GULF RESEARCH
REPORTS

Published by the

GULF COAST RESEARCH LABORATORY
Ocean Springs, Mississippi

November 1973

Gulf Research Reports

Volume 4 Issue 2

January 1973

A Quarter Century of Geology at the Gulf Coast Research Laboratory (1948-1973)

Ervin G. Otvos Jr.

Gulf Coast Research Laboratory

DOI: 10.18785/grr.0402.01

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Recommended Citation

OtvoS; E. G. Jr. 1973. A Quarter Century of Geology at the Gulf Coast Research Laboratory (1948-1973). Gulf Research Reports 4
(2); 151-165.

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A QUARTER CENTURY OF GEOLOGY AT THE GULF COAST
RESEARCH LABORATORY (1948-1 973)

by

ERVIN G. OTVOS, JR.

Gulf Coast Research Laboratory
Ocean Springs, Mississippi 39564

After its founding at the Mississippi Academy of Sciences meet-
ing on May 3, 1947 in the Buena Vista Hotel, Biloxi, Mississippi, the
Laboratory offered two biological courses during its first summer ses-
sion but no geology. Geology has been an integral part of Gulf Coast
Research Laboratory since its second year of existence. During the
first few years the abandoned Civilian Conservation Corps camp in
Magnolia State Park, east of Ocean Springs, was used for housing the
Laboratory, The activity was directed by Dr. R. L. Caylor, Chairman
of the Science Division, Delta State Teachers College in Cleveland,
Mississippi. An assortment of temporary buildings served as resi-
dence, office, library and dining hall. From surplus army equipment
additional dormitory buildings were constructed and vessels of the
Mississippi Sea Food Commission were borrowed at Ocean Springs
Harbor for field work. The Mi.ssissippi Board of Trustees of Institu-
tions of Higher Learning was induced to assume responsibility for the
Laboratory and the State Legislature established it as a state institu-
tion in 1948 and appropriated ten thousand dollars for its operation
in 1948-50.

Doctor Caylor .served for several years as director of the Labora-
tory. He was only on the grounds permanently during the summer.
He invited Dr. Richard R. Priddy, Chairman of Geology at Millsaps
College in Jackson (1948-1972) to help develop a geology teaching
program and join the Executive Board of the Laboratory, Priddy’s
participation had a profoundly beneficial effect on the geological work
for years to come. With youthful enthusiasm and interest in his sub-
ject he gradually overcame the problems created by the lack of equip-
ment, primitive, inconvenient facilities and his initial unfamiliarity
with coastal geology. Makeshift instruments (sediment corers, oxy-
gen and pH-meters, etc.) had to be manufactured. In the coming de-
cades practically all Millsaps geology majors and several chemistry
majors attended geology courses offered during the summers. These
courses, in fact, were made a recommended summer field camp sub-
stitute for the Millsaps geology majors.

While in 1947 only botany and marine biology were taught, in
August of 1948 geology became the third established course and a
two-week marine sedimentation problems class was taught by Priddy
with the assistance of C. P. Marion, a graduate student at Mississippi

151

152

Otvos

State College. Between 1949 and 1955 the course was called Marine
Sedimentation. After 1948 all geology courses lasted three weeks and
carried three hours credit. The four students in 1948 (C. A. Barton,
E. R. Campbell, F. G. Clark and W. E. Cook) were all from Millsaps
and students from this college were always very well represented in
the geology classes of the following summers (Table 1).

In the early years sediment sampling from the Biloxi Bay bot-
toms and the surrounding beaches occupied the course, along with the
description and grain size analysis of the samples. Later the class
projects became more varied and carried students to many different
locations along the Mississippi shore. Marion taught the geology
course during the summers of 1949-50. In 1952 Arthur T. Allen of
Emory University was the instructor and during the following three
summers Dr. Olin T. Brown of Mississippi Southern College (later
University of Southern Mississippi) taught the course. Priddy taught
in 1951 and again from 1955-58. In those formative years all profes-
sors practically donated their efforts and time. In the beginning the
teaching and research work at the Laboratory was largely restricted
to the summer months. As late as 1954 the year-round staff consisted
of three part-time clerical and maintenance personnel and only two
full-time .scientific staff members, including Dr. A. E. Hopkins, the
director who had become the first full-time director in 1952, It was
not until 1961 that the first full-time geologist assumed his duties.

The physical expansion of the Laboratory and its geological facil-
ities proceeded slowly but steadily. With the help of the Executive
Secretary of the Board of Trustees of Institutions of Higher Learn-
ing, the Laboratory acquired the Smart property on Davis Bayou, a
mile and a half south of the facilities in Magnolia Park. The purchase
price was $35,000. A 2V2-story main building (Fig. 1) that be-
came known as the “Big House,” was a former summer home that had
been constructed in 1900. The surrounding 49 acres of land was a
beautiful .setting on a small peninsula bounded by Halstead, Stark
and Davis bayous. In 1950, the “Big House” was occupied by Labora-
tory personnel and a fii'e in the Magnolia Park facilities during Au-
gust of that year hastened the completion of the move to the Smart
property in 1951.

One of the auxiliary structures behind the site of the future Hop-
kins Building, by the boat harbor, became the “mud shack”, a one-
room geological classroom-laboratory building. This had been a form-
er servant’s quarters. Between 1961 and 1965 the small kitchen and
pantry of the “Big House” were used for the geology office and stor-
age room. With the exception of two wooden dormitories, Laboratory
buildings were clustered on low ground along Davis Bayou. The dor-
mitories were reassembled on the slopes of the “Hill”, the densely
wooded, flat-topped “hinterland” which formed the center of the small

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Figure 1. The “Big House” shrouded by live oaks, from the pier. The
building was demolished by the 1969 hurricane and a parking lot covers
its site. (Photo by J. O. Snowden)

peninsula at 15-18 feet elevation and provided the potential for the
further expansion of the Laboratory.

In addition to sediment .studies in the Biloxi Ray area the study
of water and sediment chemistry of the bays and bayous became an
important subject during the .summer sessions. Priddy in 1952 in-
volved Dr. Franklin W. James of the Millsaps Chemistry Department
in the study of Fe, Si, Ca, Mg, nitrate, nitrite, sulfate, sulfide, organic
carbon and other components of the coastal water.s and sediments.
Priddy likewise involved Dr. Jo.seph B, Price, Chairman of the Mill-
saps Chemistry Department, in this study from 1954 until his un-
timely death in 1963, Salinity, redox-potential, pH, chlorinity and
other characteristics were .studied in some detail. A few Millsap.s
chemistry majors also took part in this study, including J. D, Powell
and Hugh Burford. Numerous abstracted progress reports and papers
originated from these investigations. Two of the papers, by Priddy
and Price, dealt with colorimetric and rapid volumetric analysis
methods of brackish waters and an issue of the Mississippi Geological
Survey Bulletin contained the summary of the analytical results of
the Mississippi Sound work. As a result of this emphasi.s on water
and sediment chemi.stry, two geology courses were offered, starting in
1956. “Marine Geology” (later renamed “Physical Marine Geology”)
dealt with physical geological and sedimentological aspects of coastal

A Quarter Century ok Geology

155

Figure 2. Dr, Joseph B. Price. Chemistry Chairman and A. D. Bishop. 1960
summer student (both of Millsaps) in the old Hopkins Building. (Photo
by J. O. Snowden)

Figure 3. Professor R. R. Priddy in the Hopkins Building (before Hurri-
cane Camille). (Photo by Dr. J. O. Snowden)

156

Otvos

geology, while “Marine Sedimentation” (later renamed “Problems in
Sedimentation”, and still later as “Chemical Marine Geology”) con-
cerned itself with the analysis of chemical components and of the
physico-chemical properties of bottom sediments and of the coastal
water bodies. Until Dr. J, O. Snowden, Jr., one of Priddy*s former
students and a member of his Millsaps faculty at the time, took over
the teaching of the chemical marine geology course in 1968, Priddy
taught it every summer with the exception of 1959. The Laboratory
hosted regional geology courses for college instructors during the
consecutive summers of 1965-68. The.se National Science Foundation-
sponsored courses were directed by Priddy in the Caylor Building.

Physical facilities were .slowly improving. Since its completion in
June 1951, the geology classes were able to utilize parts of the small
(30 by 60 feet) Caylor Building, the first brick structure on the cam-
pus. After the arrival of Dr. Gordon Gunter in 1955 to be director of
the Laboratory, a second one-story classroom-laboratory building,
named after A. E. Hopkin.s, was opened in 1956 and all of the Caylor
Building along with parts of the Hopkins Building were used for
geological purposes. Both these structures and the “Big House” were
demolished in the 1969 hurricane and only the Hopkins Building was
rebuilt thereafter. By this time, however, geology work had for sev-
eral years been occupying modern, well-equipped, air-conditioned

Figure 4. The old, un-air-conditioned Hopkins Building before Hurricane
Camille. (Photo by J. O. Snowden)

A Quarter Century of Geology

157

quarters in the Oceanoj^raphy Building on the top of the “Hill” which,
when completed in May 1965, became the center of the Laboratory
complex.

In 1959 both geology courses were taught by Dr. A. R. Cariani of
University of Mississippi. The following year when the Laboratory
still had only six resident scienti.sts, including Doctor Gunter, Dr.
David A. DeVries of Mis.sissippi Southern College taught Physical
Marine Geology, He became resident geologist in 1961 and taught the
cour.se as a member of the .staff through the summer of 1966. Between
the summer of 1967 and January 1971, this position was filled by
W. L. Siler.

Student class reports written under the supervision of DeVries
and Priddy indicate a variety of .study projects during the summers.
Modern beach sands were investigated on Horn, Ship and Round Is-
lands and on Belle Fontaine Beach; ancient beach sand.s were studied
at Belle Fontaine, changes of the Marsh Point sand spit were measured
and recorded in great detail and minor and major beach forms in-
vestigated in several areas. One of the most original projects was jet-
drilling with a centrifugal pump from a boat in Davis Bayou in estab-

Figure 5. 1960 Chemical Marine Geology class at the Laboratory. Front
row, left-to-right : Fred Lockett, Marlin Hobbs and Dr. R. R. Priddy. Back
row: Frank Brooks, Joe Snowden, A1 Bishop, Dr. J. B. Price and Billy
Moore,

158

Otvos

lishing the configuration of an incised T.ate Pleistocene stream valley-
in the continuation of the prevsent Stark Bayou. This gully was filled
in and buried by Holocene deposits. Some of the heavy mineral, mi-
crofauna and other studies which were initiated during the summer
sessions developed into masters theses and several abstracts and pa-
pers were published later. Similar abstracts resulted from the chemi-
cal studies of Priddy and his associates on the sulfur cycle, flocculent
materials, Eh-pH conditions, humus content and other aspects of the
bottom muds and the waters of Mississippi Sound and adjacent bays
and bayous.

Doctor Snowden of Louisiana State University in New Orleans
and the writer (who became affiliated with the Laboratory on a part-
time basis on July 1, 1970 and became Geology Section Head June 1,
1971) have undertaken an extensive study of the chemistry and sedi-
mentary petrology of the deposits, the chemistry and the pore-water
and overlying free-water bodies along the entire Mississippi coast
with a grant from the Water Resources Institute under the U, S. Of-
fice of Water Resources Research, The Pearl, Wolf and Pascagoula
river estuaries and Davis, Heron, Old Fort and Graveline bayous were
investigated in this re.spect during 1970-72. Instead of the time-
consuming classical titrimetric methods, atomic absorption spectro-
metry was employed and the salinity of the squeezed-out pore water
measured by the micro-resistivity method. The sulfate, nitrite and ni-
trate content of the water samples was determined by the fast tur-
bidimetric analysis. These new methods were introduced in Dr. J. 0.
Snowden’s coui'se in which water chemistry and the chemical inter-
action between interstitial pore water and the enclosing sediments has
been emphasized.

As the result of Doctor Gunter’s vigorous program of develop-
ment and expansion of the Laboratory, several large modern build-
ings were added in 1965-71. Recovery was fast from the severe dev-
astation of Hurricane Camille in August 1969 with generous federal,
state and private aid. The Instructional Facility and Oceanography
Buildings became the largest on the campus, both with approximately
18,000 square feet gross area. A phy.sical marine geology and a
chemical marine geology cla.ssroom with adjoining office and storage
rooms opened in the two-story Instructional Facility Building in 1971
(renamed Richard L. Caylor Building in 1973) and additional space
was utilized for geological research and sample storage in the new
Research Facility Building the same year. A new exhibition center
near Point Cadet, Biloxi, prompted planning of part of the future ex-
hibit to demonstrate the geological makeup and history of the coast
and the role environmental geology plays in the area.

