizomes buried in the sediment. Depending on
local conditions, various seagrass beds may support a prolific epiphytic algal flora
on the leaves. Such severe epiphytism may lead to significant decreases in seagrass productivity (Sand-Jensen, 1977). Recent work on Zostera marina L. beds
in North Carolina waters has shown that over a one-year period algal epiphytes
accounted for 24% of the total biomass and ]8% of the total productivity on an
areal basis of the seagrass-epiphyte community (Penhale, 1977). The presence of
algal epiphytes should presumably increase the nutritive value and utilization of
seagrass species by various faunal elments. Wood (1959) found that diatoms epiphytic on Australian seagrasses were an important food source for phytophagous
fish and other animals over a yearly cycle.
Published accounts of those algal species epiphytic on seagrasses within waters
of the continental United States have been rare indeed. Humm (1964) and Ballantine (1972) reported on blue-green, red, brown, and green algae epiphytic on
four seagrass species off the east and west coast of Florida, respectively. The
diatom component of seagrass epiphytic floras has received attention in only one
study on the Pacific coast (Main and McIntire, 1974) and two on the Atlantic
coast (Reyes-Vasquez, 1970; Sullivan, 1977). Therefore, our knowledge of the
diatom flora epiphytic on seagrasses in the Gulf of Mexico is nonexistent. The
present paper presents some preliminary information as a first step in alleviating
this unfortunate situation.
DESCRIPTION

OF STUDY AREA

Within Mississippi Sound seagrass beds are primarily found associated with a sandy boltom and
shallow waters along the northern shores of the four barrier islands: Cat, Ship, Horn, and Petit Bois.
The dominant seagrass species are shoal grass, Halodule beaudettei (Den Hartog) Den Hartog; turtle
grass, Thalassia lesludinum Konig; and to a lesser extent manatee grass, Cymodoceafiliforme (Klitz,)
Correll (=Syringodium filiforme Klitz.). The study area was located approximately 0.3 km north of
Horn Island. Water depth over the seagrass beds studied ranges from 0.8 to 1.4 m below MLW.
Summer water salinities in the study area generally range from 20 to 35 and average about 30"100. All
three seagrass species were present in the study area forming pure or occasional mixed beds of
variable size.

459

BULLETIN

460

OF MARINE SCIENCE, VOL. 29, NO.4,

]979

METHODS

A single preliminary collection of epiphytic diatoms from the three seagrass species was made on
20 July 1977. The salinity of Mississippi Sound on this particular date was 34%0 following a drought
of approximately 2 months duration. Heavily epiphytized leaves were removed by hand from three
individual beds, each bed being composed of a single seagrass species, Virtually all leaves present in
each bed were so completely covered with a dense growth of Ectocarplls and diatoms that the green
color of the seagrass was all but undetectable. Epiphytic diatoms were removed by scraping individual
leaves with a spatula. The leaves were examined visually following epiphyte removal and none appeared to be senescent. Three epiphytic diatom samples were thus obtained, each one representative
of a single seagrass species. Each sample was then boiled in nitric acid and potassium dichromate to
oxidize the organic matter and prepare the diatom valves for observation under both the light (LM)
and scanning electron microscope (SEM). One-half of the cleaned material in each sample was mounted in Hyrax (LM) while the remaining half was air-dried on aluminum stubs and then coated with
gold in a sputtering device (Technics, Inc.) for observation in a Hitachi HHS-2R SEM operated at an
accelerating voltage of 20 KV. Although all counts for the data analysis were made with the LM,
identification of some of the diatom taxa would have been difficult, if not impossible, in the absence
of SEM observations.

Data Analysis
At least 1,000 diatom valves were counted and identified to species or varietal rank in each of the
three samples. This number represented a small percentage of the total number of valves present in
a given sample. After each sample had been analyzed taxonomically, community diversity statistics
were calculated. The first of these was the Information index (Shannon and Weaver, 1949):

where H' (species diversity) is expressed as bits/individual, n is the number of individuals of the
i-th taxon, N is the total number of individuals, and S is the total number of taxa in the sample. The
second index of community diversity calculated was redundancy (Main and McIntire, 1974):
j

R' = H'm.x - H'

H'max - H'min

'

where
H'max

log,S,

and

N-S+l)

H'mln = 10g,N -

10g,(N - S

I).