Geology has also acquired additional personnel. In 1971 Michael
Bograd and Edmond Funel worked as temporary assistants in the
sedimentation laboratory and the field. The following year two per-

A Quarter Century of Geology

159

Figure 6. Chemical Marine fieologj’^ class (AugusU 1062) at Horn Island.
W. E. Davenport, Ethel Rad/ewicz (Mlllsaps) Marshall Kern (Mississippi
Cieol. Survey), L. D. Simpson (Mississippi State) and W. H. Allen (Uni-
versity of Mississippi). Dr. R. R. Priddy at right. (Photo by Marion
Saucier)

manent assistants replaced them: Wade Howat, assistant geologist
(B.S. 1971), and Joseph Milner. In 1972 Robin Wink replaced Milner
as geo-technician and Mrs. Kathy Peter (B.S. 1972) also stepped in
as temporary sedimentation laboratory assistant. These developments
paved the way for an expanded geology program. In 1971 geology
participated in an interdisciplinary project of earth resources data
collection from southea.stern, coastal and otfshore Mi.s.si.ssippi, rele-
vant to the cultural, physical and economic development of the State
of Mi.ssissippi, This project, which resulted in a geological bibliog-
raphy and survey of the map coverage of the subject area, was done
in conjunction with geology faculty members at Mi.ssissippi State
University and the University of Southern Mississippi for the Earth
Resources Laboratory, Mi.s,sissippi Test Facility (Bay St. Louis) of
the National Aeronautic.s and Space Administration. Since 1971 the
Geology Section has also participated in another complex research
project, involving the three state universities in Missi.ssippi under the
Office of Sea Grant Programs, in studying sedimentation and sedi-
ment chemistry in a St. Louis Bay oyster reef and in the Bay area in
general. Another study, in cooperation with the Fisheries Section,
deals with the sedimentology of Biloxi Bay and is done for the Earth

160

Otvos

Resources Laboratory, NASA. The writer was appointed by Doctor
Gunter in September 1971 to represent the Laboratory on the Uni-
versities Marine Center Marine Education Committee for Mississippi.

Since 1972 the Section has actively participated in the Environ-
mental Affairs Committee of the Laboratory. Work of this committee
has included formulating recommendations for the Mississippi Ma-
rine Resources Council about proposed construction projects in the
coastal zone which could have an impact on the coastal environment
in Mississippi. The head of the Section also participates in the work-
ings of the Point Cadet Science Advisory Committee which assists in
the development of the Laboratory’s new exhibition-research facility,
the Marine Education Center, in the southeastern corner of Biloxi. As
of 1973 the author has also been charged by Director Harold D.
Howse to represent the Laboratory in the recently established Missis-
sippi Mineral Resources Institute.

Systematic acquisition of topographical, hydrological, geological
and aerial photo maps has been started for a geological map collection
for research and teaching purpo.ses- The collection primarily consists
of maps related to Gulf and Atlantic coastal areas but also includes
archive copies of Mississippi-Louisiana area charts. Holdings of the
library in geological and related fields have been steadily increasing
and the library currently is subscribing to about twenty domestic
and foreign periodicals, dealing with different aspects of earth sci-
ences. During the past decade, as demands have periodically arisen,
staff geologists have taught physical geology evening classes at the
University of Southern Mississippi Keesler Air Force Base Resident
Center in Biloxi and on Gulf Park Campus, Long Beach. Recently,
historical and environmental geology courses were taught by the
writer.

Starting in 1971 one of the main research objectives of the Geol-
ogy Section has been the detailed study of the composition and geolog-
ical history of the Pliocene Citronelle Formation, the Pleistocene
fluvial-alluvial, brackish and nearshore-marine deposits and of the
Holocene-Recent sediments, including the relationship of these units
to each other and the surface configurations of the Miocene “base-
ment’^ unit. The problem of the older (pre-Sangamon), higher Pleis-
tocene marine deposits, barrier beach ridges was investigated not only
in the Mississippi Coast but also in adjoining states and the coastal
Florida Panhandle. Correlation problems of the Citronelle under the
present Pleistocene coastal plains and its distinction from underlying
Miocene and overlying Plei.stocene formations are among the several
problems, as is the nature and origin of “terraces” and .scarps in the
coastal zone. The T>ate Pleistocene (Sangamon) Waveland-Gulfport-
Belle Fontaine mainland barrier ridge system is now being investi-
gated in great detail, Other held studies revealed that there is no geo-
logical proof for the existence of earlier Pleistocene, higher coastal

A Quarter Century of Geology

161

Figure 7. Physical Marine Geology class in the Sedimentation Laboratory,
Oceanography Building, 1971. Mary Whitten (left) and Norma Weaver,
University of Mississippi students with author. (Photo by Catherine
Campbell)

ridges, previously widely reported from several areas of the north-
eastern Gulf coastal region.

An extensive core-drilling program, so e.ssential in coastal re-
gions with few natural outcrops, was initiated in October 1971 to
solve the numerous stratigraphic-geomorphological problems. The
most important result of this project to date has been the recognition
of a widespread, continuous, shallow stratigraphic unit under the
Sangamon-agc barrier ridge zone and under certain areas of the Late
Pleistocene coastwise alluvial plain. This fossiliferou.s unit (named
Biloxi Formation) appears to represent the marine transgression
phase of the Sangamon Interglacial and is correlative probably along
the whole northern coast of the Gulf of Mexico. Plans also call for
stratigraphic exploration drilling in Mississippi Sound and on the off-
shore barrier islands in order to learn more about the pre-Pleistocene,
Pleistocene and Holocene history of the north-central Gulf coastal-
nearshore area and its relationship with the .subsiding zone of the
Mississippi River Delta region.

Geological field work including future detailed boring in Missis-
sippi offshore barrier islands and the south Hancock County marsh-
beach ridge complex should clarify the Holocene history of the present

162

Otvos

offshore-inshore area and the changes (crosional destruction, island
migration, accretion) which profoundly affect them. Long-range ef-
fects of natural (hurricanes) and man-made changes on these fea-
tures and the resulting effects on the mainland coast and the Missis-
sippi Sound are not yet adequately knowui. Such geological studies
would contribute not only to the understanding of the formation and
destruction of barrier islands and mainland barrier beaches during
the Late Quaternary in general, but also to the clarification and possi-
ble solution of serious environmental problems along the Mississippi
Coast.

Results of geological research of the past few years were demon-
strated in May 1973 during a two-day field trip of the New Orleans
Geological Society. The trip covered a wide area between New Or-
leans East and Dauphin Island, Alabama. A guidebook published for
the occasion summarized the present knowledge about the Miocene-
Recent coastal formations and the presently active geological pro-
cesses, At the request of the Commission on Shorelines of the Interna-
tional Association for Quaternary Research, we are also collaborating
in supplying regional information for the preparation of a set of
world shoreline maps, covering the Late Pleistocene, Late Holocene
and Recent shore positions.

ACKNOWLEDGEMENTS

Dr. Gordon Gunter, Director Emeritus and Professor of Zoology
at the Laboratory, encouraged the writing of this account and re-
viewed the manuscript. Doctor Gunter and Doctor Snowden supplied
data used by the author as did Doctor Priddy, whose mimeographed
descriptions of the geological activities at Gulf Coast Research Lab-
oratory were invaluable in the preparation of this article. Michael
Bograd greatly assisted in the tabulation of the student attendance
data.

BIBLIOGRAPHY

(Geological publications and the.ses associated with the Laboratory)

(1) Papers, books and reports

Clark, S, F. 1960, Behavior of some Gulf Coa>;t muds as compared to some high-
way clays‘ Jour. Miss. Acad. Sci., Vol. 6 (1954—60), pp. 197-201.

Foxworth, R. D. and B. Z. Ellis. 1960. Preliminary study of heavy minerals of
Mississippi Sound; Jour. Miss. Acad. Sci., Vol. 6 (1954-00), pp. 217-220.

Fo.xworth, K. D., R. R. Priddy, W. B. Johnson and W. S. Moore. 1962. Heavy min-
eral.s of sand from recent beaches of the Gulf Coast of Mi.ssissippi and asso-
ciated islands: Miss. State Geol. Survey Bull., No. 93.

A Quarter Century of Geology

163

Franks, J. S., J. Y. Christmas, W. h. Silur, R, Combs, R. Waller and C. Burns.
1972. A study of nektonic and benthic faunas of the shallow Gulf of Mexico
off the State of Mississippi as related to some physical, qhcmical and geologi-
cal factors 1 Gulf Research Reports, Vol. 4, No. 1.

Holladay, C. O., J. H, Foster, J, H. Hetrick, Jr. and R. D. Foxworth. 1957. A study
of the shifting of Marsh Point Sand Spit: Jour. Miss. Acad. Sci., Vol. 6
(1954-60), pp. 203-206.

Otvos, E. G., Jr. 1970. Development and migration of barrier islands, northern
Gulf of Mexico: Reply; Geol. Soc. Amcr. Bull., Vol. 81, No. 12, pp. 3783 88.

1971, Relict eolian dunes and the age of the “Prairie” coastwise terrace,

southeastern Louisiana: Geol. Soc. Amer. Bull., Vol. 82. No. 6, pp. 1753-58.

1971. (with J. Pinson, B. W. Brown, C, M. Hoskin, T. J. Laswell, et

al.) , Geology section, p, 20, in Earth resources data and technological studies
relevant to the cultural, physical and economic development of the State of
Mississippi, Task 1 Contract NAS9-11607, Final Report, (also individual
geology report).

_. 1972. Pre-Sangainon beach ridges along the northeastern Gulf Coast-
fact or fiction?: Gulf Coast Assoc. Geol. Soc. Trans., Vol. 22, pp. 223-228.

1972. Mississippi Gulf Coast Pleistocene beach barriers and the age

problem of the Atlancic-Gulf Coast “Pamlico” — “Ingleside” beach ridge sys-
tem : Southeastern Geology, Vol. 14, No. 4, pp. 241-250.

1973. Genetic and age problems of the Moreau-Caminada Holocene

coastal ridge complex: Southeastern Geology, Vol. 15, No. 1, pp. 37-43.

1973. Geology of the Mississippi-Alabama coastal area and near-shore

zone: Field Trip Guidebook: New Orleans Geol, Society, New Orleans.

1973. Inverse beach sand texture^ — coastal energy relationship on Missis-
sippi-Alabama Coast barrier islands: Jour. Miss. Acad. Science, Vol. 18 (in
press) .

1973. Phase 111; Scdimcntology, in Cooperative Gulf of Mexico Estua-
rine Inventory and Study, Mississippi; State of Mississippi Marine Conserva-
tion Commission and Gulf Coast Research Laboratory. Ed.i J. Y. Christmas,
pp. 123-137.

Price, J. B. and G. Gunter. 1964. Studies of the chemistry of fresh and low'-salinity
waters in Mississippi and the boundary between fresh and brackish watgr:
Int. Revue ges, Hydrobiol., Vol. 49, No. 4, pp. 629-636.

Price, J. B. and R. R. Priddy. 1959. Colorimetric determinations of nitrite and
nitrate nitrogen in brackish coastal waters: Bull. Mar. Sci. Gulf and Carib-
bean, Vol. 9, No. 3, pp. 310-314.

1961. Rapid volumetric determination of sulphate, calcium and magne-
sium in high lime-magnesian brackish coastal waters: Bull. Mar. Sci. Gulf
and Caribbean, Vol. 11, No. 2, pp. 198-206.

Priddy, R. R. 1954. Recent Mississippi Sound sediments compared with some Up-
per Cretaceous sediments: Gulf Coast Assoc. Geol. Soc, Trans., Vol. 4, pp.
159-167.

Priddy, R, R, and R. M, Crisler, Jr. 1955. Preliminary survey of sediments in parts
of the Mississippi Sound: Jour. Miss. Acad. Sci., Vol, 5 (1951-53), pp. 226-
230.

Priddy, R. R., R. M. Crisler, Jr., C, P, Sebren, J, D. Powell and H. Bui-ford, 1955.
Sediments of Mj.ssissippi Sound and inshore waters: Miss, State Geol. Surv.
Bull., xNo, 82.

Priddy, R. R. and B. L. Smith. 1960. Recent sedimentation on Horn Island, Missis-
sippi: Gulf Coast Assoc. Geol. Soc. Field Trip Guidebook, pp, 5-8.

Snowden, J. 0., Jr. 1965, Settling velocities of selected clay minerals in brackish
and normal sea water; Jour. Miss. Acad. ,Sci., Vol. 11, pp. 123-126.

Upshaw, C. F,, W. B. Creaih, and F. L. Brooks. 1966. Sediments and microfauna
off the coasts of Mississippi and adjacent states: Miss. State Geol. Survey
Bull., No. 106.

164

Otvos

(2) Published lecture abstracts

DeVries, D. A. 1969. Grass-balls formed on a Mississippi beach during Hurricane
Betsy: Geol. Soc. Amer. Abstracts with Programs, 1969, Pt. 6, North-central
Section, pp. 10-11.

DeVries, D. A. and R. R. Priddy. 1964. An eight-year study of Marsh Point Sand
SpiL Jour. Miss. Acad. Sci., Vol. 10, p. 182.