R' has no units and is a useful measure of the relative degree of dominance in the sample. Values of
R' range from 0, when all taxa are equally abundant, to I, when all taxa except one are represented
by a single individual. To compare the structure of selected pairs of diatom communities the following
similarity index proposed by Stander (1970) was employed:
s

2:

PUPln

1=1

SIMI =

Pu'
~1~1

~It

PIn'

where Pi) and Pin are the proportions of the i-th taxon in the j-th and n-th samples, respectively, and
S is the total number of taxa. If the two samples being wmpared share no taxa in common, SIMI has
a minimum value of 0; whereas, if the taxa present and their relative abundance are identical in both
samples, SIMI has a maximum value of I.
RESULTS

A total of 37 diatom taxa were identified from the three samples counted under
the LM. The identity and relative abundance of each diatom taxon is listed for
each seagrass species in Table I. Additional LM scans following the cessation of

461

SULLIV AN: EPIPHYTIC SEAGRASS DIATOMS

Table I. Relative abundance of diatom taxa epiphytic on seagrasses in Mississippi Sound expressed
as number of individuals in a sample (HB = Halodule beaudettei. CF = Cymodocea filiforme,
rr = Thalassia testudinum. and ~ nl = three samples pooled as one)
Diatom Taxon

Achnanthes hauckiana Grun.

Amphora coffei/ormis (Ag.) Kiitz.
A. cymbelloides Grun.
A. cymbiformis CI.
A. exigua Greg.
A. ol'alis var. pediculus (Kiitz.) V.H.
A. proteus Greg.
A. tenuissima Hust.
Berkeleya rlllilans (Trent.) Grun.
Cocconeis deperdita Giffen
C. placentu/a var. euglypta (Ehr.) CI.
C. sculellum Ehr.
C. woodii Reyes-Vasquez
Cye/olella caspia Grun.
Dimeregramma minor (Greg.) Ralfs
Fragi/aria hyalina (Klitz.) Grun.
Grammatophora oceanica (Ehr.) Grun.
Licmophora abbrel'iala Ag.
L. cf. debilis (Klitz.) Grun.
Mastogloia exigua Lewis
M. pusil/a Grun.
Nal'icula abunda Hust.
N. aequorea Hust.
N. gregaria Donkin
N. IUI/lseni M~lIer
N. pmil/ardi Hust.
N. pseudon}' Hust.
Nitzschia constricla (Greg.) Grun.
N. dissipala (Kiitz). Grun.
N. minlllll/a Grun.
N. pa/cacea Grun.
OpepllOra pacifica (Grun.) Petit
Rhopafodia gibberufa (Ehr.) Mull.
Strialella IInipllnclala (Lyngb.) Ag.
Synedra affinis val'. intermedia Grun.
S. fasl'iculala (Ag.) Kiitz.
Trachysphenia acuminata Perag.
Total No. Individuals

HB

CF

2
1
16
37
3
47
26
5
5
349
40
23
57
28
180
1
2
20
8
3
t
I
II

In,

IT

6
2
7

6
2
12
2
77
4
159
7
10
58
41
16
24
2
720
156
67
217
133
489

25
3
46
6
9
15
10
6
171
16
37
J03
90
209

36
I

76
I

7
2
I

13
2
200
100
7
57
15
100

5
20
14
1

I
I

5
9
I
I

44
71

1
5
37
67

42

50

4
36

1
48
3

17
68
132
42
4
81
4

1,003

1,000

1,064

4
6
45
31
4
2
3
33
149
270
134
9
165
7
1
3,067

counting and extensive SEM observations were carried out for purely taxonomic
purposes, and an additional eight taxa were encountered. These taxa were Amphora robusta Greg., Diploneis obliqua (Brun) Hust., D. pseudovalis Hust.,
Navicula amphipleuroides Hust., N. flanatica Orun., N. subforcipata Hust.,
Synedra investiens W. Sm., and Trachyneis aspera (Ehr.) CI.
The four most abundant diatoms, if the three seagrass samples are pooled to
yield a single sample (N = 3067), were Fragilaria hyalina, Mastogloia pusilla,
Licmophora cf. debilis, and Opephora pacifica (Table 1). For the most part,
these taxa were among the four most abundant ones on each individual seagrass
species, and collectively accounted for 55.3% of all individuals counted. If all
rare taxa are ignored (i.e. those represented by less than 10 individuals in each
of the three seagrass samples counted in Table 1), the taxonomic composition of
the three seagrass samples was with the exception of one taxon identical. This