Moore, W. S. 1962. Eh-pH relations in sulfide deposition in Mississippi Sound:
Jour. Miss. Acad. Sci„ Vol. 8, p. 100,

Moore, W. S. and J. O. Snowden, Jr. 1967. Sulfates in Mississippi coastal waters:
Jour. Mhss, Acad. Sci., Vol. 13, pp. 59-60.

Otvos, E. G., Jr. 1972. Pre-Sangamon beach ridges along the northeastern Gulf
Coast-fact or fiction?: Bull. Amer. Assoc. Petr. Geologists, Vol. 66, No. 9,
p. 1901.

Price, J. B. and R. R. Priddy. 1960. Colorimetric determination of nitrite and
nitrate nitrogen in brackish coastal waters: Jour. Miss. Acad. Sci., Vol. 6
(1954-60), p. .349.

1960. Rapid volumetric determination of calcium and magnesium in

brackish coastal waters: Jour. Miss. Acad. Sci., Vol, 6 (1954-60), p. 351,

1960. Rapid volumetric determination of sulfate, calcium and magne-
sium in high lime-magnesium brackish coastal waters: Jour. Miss. Acad. Sci-,
Vol. 6 (1954-60), p. 399.

Price, J. B., R. R. Priddy, W. S. Moore and W. 1. Luke. 1962. Investigation of
flocculent materials in Mississippi Sound: Jour. Miss, Acad, Sci., Vol. 8,

p. 111.

Priddy, R, R. 1955. Humu.s of Mississippi Sound: Geol. Soc. Amer. Bull., Vol. 66,
No. 12, Pt. 2, p. 1717,

1960. Additional studies of Gulf Coast sediments: Jour. Miss. Acad.

Sci., Vol. 6 (19.54-60), p. 58.

1962. Mississippi Sound heavy minerals compared to other “heavies” of

the Gulf Coast and South Atlantic Coast: Jour. Miss. Acad. Sci., Vol. 8,
pp. 102-103.

Priddy, R. R,, R. M. Crisler and H. Burford. 1954, Sediments of parts of Missis-
sippi Sound: Geol. Soc. Amer. Bull., Vol. 65, No 12, Pt. 2, pp. 1366-67.

Priddy, R. R., R. J, Gentile and R H. Lyons. 1964. Observations on the sulfur
cycle in some Mississippi Sound muds: Jour. Miss, Acad. Sci., Vol. 10, pp. 49-
.50.

Priddy, R. R, and E. J. Johnson. 1966, Sulfur and sulfur bacteria in some Missis-
sippi Sound muds; Geol. Soc. Amer. Special Paper, No. 87, p. 260.

Snowden, J. 0., Jr. 1961. Recent sedimentation of Biloxi Bay, Mississippi: Progr.
Ann. Mtiig. Geol. Soc. America, Cincinnati.

Snowden, J. O., Jr. and E, Otvosi Jr. 1971, Chemical water quality and sedi-
ment-water relations in Louisiana and Mississippi estuaries: Abstl*. Second
Coastal and Shallow Water Research Conference (Univ. of Southern Cali-
fornia, University Press), p. 215.

1972. (In Press). Chemical changes in interstitial sediment water and

clay mineralogy in Louisiana-Mississippi estuaries: Clays and the Marine En-
vironment, Proc. of the 21st Conf. Clay Minerals Soc.

(3) Masters Theses

Barton, C. A. 1952. The sediments of Biloxi Bay, Mississippi: M. A. Thesis Uni-
versity of Illinois.

Foxworth, R. D. 1958. Heavy minerals of sand from recent beaches of the Gulf
Coast of Mississippi and associated islands: M. A. Thesis, University of Mis-
souri.

Lindsey, R, H., Jr. 1962. A study of recent forminifera along a salinity gradient

A Quarter Century of Geology

165

through the Mississippi Sound, Mississippi: M. S. Thesis, University of Mis-
sissippi.

Marion, C. P. 1951. A study of recent marine sediments in the Biloxi-Ocean
Springs area of the Mississippi Gulf Coast: M, A. Thesis, Mississippi State
College.

Robbins, W. H. 1961. Ecology of foraminifera of Biloxi Bay, Mississippi: M. S.
Thesis, University of Missouri.

Snowden, J. O., Jr. 1961. Geological and chemical environment of Biloxi Bay, Mis-
sissippi: M. A, Thesis, University of Missouri.

(4) Papers in Preparation

Otvos, E. G., Jr. Inverse beach sand texture — wave energy relationship, north-
east Gulf Coast barrier islands.

A Holocene beach ridge system (“Hancock County Chenier”), South-
western Mississippi Coast.

Sediments and sediment chemistry of Holocene St. Louis Bay deposits.

A late Pleistocene transgressive unit, northern Gulf Coast (Biloxi For-
mation) .

Holocene history of the Mississippi-Alabama coastal area.

Development and recent history of Mi.ssissippi-Alabama coastal barrier

islands.

The late Holocene Hancock-New Orleans coastal-submarine ridge sys-
tem.

Otvos, E. G., Jr. and J. O. Snowden, Jr. Geology of the Back Bay of Biloxi.

Snowden, J. O. and E. G. Otvos, Jr. Chemical water quality and sediment-water
reactions in Louisiana and Mississippi estuaries: Louisiana Water Resources
Research Institute Techn. Rprt. No. 5. Louisiana State Univ,, Baton Rouge,
La.

Gulf Research Reports

Volume 4 Issue 2

January 1973

The Occurrence of the Remarkable Scyphozoan, Deepstaria enigmatica, in the Gulf of
Mexico and Some Observations on Cnidarian Symbionts

Philip J. Phillips

Texas A6’M University

DOI: 10.18785/grr.0402.02

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Recommended Citation

Phillips, P. J. 1973. The Occurrence of the Remarkable Scyphozoan, Deepstaria enigmatica, in the Gulf ofMexico and Some
Observations on Cnidarian Symbionts. GulfResearch Reports 4 (2): 166-168.

Retrieved from http:// aquila.usm.edu/gcr/vol4/iss2/2

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THE OCCURRENCE OF THE REMARKABLE SCYPHOZOAN,
Deepstaria enigmatica, IN THE GULF OF MEXICO AND
SOME OBSERVATIONS ON CNIDARIAN SYMBIONTS

by

PHILIP J. PHILLIPS^
Department of Biology
Texas A&M University
College Station, Texas 77801

On 8 July 1965 one damaged specimen of the remarkable scypho-
zoan, Deepstaria enigmatica Russell 1967, was taken in the course of
a sampling program conducted by the Department of Oceanography,
Texas A&M University in a ten-foot Issacs-Kidd midwater trawl in
the Yucatan Basin. The trawl was put into water at Lat. 19° 58' N,
Long. 85° 14' W. This is the first report of Deepstaria in the Atlantic
region. This medusa is otherwise known only from the Pacific Ocean
where it has been captured by slurp gun from the submersible Deep-
star over the San Diego Trench and has been taken in midwater
trawls by the Scripps Institution of Oceanography Mid-Pacific Ex-
pedition (Barham and Pickwell 1969 and Russell 1967).

The present specimen, although badly torn, is clearly recogniz-
able as D. enigmatica. The anastomosing canal lattice has the charac-
teristic pattern described by Russell. The medusa is a deep purple-
blue in color and appears to have had a diameter in excess of 70 cm.
The mesogloea is 18 mm. thick near the center of the disc. Oral arms
are not discernible and portions of the disc are considerably distorted
by local extreme contraction. Fragments of female gonadal material
are attached to parts of the subumbrellar surface. These gonadal frag-
ments, too small to permit any observations on gross gonad morphol-
ogy! contain zygotes in various stages of early cleavage. Ova and
zygotes are small (c. 100 microns diameter) and have very little, if
any, yolk. The depth at which this jellyfish was taken cannot be stated
with any certainty since the collecting device fi.shed from the surface
to a depth of 2400 m., no '‘at depth" closing device having been used.

The occurrence of this unique scyphomedusan in the Gulf of Mex-
ico constitutes a considerable range extension. Closer examination of
midwater trawl samples from other regions and more extensive sam-
pling will probably lead to the discovery of more specimens of Deep-
staria from other areas, I strongly suspect that D. enigmatica may
have a circumglobal di.stribution in tropical and subtropical waters.
As is the case with many oceanic organisms, the zoogeographic dis-

1 Present address; North Atlantic Division, U.S. Army Corps oX Engineers,
90 Church Street, New York, N. Y. 10007.

166

Deepstaria enigmatica in the Gulf

167

tribution of Deepstaria can probably be correlated with intensity of
sampling and workers who can identify it.

Barham and Pickwell discuss a commensal or symbiotic associa-
tion between the giant parasitic isopod Anuropus and Deepstaria,
Photographs taken by Deepstwr show one of these isopods clinging to
the subumbrellar surface at the time of capture. Barham and Pick-
well state that theTnedusa ‘Mid not pulsate or show normal swimming
movements. Instead the medusa appeared flaccid and seemed to be
floating passively.” Ru.ssell noted the absence of stomach, epithelial
lining and parts of the coronal muscle, and suggested the possibility
the medusa was moribund at the time of capture. Taking into account
the occurrence of nematocysts in the stomachs of some anuropids (as
reported by Menzies and Dow 1958 and cited by Barham and Pick-
well) Barham and Pickwell suggested that the isopod feeds on the
jellyfish, incapacitates medusa movement and creates a “floating pro-
tective environment,” which they state is a process that stops “some-
what short of the reduction of salps and pyrosomes into thin-walled
houses by the well known amphipod, Phronma sedenteria'' It should
be noted that Barham and Pickwell did not demonstrate nematocysts
in the stomach contents of the Anuropvs in association with Deep-
staria, While the isopod may be traveling on an atenic host or is ac-
tually a true parasite of Deepstaria there are no criteria for establish-
ing just what can be considered normal swimming movement for this
medusa, if indeed it noi*mally does generate swimming movements, I
find it difficult to believe that Anuropus incapacitates the medusa
since my own personal and published observations (Phillips, Burke
and Keener 1969) indicate that medusae can serve as hosts for a wide
variety of crustaceans and other metazoans without incapacitation. In
actual fact the moribund state will render an organism prey for a
wide range of animals which would not normally attack it, as may
possibly be the case for Deepstaria,

Additionally, it is exceedingly dubious that Phronhna invades
salps or pyrosomes and converts them into thin-walled houses. The
structure of the amphipod house bears little semblance of that of any
tunicate and the resemblance is a superficial one at best according to
Leo Berner, Jr., (personal communication). It is much more likely
that a Phronima secretes its own domicile. Examination of Phronima
houses evinces no evidence of zooids of any type ever having been em-
bedded in the gelatinous matrix. There is also a very striking correla-
tion between house proportions and amphipod size, a situation that
would not be so well defined if the Phronima did not secrete it.

Symbiotic relationships between Scyphozoa and other metazoans
are very common. Pickwell and Barham cite seven such relationships
and their list can be considerably lengthened. Fish-jellyfish associa-
tions are probably the best known (Marisueti 1963 and Phillips, Burke
and Keener 1969) and many are known that involve Crustacea and

168

Phillips

medusae (Outsell 1928 and Phillips, Burke and Keener 1969). It
should be noted that there are trematode and cestode parasites of
medusae (Dollfus 1931 and Stunkard 1969) as well as vermiform
parasites of unknown phyletic aflfinilies which are commonly found in
some rhizostomes (Moestafa and McConnauj^hey 1966 and Phillips
and Levin, in prep.), I have also found on very rare occasions hyperiid
amphipods embedded in the mesogloea of two species of calycophoran
siphonophores (Diphyes dispar and Chelophyes appendiculata) . Me-
tazoan parasites or symbionts of Cnidaria are not at all rare; they
are merely seldom looked for and even less seldom investigated.

It remains to be seen whether or not life cycle completion in
Anuropm is dependent on cnidarian hosts. Data presented by Barham
and Pickwell regarding the association between Annropus and Deep-
stama are insufficient for drawing conclusions.

ACKNOWLEDGEMENTS

I would like to thank Dr. Leo Berner, Jr. (Department of Ocean-
ography, Texas A&M University) and Dr. Sewell H. Hopkins (De-
partment of Biology, Texas A&M University) for critical review of
the manuscript. This study was supported in part by NSF Grant GP
3555. The Gulf of Mexico Deepstaria enigmatica specimen was made
available to me through the courtesy of Dr. W. E. Pequenat and Dr.
L. Berner, Jr. and has been placed in the U. S. National Museum,
Washington, D. C., U.S.A. This paper in modified form was part of
a dissertation submitted in partial fulfdlment of the requirements for
the Ph.D. degree in biology at Texas A&M University.

LITERATURE CITED

Barham, E. G. and G. V. Pickwell. 1969. The giant isopod, Anuropus: a scypho-
zoan symbiont. Deep Sea Research. 16:525-9.

Dollfus, R. P. 1931. Nouvel addendum a mon “Enumeration des cestodes du planc-
ton et des invertebres marins” (1) Annales de Parasitologie Humainc et Com-
paree. 9(6) :552-60.

Gutsell, J. S. 1928. The spider crab, Libinia dubia, and the jellyfish, Stomolophiis
meleayris found associated at Beaufort, North Carolina. Ecology. 9(3) :358-9.
Mansueti, R. 1963. Symbiotic behavior between small fishes and jellyfishes with
new data on that between the stromateid, Pepriius alepidotus and the Scy-
phomedusa, Chrysaora quinquecin ha, Gopeia. (1) ;40-80.

Moestafa, S. H. and B. H. McConnaughey. 1966. Catostyliis ouwensi (Rhizostomae,
Catostylidae) , a new jellyfish from Irian (New Guinea) and Ouwensia cato-
styli n. gen., n. sp., parasitic in C. otiwcnsi, Treubia. 27(1) :l-9.

Phillips, P. J., W, D, Burke and E. J. Keener, 1969, Observations on the trophic
significance of jellyfishes in Mississippi Sound with quantitative data on the
associative behavior of small fi.shes with medusae. Transactions American
Fisheries Society. 98(4) :703-l2.

Russell, F. S. 1967. On a remarkable new seyphomedusan. Journal of the Marine
Biological Association of the United Kingdom, 47(3) ;469-73.

Stunkard, H. W. 1969. The morphology and life history of Neopechona pyriforme
(Linton 1900) n. gen., n. comb. (Trematoda: Lepocreadiidae) . Tke Biologi-
cal Bulletin. 136:96-113.

Gulf Research Reports

Volume 4 Issue 2

January 1973

Stranding Records of a Finback Whale^ Balaenoptera physalus, from Mississippi and the
Goose-Beaked Whale^ Ziphius cavirostris, from Louisiana

Gordon Gunter

Gulf Coast Research Laboratory

J.Y. Ghristmas

Gulf Coast Research Laboratory

DOI: 10.18785/grr.0402.03

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Part of the Marine Biology Commons

Recommended Citation

Gunter; G. andj. Christmas. 1973. Stranding Records of a Finback Whale^ Balaenoptera physalus, from Mississippi and the Goose-
Beaked Whale, Ziphius cavirostris, from Louisiana. Gulf Research Reports 4 (2); 169-173.

Retrieved from http;// aquila.usm.edu/gcr/vol4/iss2/3

This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean
Research by an authorized administrator of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

STRAJNDING RECORDS OF A FINBACK WHALE, Balaenopiera
physalm,, FROM MISSISSIPPI AND THE GOOSE-BEAKED
WHALE, Ziphius eavirostris, FROM LOUISIANA

by

GORDON GUNTER and J. Y. CHRISTMAS
Gulf Coast Research Laboratory
Ocean Springs, Mississippi

INTRODUCTION

Stranding records are sometimes the only source of data con-
cerning some species of cetaceans and it is appropriate that these in-
cidents be reported in some detail. This is particularly true of rare or
little known species. Bottle-nosed dolphin (Tursiops truncahis) car-
casses stranded on the Mississippi and Louisiana coasts are so com-
mon that they receive little attention. Other species occur and have, no
doubt, been overlooked or confused by laymen with I'nrsiops. Even
the rare stranding of large whales in this area may go unreported ex-
cept for news stories that fail to identify the animal properly and
which are not readily available to cetologists.

The stranding in 1967 of one of the first definitely reported fin-
backs, Dalaerioptera physahis, in the Gulf of Mexico is given here. It is
the first Mississippi record,

Moore (1953) noted that “only six specimens of the goose-beaked
whale have been reported on the eastern coast of North America” and
provided four new records from the Florida west coast. The present
report includes a Louisiana stranding in 1969.

SPECIES ACCOUNT

Balaenopiera physalus

The Gulfport Ship Channel extends from the western end of Ship
Island, a barrier island off the Mississippi coast, northward across
Mississippi Sound for 11,5 nautical miles to the Gulfport Harbor. It
has a limiting depth of 40 feet. On 7 April 1967 a baleen whale in-
vaded Mississippi Sound by the ship channel for the first time that
this event has ever been reported. This is also probably the first time
a whale has entered the bay during the past several hundred years
because the waters were too shallow until the ship channel was dug.

The general events have been described by Heiliger (1967) as fol-
lows : On the late afternoon of April 7, Captain Peter Skrmetta and
Peter B. Lasanen were returning on the Pan-American Clipper, an
excursion boat which operates between Gulfport and Ship Island,
when they saw a whale spouting near the mouth of the Gulfport Har-

169

170

Gunter and Christmas

Stranding Records of Whales

171

bor. Needless to say, this was most surprising. The next day this
whale was found to have run ashore and died inside of Ship Island.
It was later towed closer to Fort Massachusetts, the landing point on
the island, as an attraction to tourists and excursionists aboard the
Pan-American Clipper. This attraction lasted until the odor of the
decaying animal became overwhelming, when Captain Skrmetta
moved the carcass to the Cfulf beach and partially buried it with the
help of a dragline dredge mounted on a barge.

A group from the Gulf Coast Research Laboratory, including the
senior author, visited Ship Island on April 11 and inspected this
whale by going around it in a skiff. Heiliger (1967) had previously
estimated the length as 45 feet and the weight at about 12 tons. The
animal was a maturing male with a protuberant penis well over a
meter long. There w'as a good bit of blotchy white on the right side of
the anterior belly and the lower side of the flukes was white. The in-
ner flippers were also white. Some of this can be seen in the tw^o ven-
tral view photographs (Plate I, a & b). These and all other aspects of
the specimen corresponded to the finback whale, Balaenoptera phy-
salus.

After this whale had been buried for a few months its head was
dug up and exhibited at Fort Massachusetts for the delectation of the
excursionists to Ship Island. The actions of thieves and the Hurricane
Camille on August 17-18, 1969 have reduced the skull (25 June 1972)
to a remnant of one jawbone.

Records of Balaenoptera sp. from Louisiana have been summa-
rized by Lowery (1943), The Louisiana State University Museum had
a piece of baleen from a probably Louisiana or Mississippi specimen
of B. physalns taken about 1928. Several more probable and a couple
of definite records from Louisiana will be summarized by Lowery (in
press) .

Zi phiiis cu inrostris

We first received the report of a whale stranded on the Chande-
leur Islands of Louisiana from Mr. Bob Stevens who saw it on a rou-
tine inspection flight of the Gulf Islands National Wildlife Refuge in
April 1969. The carcass was located later on 15 May 1969 by J. Y.
Christmas, Tom Mcllwain and Lionel Eleuterius. It was partially
buried in the berm about five miles south of Chandeleur Lighthouse
on the Gulf beach (approx, lat. 29° 58' N. long. 88° 49' W.), Heavy
surf prevented landing on the Gulf beach, but Captain Kenneth Mel-
vin w'as able to put the party ashore from Chandeleur Sound, across
the island.

The skin had sloughed off the carcass so that muscle tissue was
exposed on some areas of the body. It lay on the right side with dorsal
fin and flukes buried. The beak was about half buried with jaws open
at 10-15°. The whole body was blackened so that natural color could

172

Gunter and Christmas

Plate 11. Upper view of skull (a) and mandibles (b) of a specimen of the
goose-beaked whale found on the Chandeleur Islands, Louisiana in April
1969. Courtesy of Dr. G. H. Lowery, Jr.

Stranding Records of Whales

173

not be ascertained. The viscera were pushed into the mouth and an
area about 46 cm in diameter around the anus was distended.

When dug out, the dorsal fin and flukes were essentially intact
but their shape was not clearly discernible. Teeth could not be seen.
Somewhat crude measurements made with a meter stick were as
follows:

Total length — 559 cm.
(18.1ft.)

Maximum body depth — 106
cm.

Tip of beak to dorsal base —
336 cm.

Tip of beak to eye — 71 cm.
Tip of beak to pectoral base-
132 cm.

Lower jaw length — 38 cm.
Upper jaw length — 35.6 cm.
Flukes, Maximum width —
112 cm.

Blowhole (located behind
eye) — 7.6 cm.

The carcass was left intact because any parts that could have
been removed could not be carried across the island to the boat.

About a month later, J. Y. Christmas and Richard Waller re-
turned and landed on the Gulf beach. Decomposition was considerably
advanced. The head, tail and left pectoral were removed, placed on a
sheet of polyethylene and dragged into the water. A piece of nylon
webbing was secured around the whole bundle and it was lifted to
the trawl deck of the boat. The return trip stank,

At the Gulf Coast Research Laboratory the bundle was buried
above normal tides. A few days after Hurricane Camille had inun-
dated the whole area we found that only shallow erosion had occurred
at the burial site, exposing the end of the beak. One tooth was found
on the ground. The head was still putrid. It was covered and not dis-
turbed again until Dr. George H. Lowery, Jr. removed the material to
Louisiana State University for museum preparation.

He has kindly furnished photographs of the cleaned specimen
which is now LSUMZ15609. Plate 11 is an upper view of the .skull (a)
and the mandibles (b) of this specimen. Doctor Lowery (personal
communication) was able to confirm our tentative identification as
Ziphi'US cavirostris. lie later carried photographs to London where
F. C. Fraser at the British Museum concurred in this identification.

LITERATURE CITED

Heiliger, Dudley P. 1967. Outdoors along the coast. (Newspaper column). The
Dixie Guide, May 1967, p. 16, (A monthly newspaper published in Gulfport,
Mississippi, now defunct.)

Lowery, George H., Jr, 1943. Check-list of the mammals of Louisiana and adjacent
waters. Occasional Papers of the Museum of Zoology (Louisiana State Uni-
versity), Number 13:213-57.

(In Press). The mammals of Louisiana and adjacent waters. Louisiana

State University Press.

Moore, Joseph Curtis. 1953. Distribution of marine mammals to Florida waters.
The American Midland Naturalist, 49(1) : 117-58.

Gulf Research Reports

Volume 4 Issue 2

January 1973

Some Effects of Hurricanes on the Terrestrial Biota^ With Special Reference to Camille
(Reprint)

Gordon Gunter

Gulf Coast Research Laboratory

Lionel N. Eleuterius

Gulf Coast Research Laboratory

DOI: 10.18785/grr.0402.04

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Marine Biology Commons

Recommended Citation

Gunter^ G. and L. N. Eleuterius. 1973. Some Effects of Hurricanes on the Terrestrial Biota^ With Special Reference to Camille
(Reprint). Gulf Research Reports 4 (2): 174-185.

Retrieved from http:// aquila.usm.edu/ gcr/vol4/iss2/ 4

This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean
Research by an authorized administrator of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

SOME EFFECTS OF HURRICANES ON THE TERRESTRIAL
BIOTA, WITH SPECIAL REFERENCE TO CAMILLE'

by

GORDON GUNTER and LIONEL N. ELEUTERIUS
Gulf Coast Research Laboratory
Ocean Springs, Mississippi

INTRODUCTION

There have been very few articles concerning the effects of hur-
ricanes upon marine and shore organisms. Some effects on fishes have
been described by Hubbs (1962) and in that paper he reviewed some
of the previous references.

Information on animals killed or injured by hurricanes is scarce
because potential observers in areas where they strike are generally
more concerned with practical personal matters than biological stud-
ies right after a bad storm. The senior author has been in or very
close to seven West India hurricanes as they came ashore. Each time
he was somewhat forewarned and had determined to make some type
of quantitative appraisal of killed animals following these storms.
However, on no occasion has this been done. Nevertheless, the tw^o
waiters have collected some fragmentary information worth record-
ing.

Some Damages to the Fauna on the Mississippi Coast
Hurricane Betsy — 9-10 September 1965

When Betsy passed the Mississippi Coast on its way to devastating
areas of New Orleans, the center was approximately 50 miles south of
Horn Island. Thus the wind blew more or less from east to west along
the Ocean Springs^Pascagoula coastal area. Following this storm hun-
dreds of sea balls mostly of the marsh grass Spartina were found on
the mainland beach of Mississippi Sound near Gulfport.

The water rose to a height of about six feet on the Laboratory
grounds and left a strand of debris along the beach. In areas where
there was marsh grass, thousands of little drowmed mice, the Eastern
Harvest Mouse, Reithrodontomys humuliSy lined the shore in a little
browm wundrow which w'as sometimes 100 yards long without a break.
Also lying on the beach every ten yards or so w^as a dead raccoon, Pro-
cyon loto7\ There were so many on the Laboratory grounds that they

^ This is a verbatim reprinting of the paper with the same title published in
Volume 3, Number 2 of this journal, which was so improperly laid out that re-
prints with the plates could not be made and the legends were missing.

174

Hurricane Effects on Biota

175

had to be hauled away. Raccoons are good swimmers and they cer-
tainly did not come from the surrounding nearby marsh-land, Horn
Island, which lies eight miles offshore, was either completely or nearly
submerged during this storm. The most reasonable assumption is that
these coons were drowned in Mississippi Sound, after being washed
off of Horn Island and their bodies were finally blown onto the main-
land shore.