BULLETIN

462

OF MARINE SCIENCE, VOL. 29, NO.4,

Table 2. Species diversity (H' in bits/individual),

terizing epiphytic diatom samples from Mississippi
= mean)

Statistic

H'

S
R'

1979

number of taxa (S), and redundancy

(R') characSound (Seagrass symbols same as in Table I and

HB

CF

IT

3.473

3.732
29

3.7]8
32

29
.304

.248

.276

3.641

30
.276

one exception involved Cocconeis scutellum, which was not present in the ThaLassia sample of 1,000 individuals. The significance of this "taxonomic sameness"
becomes apparent when it is seen that there were 18 rare taxa and the reamining
19 taxa accounted for 97.3% of all individuals counted in the three seagrass
samples (Table 1).
Table 2 presents values of three community diversity statistics characterizing
each seagrass sample. Each statistic was calculated from the data listed in Table
I. Regardless of which statistic is examined the conclusion is the same; namely,
that for all practical purposes the three samples exhibited virtually equal community diversity values. In addition, species diversity (H') is relatively high and
dominance in the samples (R') relatively low.
Comparisons of the structural similarity of selected epiphytic diatom samples
by a selected similarity index (SIMI) appears in Table 3. Values of SIMI for all
possible sample pairs ranged from 0.825 to 0.879, or put in another way, any two
samples shared between 82 and 88% of the maximum similarity possible.
DISCUSSION

An examination of community structure characterizing epiphytic diatom samples taken from three seagrass species in Mississippi Sound on 20 July 1977
revealed the following information: (1) Except for the rare taxa (2.7% of all individuals counted) and the absence of Cocconeis scutellum from one sample, the
taxonomic composition of the three samples was identical. (2) Values for three
different community diversity statistics were virtually identical for the three samples indicating an inherent sameness in species-numbers relationships. (3) Comparison of samples by SIMI showed that the three samples were structurally very
similar with respect not only to the kinds of taxa present but also the apportionment of individuals amongst constituent taxa. Therefore, the available taxonomic
and mathematical information indicates that Halodule beaudettei, Cymodocea
filiforme, and Thalassia testudinum supported a single, nearly homogeneous epiphytic diatom community on the above date in the seagrass beds sampled, despite
considerable differences in leaf morphology. Furthermore, it would appear that
the species pool for wlonization of the seagrass leaves by epiphytic diatoms is
the same for the above three angiosperms and that the absence of the rare species
from the sample counts is due to stochastic, rather than biological, processes.
Finally, even if the seagrass species are interacting nutritionally with their epiphytes, this relationship was not significantly different between the seagrasses on
20 July as no host specificity was apparent.
The questions naturally arise as to whether or not the epiphytic diatom community is largely or entirely homogeneous throughout the growing season of the
three seagrass species sampled, and what are the horizontal and vertical extents

SULLIVAN:

463

EPIPHYTIC SEAGRASS DIATOMS

Table 3. Similarity values (SIMI) for comparisons

Sound (Seagrass symbols same as in Table I)

of epiphytic

diatom

samples

Comparison

SIMI

HB vs. CF

.879
.873
.825

HB vs. IT
CF vs. TT

from

Mississippi

of this possible homogeneity in community structure. At present it is not possible