Hurricane Camille — 17-18 August 1969

This has been publicized variously as the worst storm that ever
struck North America, or as the worst that has come ashore in this
country in 143 years. Old, indefinite accounts indicate that something
like this struck Florida in the 1700s. It is quite certain that Camille
was the most powerful hurricane that has struck a w’ell-populated
shore of the United States. The weather planes flying through it
clocked the winds at 218 mph and recorded the lowe.st natural baro-
metric pressure that has ever been read (26,01 inches) .

The ''Big House,” an old landmark of the Laboratory and of the
coast, was splintered and even the brick pillars upon which it stood
were washed aw'ay.

We have no way of quantifying the destruction of animals, ex-
cept to say that the clean-up agency, the 43rd Battalion, Corps of
Engineers, U. S. Army, reported removing 28 tons of animals from
the beach between Biloxi and Uulfport on August 22-24, Most of
these animals were dogs and cats, but some horses and cattle were
mixed in. After the storm many dogs were homeless and many were
systematically shot because they were starving.

Following the storm, the writer was waked up by a bird singing
lustily just outside his window; this was the only bird seen for about
a w^eek. An unknown number of wild animals, birds, dogs and other
life including human beings lost their lives in the storm; after about
three days, the odor of decaying animals was noticed in the atmos-
phere and lasted about a week before it gradually went away.

The bird population and the squirrel population virtually disap-
peared but both came back after a few weeks time, most noticeably
the jaybirds and a few gray squirrels {Sciurus carolinensis) . The
birds disappeared again, probably because they could find nothing to
eat. This was certainly true of the squirrels and they w^ere reduced to
gnawing the bark off of felled water oaks (Gunter and Eleuterius
1971).

When the storm struck, the seeds of various nut trees — chiefly the
hickory, black walnut, and thousands of pecans which are planted in
this area^ — were just beginning to mature. Many of these were blown

176

Gunter and Eleuterius

down and approximately half of the foliage of those remaining was
denuded by breakage of the limbs. The same thing was true of oaks
and acorns. Additionally^ the nuts themselves were beaten off the
trees that remained standing. Presumably for that reason the Eastern
Gray Squirrel, which was quite common, had not returned in its for-
mer numbers by April 1970. Before the storm it was quite common to
see as many as eight of these at one time in a relatively .small area of
trees in the senior author’s front yard. After the storm, he saw none
for one week and then he saw a lone squirrel. The squirrel population
apparently increased in about three weeks to a month after the storm,
then declined again. This observation would bear out the supposition
that squirrels moving in from other areas could not find sufficient
food and moved out again. The same thing apparently was true of the
jaybirds.

There was a decided diminution in the number of birds which
came to feeder stations during the following winter. For instance,
dozens of birds and sometimes a few' hundred in one afternoon for-
merly fed at a home facing the beach just to the side of the Labora-
tory grounds. The most numerous species, sometimes present in the
dozens at a time, was the Savannah Sparrow. During the past winter
only three or four have appeared at a time. The owner, before Camille,
had to keep watch on the Starlings and jaybirds because they dis-
turbed and ran off the others, but has had no trouble since the storm.
General observations show' that the Brown Thrashers, the jaybirds
and the Cardinals are present in very diminished numbers even today
(April 1970).

These facts have been noticed by other people and recorded, espe-
cially in The Dixie Guide by Mr. Clayton Rand who has gone through
three bad hurricanes at his home in Gulfport. Mr. Rand has men-
tioned in his paper several times, the last being February and March
1970, that during former hurricanes there were many snakes and
frogs everywhere in the area and that the mosquitoes were quite bad.
He has remarked three times in his monthly newspaper that there was
a great absence of life following Camille, even of the birds.

To the senior writer, however, the most amazing thing has been
the disappearance of the ants up until this time (April 1970). The
black carpenter ant and the Argentine fire ant and several other
smaller species were quite common in his yard. Apparently they all
succumbed to the storm, e.\cept for a minute yellow species that goes
by the name of sugar ant, which has been seen one time. Bread and
other foods set out for dogs and cats w'ere formerly covered with ants
in a matter of minutes; but, even this long after the storm, they may
remain untouched by ants for days. We do not know the extent of de-
struction of the Argentine fire ant, but locally they are gone.

It is to be expected that termites and termite feeding animals and

Hurricane Effects on Biota

177

possibly woodpeckers would increase greatly in numbers due to the
thousands and thousands of felled trees and rotting timber, a good bit
of which, after having had the top broken off, is still upright.

Damages to the Flora on the Mississippi Coast

There are very few reports of the effects of hurricanes, typhoons,
or cyclones (tornadoes) on coastal vegetation. Sauer (1962) reported
the effects of cyclones on the coastal vegetation of a tropical island
(Mauritius) in the Indian Ocean. Chamberlain (1959) and the U. S.
Department of Agriculture (1960) reported some of the effects of
Hurricane Audrey on the vegetation of south Louisiana. Previous hur-
ricanes which struck the Mississippi coast inflicted minor damage to
the vegetation; one of the worst of these storms known to the junior
author occurred in 1947.

The “eye” or center of Hurricane Camille came ashore in the
Pass Christian-Bay St. Louis area and the path was well marked by
the effects of the storm on vegetation. The most apparent and obvious
effect was the destruction of the ti’ees. In Jackson County most of the
trees blown down were oriented with the tops pointing toward the
northwest. In Harrison County near Gulfport, the trees became ori-
ented with the tops pointing toward the west-northwest and in the
Pass Christian-Bay St. Louis area, they were oi’iented in an east-west
direction, but some tree tops pointed eastward and some pointed to the
west and the trees were nearly parallel in alignment (Figs. 1 and 2).
The paradoxical alignment was apparently a result of the initial
winds from the east, followed after the “eye” passed over the area, by
winds from the west. Trees west of Bay St. T^ouis near Pearl River
were oriented with the tops toward the east-northeast and near Sli-
dell, Louisiana, they were down in a northeast direction.

The intensity of winds from Hurricane Camille could be seen in
the number of trees felled, the number increasing as the wind velocity
increased toward the path of the “eye.” In fact, without referring to
other data, one could determine the storm’s path by observing the
east-west direction in which the trees were blown down and by the
gradual increase in the numbers of trees destroyed as the center of
the path was approached.

Tornadoes or extremely turbulent winds ripped through many
areas on the periphery of the hurricane and the paths of their “touch
downs” were well documented in the vegetation. In Magnolia State
Park, which almost adjoins the Laboratory propei*ty, there is one area
50 feet wide and 17 tree lengths long, which the second author at-
tributed to these tornadic gusts.

The junior author conducted two vegetatioiial surveys to compare
the intensity of damage to areas on the periphery of Hurricane Ca-

178

Gunter and Eleuterius

mille with areas nearer the center. In Jackson County, these surveys
showed that in one tract, 4% of the trees were blown down and 10%
were damaged to the point that survival was in question. The plant
community was dominated by Quercus yiigra (water oak) with Piniis
elliotii (slash pine), Carya glabra (hickory) and Qmrcus rubra (red
oak) being the subdominant species. This 40-acre tract in Magnolia
State Park was approximately 22 feet above sea level. Destroyed trees
in decreasing order were: red oak, slash pine, water oak, and hickory.
It was noted that the heart wood (xylem) of the red oaks had been
weakened by pathogenic attack and were rotted. Dess than 10% of
the pines destroyed were uprooted; they were twisted or broken off
at heights ranging from 5 to 20 feet above the ground. The large tap-
root characteristic of the pines apparently held the trees up; they
were not blown down easily, but could be broken. Other trees blown
down in adjacent plant communities were Magnolia grandiflora (mag-
nolia), Nyssa biflora (black gum), Liquidamhar styraciflora (sweet
gum) , and Lireodendrum tulipera (tulip tree or yellow poplar) .

Another survey was conducted on 87 acres of forested land north
of Pass Christian in Harrison County, bordering the Wolf River and
Red Creek Road. Approximately 10 acres here was bottomland forest
along the river and adjacent low-lying drainage areas. The rest of the
land was approximately 25 feel above .sea level and covered with Piniis
elliotii (slash) , Pinus taeda (loblolly) and Pimis palustris (longleaf)
in various stages of growth. The owner considered the area a game
reserve and left it undisturbed. Results of a sample showed that ap-
proximately 70% of the bottomland species were blown down. The
species were Magnolia virginiana (sweet bay) Liquidamhar styraai-
flora (sw'eet gum), Taxodium distichum (bald cypress), Acer rubrum
(red maple), and the area was dominated by Quercus nigra (water
oak) . Ninety per cent of the trees in the low-lying area had diameters
greater than 24 inches at breast height and there were between 100
and 150 trees per acre. An estimated total of 201,000 board feet of
hardwood timber was lost.

Approximately 10,000 slash, loblolly, and longleaf pine trees with
diameters greater than 10 inches were present on the higher sites and
there were only 300 of these trees that were not damaged, i.e. 97 %
were destroyed. Many of those standing were not expected to survive
due to lack of limbs, missing tops or split trunks. A total of 607,600
board feet of pine was estimated as lost. Many young trees were
crushed by the falling trees, and other understory plants and habitats
for wildlife were destroyed. At the time of the survey (March 1970),
beetles, especially Ips avuhu^, bps grandicoUis, and Ips calligraphns,
had infested many of the downed trees and rot had begun. The specific
names of the beetles were furnished by Dr. Virgil Smith, entomologist,
U. S. Forest Service, Gulfport, Mississippi. Twisted and split saw logs
could not be salvaged for use. Paper wood operations were expected to

Hurricane Effects on Biota

179

be hindered by the tangled mass of trees. Practically all of the pine
trees were second growth and ranged from 16 to 68 years old. The
water oaks and other hardwoods were much older, ranging from 100
to 125 years.

These two tracts simply show by comparison that the most dam-
age to the vegetation was caused by winds occurring near the center
of Hurricane Camille’s path.

Another observation was the destruction of Querciis virginiana
(live oak) along the Peach front from Biloxi to Pass Christian. Ap-
proximately 25,000 live oaks were growing along the beach before
Camille and one-fourth were destroyed by wind and water and one-
half were damaged. Those trees nearest the beach were partially in-
undated and the roots eroded by wave action. The immediate beating
action of wind and the physiological “drought” resulting from the
salt spray reduced these evergreens to bare branches (Figs. 3 and 4) .

Many slash and longleaf pines may have been killed as a result of
the inundation of low-lying areas near the mouth of the Wolf and
Jourdan Rivers. The trees are dead but standing; however, this could
be the result of other, internal damage since many trees on the barrier
islands were covered by salt w'ater and survived. This observation
needs further study.

The Corps of Engineers, U. S. Army, estimated that a total of
1.2 million board feet of saw timber and one million cords of pulp-
wood in MissLssippi w'ere lost. On the Mi.ssissippi Test Facility in Han-
cock County, an estimated 6,000 cords of pulpwood were damaged and
only 60 of the downed trees could be salvaged for lumber. It has
been reported that a total of 290 million cubic feet of pine alone was
lost in South Mississippi (Van Hooser and Hedlund 1969).

The barrier islands presented a pattern of destruction similar to
that on the mainland. Petit Bois Island was affected relatively little
but there was a gradual increase in damage on the islands to the west.
Horn Island was heavily eroded on the outside beaches. The marsh
vegetation was pushed dowm and pressed to the soil .surface by the
w’ater as it passed over the island (Figs. 5 and 6). Ship Island was
cut into three pieces and more than one-third of the vegetation, most
of which was herbaceous, was removed. Cat Tsland was heavily dam-
aged. Large oaks were uprooted by wave action and many pines were
broken by the wind. Large sand dunes were leveled, the sand redis-
tributed over much of the adjacent low-lying marsh. Tons of plant
materials .swept from the Louisiana marshes and the barrier islands
were deposited on the mainland in large windrows.

Marshlands were affected insignificantly because the water cov-
ered them early in the hurricane and they were not exposed to the
terrific beating of wind and wave that occurred later. Spartina alter-

180

Gunter and Eleuterius

niflora (smooth cord grass) flowered on schedule (September through
November). Shrubs found along the periphery of marshes, where they
formed thickets, acted as baffles and protected trees and, in some
cases, homes. Many upland understory areas were denuded of her-
baceous and woody shrubs where they were located near water.

The botanical regime of South Mississippi was disturbed by Hur-
ricane Camille of August 1969, probably to a greater extent than by
any other hurricane in the history of Mississippi, and the greatest in-
fluence on the terrestrial vegetation was the destruction of the trees.

Hurricane Effects on Biota

181

Figure 1. Heavily damaged pine stand in Hancock County show-
ing parallel but opposite direction alignment of fallen trees. Note
direction in which standing trees are leaning.