to answer these questions since only three samples of epiphytic diatoms were
collected on one date from three adjacent seagrass beds. However, studies of the
epiphytic flora of both marine angiosperms and macroalgae by other workers
suggest that both a temporal and spatial homogeneity may characterize the epiphytic diatom flora of seagrasses in Mississippi Sound if a long-term, more comprehensive study is conducted in the future. BaJlantine (1972) sampled the nondiatom algal flora epiphytic on the same three seagrass species (if synonymy is
taken into account) considered here, in addition to Halophila engelmannii Ascherson, off the west coast of Florida. Sixty-five taxa were recorded and the 20 to
25 most common epiphytic taxa were collected from all seagrass species. He
concluded that the algae simply utilized the seagrasses present in a particular
locality for attachment purposes since none showed any preference for a particular host. Sullivan (1977) examined the leaves and internodes of Ruppia maritima
L. growing in New Jersey salt marsh pools and showed that a single, nearly
homogeneous epiphytic diatom community was distributed over the entire angiosperm host. The epiphytic diatom flora of two Sargassum species collected from
seven widespread stations in the western Sargasso Sea was studied by Carpenter
(1970). Only 13 taxa could be found in the samples and except for a single station,
the dominant diatom was Mastogloia binotata (Grun.) Cl. Carpenter stated from
this and other distributional data that the epiphytic diatom flora of this large body
of water appeared to be of a relatively homogeneous nature. Main and McIntire
(1974) concluded from their work with Zostera marina and various macroalgal
species in Yaquina Bay Estuary, Oregon that macrophytes simply increase the
surface area available to diatoms for attachment, since associations between epiphytic diatoms and host macrophytes were a result of similar responses to the
physical environment rather than to nutritional interactions.
The results of the preliminary study reported here represent the first published
account of the epiphytic diatom flora on seagrasses in the Gulf of Mexico and
therefore is important for distributional records. Although temporal and spatial
extrapolations are not justified from the data, the essential findings are in agreement with similar, more comprehensive, studies in the listerature. The need for
further research in MIssissippi Sound and adjacent water masses in the Gulf of
Mexico is apparent as far as the epiphytic diatoms are concerned.
ACKNOWLEDGMENTS
The assistance
of Dr. L. N. Eleuterius and personnel of the Botany Section, Gulf Coast Research
Laboratory,
Ocean Springs, MS in collecting
the epiphytic
diatoms from the R/V HALOPHILA
is
gratefully acknowledged.
The work upon which Ihis report is based was supported
in part by funds
provided by the United States Department
of the Interior as authorized
under the Water Resources
Research Act of 1964. as amended,
and administered
by the Water Resources
Research Institute of
Mississippi.

464

BULLETIN

OF MARINE SCIENCE,

LITERATURE

VOL. 29, NO.4,

1979

CITED

Ballantine. D. 1972. Epiphytes of four Florida seagrass species in the Anclote Anchorage, Tarpon
Springs, Florida. M.A. Thesis, University of South Florida, Tampa. 87 pp.
Carpenter, E. J. 1970. Diatoms attached to floating Sargassum in the western Sargasso Sea. Phycologia 9: 269-274.
Humm, H. J. 1964. Epiphytes of the sea grass, ThaJassia testudinum, in Florida. Bull. Mar. Sci.
Gulf Carib. 14: 306-341.
Main, S. P., and C. D. McIntire. 1974. The distribution of epiphytic diatoms in Yaquina Estuary.
Oregon (U.S.A.). Bot. Mar. 17: 88-99.
Penhale, P. A. 1977. Macrophyte-epiphyte biomass and productivity in an eelgrass (Zostera marina
L.) community. J. Exp. Mar. BioI. Ecol. 26: 211-224.
Reyes-Vasquez, G. 1970. Studies on the diatom flora living on Tilalassia testudinum Konig in Biscayne Bay. Florida. Bull. Mar. Sci. 20: 105-134.
Sand-Jensen, K. 1977. Effect of epiphytes on eelgrass photosynthesis. Aquat. Bot. 3: 55-63.
Shannon, C. E., and W. Weaver. 1949. The mathematical theory of communication. Univ. Illinois
Press, Urbana. 117 pp.
Stander, J. M. 1970. Diversity and similarity of benthic fauna off Oregon. M.S. Thesis. Orcgon State
University, Corvallis. 72 pp.
Sullivan, M. J. 1977. Structural characteristics

of a diatom community epiphytic on Ruppia maritima.

Hydrobiologia 53: 81-86.

Wood, E. J. F. 1959. Some aspects of the ecology of Lake Macquarie. N.S.W .. with regard to an
alleged depletion of fish. VI. Plant communities and their significance. Aust. J. Mar. Freshwater
Res. 10: 322-340.
DATE ACCEPTED: May 22, 1979.
ADDRESS: Biology Department, P.O. Drawer Gr, Mississippi State, MS 39762.