Figure 2. Heavily damaged pine stand illustrating parallel but
opposite direction alignment of fallen trees. This effect was
caused by passage of the “eye” of Hurricane Camille through
Bay St. Louis-Pass Christian area.

182

Gunter and Eleuterius

Figure 3. Damaged live oaks (Quercus virginiana) along Highway 90 near Long Beach,
Mississippi.

Hurricane Effects on Biota

183

Figure 4. Damaged home and live oak (Quercus virginiana)
along Highway 90 and open waters of Mississippi Sound at
Long Beach, Mississippi.

184

Gunter and Eleuterius

Figure 5. Marsh near the south beach of Horn Island. Altitude
approximately 1,500 feet.

Figure 6. Low altitude view (600 feet) of same marsh shown in
Figure 5. Note flattened plants of J uncus roemeriamis and Spartina
alterniflora as a result of wave action across island.

Hurricane Effects on Biota

185

LITERATURE CITED

Chamberlain, J. L. 1959. Influence of Hurricane Audrey on the coastal marsh of
southwestern Louisiana. Coastal Studies Institute, Louisiana State Univer-
sity, Technical Report, lOB, ONR 35608.

Gunter, G. and L. Eleuterius. 1971. Bark eating^ by the common gray squirrel fol-
lowing a hurricane. Amer. Mdl. Nat. 85(1) :235.

Hubbs, Clark. 1962. Effects of a hurricane on the fish fauna of a coastal pool and
drainage ditch. Tex. Jour. Sci. 14(3) :289-96.

Sauer, J. D. 1962. Effects of recent tropical cyclones on the coastal vegetation of
Mauritius. J. Ecol, 50:275-90.

U. S. Dept, of Agr. Soil Conservation Service. 1960. Effects of saline water from
Hurricane Audrey on soils and vegetation. Alexandria, La., Special Rept.
(Minco).

Van Hoqser, Dwane D, and Arnold Hedlund. 1969. Timber damaged by Hurricane
Camille in Mississippi. U. S. Forest Service Res. Note. 80-96:1-5. Southern
Forest Experiment Sta., New Orleans, La.

Gulf Research Reports

Volume 4 Issue 2

January 1973

Protozoan Symbionts from the Anemone Bunodosoma cavernata from Galveston Island^
Texas

Philip J. Phillips

Texas A6’M University

DOI: 10.18785/grr.0402.05

Follow this and additional works at; http://aquila.usm.edu/gcr

Part of the Marine Biology Commons

Recommended Citation

Phillips^ P. J. 1973. Protozoan Symbionts from the Anemone Bunodosoma cavernata from Galveston Island; Texas. Gulf Research
Reports 4 (2): 186-190.

Retrieved from http:// aquila.usm.edu/ gcr/vol4/iss2/ 5

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PROTOZOAN SYMBIONTS FROM THE ANEMONE
Bunodosoma cavernata FROM GALVESTON ISLAND, TEXAS

by

PHILIP J. PHILLIPS!

Department of Biology
Texas A&M University
College Station, Texas 77801

Hargitt (1904), in his classic paper on the medusae of the Woods
Hole region, noted the susceptibility of medusoid forms to various pro-
tozoan parasites. He stated that “when working upon the regeneration
of medusae I found several species of Protozoa very closely associated
with them and, under the limitations of the aquaria, often exceedingly
troublesome, seriously interfering with the progress of the experi-
ments. This suggested the possibility of a parasitic relation.” Unfor-
tunately Hargitt made no mention of the taxa of Protozoa involved.
Studies regarding protozoan infestations of Cnidaria have been few.
Other than the passing references to protozoan infestations by Har-
gitt (1904) and Kudo (1966) there have been no intensive studies of
protozoan parasites or symbionts of marine Cnidaria.

The anemone Bunodosoma cavernata is a common inhabitant of
the Texas coast and cursory observation revealed a large complement
of protozoan associates. These included Saprodinium, Euplotes^ Pa-
raeuplotes, Vorticella, Cohnilembun, Aiiophrys, Uronema and Vahl-
kampfia.

Approximately forty anemones were collected from jetty rocks on
Galveston Island on 3 November 1970. Note was made within one day
of protozoans found on the external surfaces, in the coelenteron and in
surrounding sea water. The anemones were kept until 7 December
1970 under varying conditions and bi-weekly note was made of the
associated protozoan fauna.

One group of six anemones was placed in a 29-gallon aquarium
in which the salinity was maintained at 28 g/kg and the temperature
ranged from 15 to 18 °C. These anemones were considered “normal.”
No attempt was made to construct a facsimile of the Galveston littoral
environment from which the anemones were removed. Other groups of
three to eight anemones were kept in 1.5-gallon aquaria (salinity 28
g/kg, temp. 18 °G) and in 12-inch finger bowls wherein salinity and
temperature were varied. For one group (GP 1) the salinity was al-
lowed to rise by evaporation from 20 g/kg to 40 g/kg over a 12-day
period. In a second group (GP 2) the sea water was flushed out and

! Present address: North Atlantic Division, II.S. Army Corps of Engineers,
99 Church Street, New York, N. Y. 10007.

186

Protozoan Symbionts of Anemone

187

replaced with clean sea water of the same salinity every 3 days for a
3-week interval. In the third group (GP 3), the salinity was constantly
maintained at 24 g/kg for 4 weeks. All three groups were maintained
at 18 ”C. In a fourth group temperature was maintained at 24° C. Sa-
linity was routinely monitored on a daily basis with a Goldberg re-
fractometer. Microscopy examinations were made every 2 to 4 days
using wet mounts of anemone mucus, coelenteron contents, tissue
squashes and the external medium.

Protozoans found as epizoites or symbionts on the anemones as
well as site of infection and relative abundance are listed in Table 1.

Table 1.

Protozoa Found in Association with Anemones after Two Weeks in

the Laboratory

Protozoan

Site

Wounds

of Infection
Coelen- Mucus

teron Coat

Size Range
(microns)

Vahlkampfia

XXX

XX

XX

7-30

Saprodinium

XX

X

X

21-25

EupZotes

XXX

XXX

XXX

70-140

Paraouplotes

X

X

X

50-80

Varameeiwn

X

100-150

VoYtioella

XX

XX

XX

C.150

Cohni lembus

X

X

XX

30-80

Anophrys

XX

XX

XX

60-90

Vronema

X

X

X

20-40

X indicates rarity; XX indicates frequent occurrence;
XXX indicates dominant organisms numerically.

These data were taken 2 weeks after the anemones were removed from
the natural environment and introduced into the laboratory. Initially
when first placed in aquaria all Protozoa found in association with the
anemones occurred in very low numbers when compared to the situa-
tion after 2 weeks. Protozoa in habitat sea water included very sparse
populations of Eiiplotes, Pammecmm and Vorticella. The normal ane-
mones maintained a numerically smaller symbiont population than
those in the finger bowls. Those in the 1.5-gallon aquaria had a “nor-
mal” symbiont complement also. In GP 2, where the water was flushed
out and replaced periodically, the anemones retained the normal fauna

188

Phillips

and did not support large populations of any species of protozoan. In
the group maintained at 24°C the anemones became moribund after
two weeks and developed extremely large populations of ciliates, es-
pecia.\\y Eitplotes, Arwphrys and Saprodmiimi, as well as large num-
bers of the naegleriid amoeba Vahlkampfia. In addition to being found
in necrotic and normal anemone tissues this amoeba was also found in
large numbers in the fouled medium. Both cysts and trophozoites of
Vahlkampfia were extremely numerous at sites of necrosis. In group 3
(GP 3), wherein conditions were constant, the anemones survived up
to 4 weeks at which time they became moribund and died. The same
situation prevailed here as with respect to GP 2. In GP 1 increased
salinity had no apparent effect on protozoan numbers although indi-
vidual EuiAotes showed considerable size increase (up to 250 p in
length). Size ranges of all associative Protozoa encountered are given
in Table 1.

All anemones within the finger bowls died within 5 weeks. In all
cases the anemones were infiltrated by large numbers of bacteria and
large populations of Enplotes and Vahlkampfia. Approximately half
the Euplotes in association with necrotic anemone tissue, were in the
process of conjugation.

Three morphological types of Vahlkampfia were observed. Most
amoebae were 7-10 p in length and formed only one broad, fan-shaped
pseudopod which, moreover, consisted mainly of a large hyaline area.
P^ormation of a new pseudopod was preceded by flooding the old pseu-
dopod with granular endoplasm and the formation of a new^ hyaline
bulge. Rate of pseudopod formation was hastened by the heat of a
substage illuminator. The second observed trophozoite stage was con-
siderably larger (25-30 p) and had numerous psuedopodia. Their
morphology approached that of the former type after being on a mi-
croscope slide (above an illuminator) for several minutes. These
larger amoebae underw'ent division and “reverted’’ to the smaller, mo-
nopseudopodial type. On two occasions a nucleus (2-3 p in diameter)
with a large endosome was observed in the large amoebic variant.
Large numbers of Vahlkampfia cysts were found in decaying and
moribund anemones as well as in putrefying media. After being on a
microscope slide for about 5 minute.s, above a substage illuminator,
these cysts released one amoeba each. Cysts were rounded, consider-
ably flattened, had wrinkled surfaces and Nvere between 5-9 p in diam-
eter. Vahlkampfia invades anemone tissue and will emerge from tissue
fragments when heated by substage illumination for a few minutes.
The amoebae can be demonstrated in both normal and necrotic
tissue, the greatest concentration of amoebae occurring in necrotic
tissues with a large bacterial population. The amoebae were observed
to ingest bacteria, indicating that greater degree of association with
necrotic tissue may be due to greater availability of food organismvS in
necrotic as opposed to healthy tissue. Other Protozoa, especially Eu-

Protozoan Symbionts of Anemone

189

plates, showed greater abundance where there w^as high bacterial
density.

Vahlkampfia and at least a few of the ciliates (Euplotes, Paraeii-
plotes, Saprodinium and Vorticella) are not obligate symbionts. Nae-
gleriid amoebae are known as facultative parasites from a wide vari-
ety of organisms including rodents (Wilson et al. 1967), molluscs
(Hogue 1921 and Richards 1970), insects (Page 1970) and man
(Cerva and Novak 1968 and Duma et al. 1969). Apparently amoebae
of this group can be demonstrated to be symbiotic with most any
metazoan. Although Naegleria gruberi is the only naegleriid amoeba
definitely known to be a causative agent in human amoebic meningoen-
cephalitis (Duma et al, 1969 and Cerva 1970), the remaining naegle-
riids cannot be discounted as potential pathogens. Wilson et al. (1967)
demonstrated that Vahlkampfia as well as many other genera of amoe-
bae can cause systemic amoebiasis in rodents following hypodermic
inoculation. Available data indicate that Vahlkampfia, and at least
some of the ciliates in this study feed on broken tissue and associated
bacterial flora. The external mucus coat or pellicle of the anemone on
normal anemones has a relatively large bacterial population and this
may account for the establishment of larger populations of Protozoa
in this region when compared to other anatomical regions of the
anemone.

The hymenostomid genera Cohnilemhiis and Anophrys are inter-
esting in that they are known to-be intestinal symbionts of echinoids
(Kudo 1966). These two genera and Uronema predominantly occur in
the coelenteron and mucus coat. Paraeuplotes, found here on anem-
ones, is also a known epizoite of corals (Kudo 1966).

In the normal littoral habitat the anemones are periodically ex-
posed to the air during tidal cycles and there is considerable scouring
of the anemone surface as well as drastic short term salinity and tem-
perature changes in the immediate environment of the anemone.
These environmental factors probably serve to control bacterial and
protozoan populations epizootic on the anemones. Conversely the mu-
cus sheath of the anemone, which serves for attachment of protective
shell fragments and other debris to the anemone surfaces, may pre-
vent desiccation at low tide and may act as a suitable substrate for the
establishment of epizoites. When exposed the anemones close the oral
apparatus, trapping sea water and any associated microfauna possibly
allowing survival of some associative forms. It remains to be deter-
mined whether or not any of these symbionts or epizoites are depen-
dent on the anemone for life cycle completion.

Euplotes is by far the most abundant of all the ciliates encoun-
tered and conjugation is a commonplace phenomenon especially when
putrefaction is well advanced. It is possible that death of the anemone
enables or induces conjugation in Euplotes,

190

Phillips

Anemones of the littoral zone support an extensive protozoan
fauna. Although none of the Protozoa involved are definitely known
to be obligate symbionts there is a distinct association. Most of the
forms are facultative symbionts, especially VahlJcampfia and Euploteis.
Vahlkampfia invasion of anemone tissues can to some extent be asso-
ciated with necrotic tissue changes and increased bacterial popula-
tions, Most probably bacteria initiate the necrosis and the large bac-
terial populations allow for increase in the amoeba population. No
causal relationship has been established.

ACKNOWLEDGEMENTS

I would like to thank Doctors K. Horvath and S. H. Hopkins (De-
partment of Biology, Texas A&M University) for critical review of
the manuscript.

LITERATURE CITED

Cerva, L. 1970. Comparative morpholog-y of three pathogenic ^strains of Nacgleria
gruberi. Folia Parasit. 17:127-33.

Cerva, L. and Novak. 1968. Amoebic meningoencephalitis: sixteen fatalities.
Science 160:92.

Duma, R. J., H. W. Ferrell, E. C, Nelson and M. M. Jones, 1969. Primary amoebic
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Gulf Research Reports

Volume 4 Issue 2

January 1973

Some Analyses of Twentieth Century Landing Statistics of Marine Shrimp of the South
Atlantic and Gulf States of the United States

Gordon Gunter

Gulf Coast Research Laboratory

Katherine McGraw

Gulf Coast Research Laboratory

DOI: 10.18785/grr.0402.06

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Marine Biology Commons

Recommended Citation

Gunter^ G. and K. McGraw. 1973. Some Analyses of Twentieth Century Landing Statistics of Marine Shrimp of the South Atlantic and
Gulf States of the United States. Gulf Research Reports 4 (2): 191-204.

Retrieved from http;// aquila.usm.edu/gcr/vol4/iss2/6

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SOME ANALYSES OF TWENTIETH CENTURY LANDING
STATISTICS OF MARINE SHRIMP OF THE SOUTH
ATLANTIC AND GULF STATES OF THE
UNITED STATES

by

GORDON GUNTER and KATHERINE McGRAW
Gulf Coast Research Laboratory
Ocean Springs, Mississippi

ABSTRACT

There is a strong correlation between the total catch of white and
brown shrimp with dockside prices on the United States Gulf Coast
since 1902, but there is no significant correlation between South At-
lantic production and prices, probably because the South Atlantic
shrimp stocks have been over-fished since the 1920s. There is no nega-
tive or positive correlation between the catch statistics of brown and
white shrimp of the United States, and these species seem to be weak-
ly competitive, if at all There is a significant correlation between the
annual production of South Atlantic and Gulf white shrimp, but there
is none between South Atlantic and Gulf brown shrimp, possibly be-
cause the brown shrimp live generally in deeper water and are not so
much influenced by short term variations in climatic conditions as the
white shrimp are in shallow water. In furtherance of this idea, there
is some indication that the brown shrimp production is less variable
than the white shrimp production.

INTRODUCTION

Three species of shrimp of the Family Penaeidae (Genus Pe-
naeiis) are present in considerable numbers and in overlapping dis-
tributions in the bays and oceanic shallow waters from Cape Hatteras,
North Carolina, south to Texas and beyond. These are the white
shrimp Penaeus fluviatilis, the brown shrimp P. aztextis, and the pink
shrimp P. duorarivm.

Another shallow water penaeid Penaeus brasiliensis exists only in
such small numbers off Miami, Florida that it was overlooked by bio-
logists of the area until discovered there by Eldred (1960), A fifth
species of the Penaeidae, XiphopeneMs kroyeri, is almost entirely shal-
low oceanic in divStribution with a few entering the bays in cool weath-
er (Gunter 1950) . It is not found along the South Atlantic part of the
United States in commercial concentrations, but has been fished in the
Gulf since boats and seines large enough to fish the shallow offshore
waters have been available.

191

192

Gunter and McGraw

The white shrimp grows to large size in shallow waters of the
bays. The other two species of commercial shrimp, P. aztecus and P.
dnorarum, do not grow so large in the bays and shallows and do not
school as strongly as the white shrimp and the seabob. They also go
into deeper waters when they move into the open ocean.

SOME HISTORICAL ANTECEDENTS

Indians caught shrimp with the use of dipnets, seines and leafy
weirs such as are still employed in the Rio Soto la Marina, Mexico.
Shrimp from the North Carolina waters were caught and transported
to the Philadelphia market w'hen Thomas Say (1817) first described
the North American white shrimp.

Catch statistics on the commercial fisheries were collected only
after the organization of the United States Fish Commission by S. F.
Baird and others in 1871. We may assume with complete assurance,
however, that shrimp production grew with the increase in population
up until recent years. Even in the early part of this century the catch-
ing of shrimp was by means of dipnets, seines, and castnets. For this
reason only the white shrimp P. ftiiviatilis and the seabob Xiphope-
neiis kroyeri were taken, because they were schooling shrimp. Even
so the seabob has been taken in small numbers amounting to about
1.2% of the Gulf catch, (cf. Gunter 1962) partly because of its small
size and its open ocean distribution. This shrimp is much more im-
portant, relatively, in South American waters (cf. Lindner 1957).

The otter trawl came into use along with motor vessels on the
South Atlantic Coast during the period of World War I and spread
quickly to the Gulf Coast. This permitted the fishing of deep waters
and larger shrimp, which move out as they grow older. Thus, produc-
tion gradually rose with the increase of demand and the more efficient
otter trawl put the large seine crews out of business in Louisiana in
the early 1930s.

From 1902 the shrimp production in this country increased into
the early 1950s. In the 1940s an extreme drought caused a great short-
age of white shrimp, especially in Texas waters, and there fishermen
turned to the previously unfished brown shrimp which were caught
predominantly at night. Most states had laws against shrimping at
night for the protection of the white shrimp, the idea being that they
should not be harassed all hours of the twenty-four. The large brown
shrimp generally bury in the bottom during the day. Recognition of
these facts led to exploitation of the brown shrimp and after the early
'50s it has yielded more than the white shrimp. This development be-
gan in Texas waters in 1947 and spread quickly to other areas on the
Gulf and South Atlantic Coast. Even so, the separation of the brown
and white shrimp was not begun in the federal fisheries statistics until

Shrimp Landing Statistics

193

1957. Therefore, we may say that the shrimp production figures used
here were comprised almost entirely of white shrimp from 1903 to
1948, with about being seabobs. From 1948 to 1957 there was a
period of production when the brown shrimp and white shrimp were
not separated. After 1957 these shrimp have been separated in the
catch statistics of the South Atlantic and Gulf Coasts. At that time
the seabobs were also separated in the statistics.

From 1951 to 1956 inclusive, the heads-off weight of white and
brown shrimp produced ranged between 126 and 146 million pounds
and in the 1967-71 period it ranged from 125 to 137 million pounds.
These are the only years, except for 1963, that the United States
shrimp production has ever ranged above 100,000,000 pounds of head-
less shrimp. The 1951-56 high production was due to the exploitation
of the previously unfished population of brown shrimp plus the white
shrimp. The more recent high production seems to be due to an in-
crease in the white shrimp population, cau.sed po.s.sibly by a recent
hyperfertilization of the bays.

DISTRIBUTIONS AND CATCH RECORDS

There are many interesting things about the distribution of the
shallow water penaeids along the coasts of the South Atlantic and
Gulf states and Mexico, but here we are concerned only with the
brown shrimp P. aztecus and the white shrimp P. fltiviatilis^ because
these two have been the chief commercial producers and they both
grow up in estuarine areas. Furthermore the United States population
of brown and white shrimp are quite discrete and disconnected from
other populations, and we have United States production of these two
species unmixed with foreign populations.

The white shrimp population of the United States is divided into
two distinct parts. The South Atlantic component runs along the coast
from North Carolina with the greatest abundance in Georgia and
gives out at about the St. Lucie inlet in south Florida (Gunter and
Hall 1963). The second population extends from the west Florida
panhandle to Aransas Bay, Texas.

The brown shrimp has roughly the same distribution but it is
le.ss numerous on the Atlantic and extends farther south seasonally
in the Mexican waters. Its abundance is greater in the salt waters of
Texas than that of the white shrimp, which is most abundant in Lou-
isiana because of the lower salinities in that region. In Texas waters
brown shrimp are not raised in appreciable numbers farther south
than the Aransas-Corpus Christi Bay system, which is connected to
the Gulf by Aransas Pass. During the fall both species leave the bays
and go to outside waters. Gunter (1962) showed by following the
seasonal catch statistics of four areas on the coast that the white and

194

Gunter and McGraw

brown shrimp go south on the Texas coast in the early fall and winter.
Some go into Mexico and return in diminished numbers in the spring
to Texas waters. Catches made off northern Mexico are returned to
United States ports. This movement apparently begins off Galveston
Bay and covers a distance of some 400 miles and it is virtually a
parallel case to the seasonal north to south white shrimp migration
and return from Georgia to the region of Cape Canaveral discovered
by Weymouth, Lindner and Anderson (1933) (Lindner and Anderson
1956).

Pink shrimp exist in fair concentrations off North Carolina and
in heavy concentrations off the Tortugas. There are also large concen-
trations in the Bay of Campeche, Mexico, which were formerly fished
by Florida, Texas, Cuban, and Mexican fishermen, and adequate statis-
tics are not available. Former United States catch statistics of this
species were confused by Florida and Texas boats bringing in Cam-
peche shrimp. Furthermore Gulf and Atlantic catches were confused
by shrimpers carrying some shrimp from Tortugas to Atlantic ports.
For these reasons we have avoided use of pink shrimp statistics. As
grooved shrimp they were mixed with the browns to a small extent in
the late 1950s but not enough to vitiate the brown shrimp statistics.

SUMMARY OF THE PROBLEM

The brown and white shrimp both grow up in the bays of the
northern Gulf Coast and the South Atlantic states. They have a differ-
ential distribution with relation to salinity and season (Weymouth,
Lindner and Anderson 1933, Gunter 1950, 1961, Gunter, Christmas
and Killebrew 1964) . The white shrimp come in and move out later in
the year. Furthermore the white shrimp grow to larger size in the
estuaries and, therefore, are more heavily fished before they move out-
side. As a matter of fact the whole shrimp industry grew up in the
shallows and gained technical experience on the white shrimp before
moving to the open sea.

Because of the overlapping life history of these two species of
commercial shrimp, both in time and place, the question has arisen
concerning their competition. Therefore, some who have been con-
cerned with shrimp biology have discussed the.se matters for years,
mostly with the suspicion that there was some kind of competition
that opposed one shrimp population to the other. These ideas were the
genesis of the analyses offered here.

All shrimp statistics used here were taken from the annual Fish-
ery Statistics of the United States and its predecessors, of which the
latest issue is Lyles (1969) , and preliminary pamphlets.

Shrimp Landing Statistics

195

PRICES AND PRODUCTION

One would think that prices increased with expansion of produc-
tion, the demand for shrimp, etc., and such is the case where total
United States production and price are concerned. The coefficient of
correlation, r, for the figures shown in Table 1 is 0.691 with 39 obser-

Table 1.

The Total Catch of White and Brown Shrimp of the Gulf and South
Atlantic Coasts of the United States in Thousands of Pounds and the
Dockside Value in Thousands of Dollars

Catch in Catch in

Year Pounds Value Year Pounds Value

1902

10,506

1908

11,855

1918

40,632

1923

45,987

1927

64,200

1928

74,986

1929

70,487

1930

57,219

1931

62,628

1932

57,313

1934

77,479

1936

76,520

1937

90,866

1938

96,150

1939

96,150

1940

97,754

1945

122,743

1950

122,048

1951

143,780

1952

145,414

286

- 1953

408

1954

1,746

1955

2,593

1956

3,518

1957

4,550

1958

4,435

1959

2,996

1960

2,731

1961

2,036

1962

3,067

1963

3,778

1964

5,009

1965

4,848

1966

4,848

1967

5,895

1968

21,289

1969

43,144

1970

51,518

1971

54,755

145,414

76,267

172,596

60,535

156,454

61,404

142,297

70,305

90,364

72,438

89,903

71,829

108,548

56,875

112,088

66,143

64,234

50,589

77,788

71,832

112,535

68,785

95,813

69,328

111,643

81,067

107,041

93,784

137,837

99,584

124,480

109,833

126,331

117,317

139,437

119,569

148,125

143,362

vations and 37 degrees of freedom. This means that prices and pro-
duction have grown together, and the correlation is significant within
the 1 % level.

A further breakdown shows that the correlation, r, between price
and production on the Gulf Coast amounts to 0.737 which is even more
significant (Table 2). The Gulf correlation is higher than that of price

196

Gunter and McGraw

Table 2.

The Catch of United States Gulf Coast Brown and White Shrimp in
Thousands of Pounds and Thousands of Dollars

Year Pounds Value Year Pounds Value

1902

8,031

1908

8,156

1918

30,466

1923

30,595

1927

44,725

1928

53,357

1929

50,468

1930

40,203

1931

46,075

1932

42,427

1934

60,621

1936

54,723

1937

73,050

1938

73,108

1939

78,173

1940

83,012

1945

94,444

1950

98,359

1951

125,747

1952

128,745

199

1953

270

1954

1,276

1955

1,771

1956

2,344

1957

3,092

1958

2,986

1959

2,017

1960

1,817

1961

1,400

1962

2,278

1963

2,756

1964

4,181

1965

3,725

1966

3,991

1967

5,141

1968

17,305

1969

33,112

1970

44,136

48,170

1971

145,781

66,336

153,995

53,652

137,923

54,465

125,727

62,499

74,760

63,288

76,992

63,871

94,362

50,348

94,276

57,631

53,574

43,650

64,582

60,557

103,067

63,539

86,139

62,695

96,010

70,907

93,886

82,971

125,862

90,574

109,799

95,837

110,723

101,131

126,897

108,183

129,850

123,770

and production of total shrimp, of the South Atlantic and Gulf com-
bined.

In contrast, the correlation between price and total catch on the
South Atlantic Coast, Table 3, is 0.067, which is not significant at all.
This somewhat anomalous conclusion becomes clear if the shrimp of
the South Atlantic Coast were over-fished rather early in the develop-
ment of this fishery and have been over-fished for years. This explana-
tion was advanced by Mr. Milton J. Lindner, whose experience with
the South Atlantic shrimp fishery began in 1930. Examination of
Table 3 shows that high production in white shrimp on the Atlantic
Coast was attained in the 1920s. Apparently these shrimp were fished
to the very limit of their yield and have been for a great number of
years. This seems to be the only reasonable explanation of the fact
that price level and shrimp production have not increased together on

Shrimp Landing Statistics

197

Table 3.

The Catch and Values of White and Brown Shrimp in Thousands of
Pounds and Thousands of Dollars for the South Atlantic

Year

Pounds

Value

Year

Pounds

Value

1902

2,475

87

1953

21,385

9,931

1908

3,699

138

1954

18,601

6,883

1918

10,166

470

1955

18,531

6,939

1923

15,392

822

1956

16,570

7,806

1927

19,475

1,174

1957

15,604

9,150

1928

21,629

1,458

1958

12,911

7,958

1929

20,019

1,449

1959

14,186

6,527

1930

17,016

979

I960

17,812

8,512

1931

16,553

914

1961

10,660

6,939

1932

14,586

636

1962

13,206

11,275

1934

16,858

789

1963

9,468

5,246

1936

21,797

1,022

1964

9,674

6,633

1937

17,816

828

1965

15,633

10,160

1938

17,899

821

1966

13,155

10,813

1939

17,977

857

1967

11,975

9,010

1940

14,742

754

1968

14,681

13,996

1945

28,299

3,984

1969

15,608

16,186

1950

23,689

10,032

1970

12,541

11,386

1951

1952

18,033

16,669

7,382

6,585

1971

18,275

19,592

the South Atlantic Coast, but have increased together on the Gulf
Coast.

It may be further assumed that if the Gulf fishing continues at
a high level with a continued price rise, that the production of Gulf
shrimp will reach a limit, if it has not already done so, and that in
future times price and shrimp production on the Gulf Coast will no
longer show a correlation.

PRODUCTION FIGURES BY AREAS AND SPECIES

Because of previous correlations noted between the production
of white shrimp and rainfall in the State of Texas (Gunter and Ed-
wards 1969) and the apparent preference of brown shrimp for higher
salinities, we determined the correlations between the catch of whites

198

Gunter and McGraw

and browns in the State, r equaled — 0.2151, but with only 14 degrees
of freedom it was not significant.

Similarly there was no significant correlation between the catch
of browns and whites on the South Atlantic Coast, the Gulf Coast, or
the total of both areas. This means apparently that the production of
these two shrimp are not closely related to one another and that they
have different ecological niches and are weakly competitive, if at all.

On the other hand, there is a correlation between the total annual
production of shrimp of the South Atlantic; wdth the total annual pro-
duction in the Gulf, in which r equals 0,3261 with 37 degrees of free-
dom (Table 4). This is .significant at the level of 5.0%. This would
mean that when conditions are generally good for shrimp production
on the Gulf, they are also good on the Atlantic. Most likely these con-

Table 4.

Comparison of South Atlantic and Gulf Catches of White and Brown
Shrimp in Thousands of Pounds

Year

Atlantic

Gulf

Year

Atlantic

Gulf

1902

2,475

8,031

1953

21,355

145,781

1908

3,699

8,156

1954

18,601

153,995

1918

10,166

30,466

1955

18,531

137,923

1923

15,392

30,595

1956

16,570

125,727

1927

19,475

44,725

1957

15,604

74,760

1928

21,629

53,357

1958

12,911

76,992

1929

20,019

50,468

1959

14,186

94,362

1930

17,016

40,203

1960

17,812

94,276

1931

16,553

46,075

1961

10,660

53,574

1932

14,586

42,727

1962

13,206

64,582

1934

16,858

60,621

1963

9,468

103,067

1936

21,797

54,723

1964

9,674

86,139

1937

17,816

73,050

1965

15,633

96,010

1938

17,899

73,108

1966

13,155

93,886

1939

17,977

78,173

1967

11,975

125,862

1940

14,742

83,012

1968

14,681

109,799

1945

28,299

94,444

1969

15,608

110,723

1950

23,689

98,359

1970

12,541

126,897

1951

18,033

125,747

1971

18,275

129,850

1952

16,669

128,745

Shrimp Landing Statistics

199

ditions are of a broad climatic nature, involving such things as cool
and warm years, high rainfall and droughts, and even hard cold
waves. It would be quite difficult to get some of these factors into
figures or numbers, especially comparable figures for statistical calcu-
lations, even if the climatic events were recorded years ago as many
w'ere not. Therefore, we will pass this question by.

Similarly there is a very strong correlation between the white
shrimp production of the Atlantic Coast and Gulf Coast (Table 5).

Table 5.

Catch Figures for South Atlantic and Gulf White Shrimp in Thou-
sands of Pounds

Year Atlantic Gulf Year Atlantic Gulf

1902

2,475

1908

3,699

1918

10,166

1923

15,392

1927

19,475

1928

21,629

1929

20,019

1930

17,016

1931

16,553

1932

14,586

1934

16,858

1936

21,797

1937

17,816

1938

17,899

1939

17,977

1940

14,742

8,031

1945

8,156

1957

30,466

1958

30,595

1959

44,725

1960

53,357

1961

50,468

1962

40,203

1963

46,075

1964

42,727

1965

60,621

1966

54,723

1967

73,050

1968

73,108

1969

78,173

1970

83,012

1971

28,299

94,444

9,554

11,129

7,204

25,740

8,326

24,574

12,200

28,381

9,113

14,421

7,879

23,166

4,719

47,087

5,272

43,978

10,587

33,599

5,948

29,917

7,020

24,960

11,004

30,918

10,294

44,959

8,111

45 ,962

12,077

42,010

The total series stemming from 1902 to 1971 has 29 degrees of the
freedom, because the years 1948 to 1957 were excluded when brown
shrimp and white shrimp were not properly separated in the fisheries
statistics. The correlation r was found to be 0.655 and significant at
the 1% level.

In contrast, no such correlation can be shown between the brown
shrimp catch of the South Atlantic and Gulf (Table 6). We may spec-

200

Gunter and McGraw

Table 6.

Atlantic and Gulf Brown Shrimp Production in Thousands of Pounds

and the Totals

Year

Atlantic

Gulf

Atlantic
and Gulf
browns
combined

1957

6,050

63,631

69,681

1958

5,707

51,252

56,959

1959

5,860

69,788

75,648

1960

5,612

65,895

71,507

1961

1,547

39,153

40,700

1962

7,164

41,416

48,580

1963

4,749

55,980

60,729

1964

4,402

42,161

46,563

1965

5,046

62,411

67,457

1966

7,207

63,969

71,176

1967

4,955

100,902

105,857

1968

3,677

78,881

82,558

1969

5,314

65,764

71,078

1970

4,430

80,934

85,364

1971

6,060

87,788

93,848

ulate here that brown shrimp spend a shorter time in the bays, and
live in deeper water in the ocean, and for that reason would be less
aifected by climatic variations than the white shrimp in shallower
water. Thus production would be less subject to parallel variations
induced by climatic variables in shallow water, all leading to greater
correlations of the white shrimp catch on the two coasts.

We pursued this idea a little further and compared the coefficient
of variation of the brown and white shrimp catches (Table 7). The
coefficient of variation for the brown shrimp was 25.918 and for the
white shrimp was 28.569. A comparison of the significance of differ-
ences between two variants showed that this was significant at the
classical 95% level. This means that the brown shrimp production is
probably less variable than the white shrimp production on the United
States coast, and possibly a longer series of data will clarify this point.

A list of significant correlations determined in this study and a
list of correlations which are not statistically significant are given in
Tables 8 and 9, respectively.

Shrimp Landing Statistics

201

Table 7.

Total Brown and White Shrimp Catches of the United States in

Thousands of Pounds

Year

Browns

Whites

1957

69,681

20,683

1958

56,959

32,944

1959

75,648

32,900

1960

71,507

40,581

1961

40,700

23,534

1962

48,580

29,208

1963

60,729

51,806

1964

46,563

49,250

1965

67,457

44,186

1966

71,176

35,865

1967

105,857

31,980

1968

82,558

41,922

1969

71,078

55,253

1970

85,364

54,073

1971

93,849

54,087

Table 8.

A List of Significant Correlations Determined in this Study

Degrees of
Freedom

r

Signifi-

cance

1.

South Atlantic and
Gulf browns and
whites vs. values...

37

0.6912

1.0%

2.

Gulf browns and
whites vs. values...

37

0.7368

0.1%

3.

South Atlantic
browns and whites
vs. Gulf browns
and whites.

37

0.3261

5.0%

4.

Atlantic whites
vs. Gulf whites

... 32

0.6550

1.0%

202

Gunter and McGraw

Table 9.

A List of Correlations Determined in this Study that are not

Statistically Significant

Degrees of
Freedom

r

1.

South Atlantic grooved
vs . whites.

15

-0.003

2.

Gulf grooved vs. white
shrimp

16

0.094

3.

South Atlantic browns
vs . whites . .

15

-0.065

4.

Gulf browns vs. whites..-

16

0.051

5.

South Atlantic and

Gulf browns vs. South
Atlantic and Gulf
whites

13

0,2790

6.

Texas grooved vs.
whites

16

-0.148

7.

Texas browns vs.
whites

16

-0.215

8.

South Atlantic browns
and whites vs. value,...

39

0.0674

9.

South Atlantic browns
vs. Gulf browns

15

0.121

SUMMARY

There are five species of commercial penaeid shrimp extending
from Cape FTatteras, North Carolina to northern Mexico. One is lo-
calized in Biscayne Bay, Florida and one is only produced in low per-
centage (less than 2 %) of the total catch in the Gulf of Mexico. A
third species, the pink shrimp, has had foreign catches so mixed with
the domestic production that local figures on the Gulf Coast for past
years are not reliable. Fairly adequate production figures for white
shrimp are available for the years 1902 to 1947 and 1958 to the pres-
ent, From 1948 to 1967 the brown and white shrimp catches were
mixed and to some extent with the pinks. After 1958 all species were
separated in the catch records.

Shrimp Landing Statistics

203

There is a strong positive correlation between total shrimp pro-
duction of the United States and value (dockside price) of the shrimp,
and an even more significant correlation between Gulf production and
value. In contrast the much smaller South Atlantic shrimp catch shows
no correlation with prices, probably because the stock has been fished
to capacity since the 1920s, when production limits seem to have been
obtained.

A strong correlation exists between white shrimp production of
the South Atlantic and the Gulf, while none was found for the brown
shrimp production of the two areas. A possible explanation for this
fact is the deeper w’ater distribution of the brown shrimp, which
means a more stable environment, less affected by general climatic
oscillations which influence white shrimp in shallow waters and cause
similar variations in the two populations.

There is no significant correlation between the total United States
production of white and brown shrimp, either positively or negatively,
nor are there any correlations of the South Atlantic and Gulf areas
considered separately. This means that the brown and white shrimp
are weakly competitive, if at all.

We wish to thank Mr. Paul Poole, data processor at the Gulf
Coast Research Laboratory, for his assistance • in the statistical
analyses.

LITERATURE CITED

Eldred, Bonnie, i960. A note on the occurrence of the shrimp, Penaeus brasiliensis
Latreille, in Biscayne Bay, Florida. Quarterly .Tournal of the Florida Acad-
emy of Sciences 23(2) : 164-65.

Gunter, Gordon. 1950. Seasonal population changes and distributions as related to
salinity of certain invertebrates of the Texas coast, including the commercial
shrimp. Pub. Inst. Mar. Sci. 1(2) t7-51.

1961. Some relations of estuarine organisms to salinity. Limnol. Ocean-

og. 6(2) : 182-90.

1962. Shrimp landings and production of the State of Texas for the

period 1956-1959, with a comparison of other Gulf states. Publ. Inst. Mar.
Sci. 8:216-26.

, J. Y. Christmas, and R. Killebrew. 1964. Some relations of salinity to

population distributions of motile estuarine organisms with special reference
to penaeid shrimp. Ecology 45(1) : 181-85.

, and .Judith C. Edwards, 1969. The relation of rainfall and freshwater

drainage to the production of the penaeid shrimps (Penaeus fiuviatilis Say
and Penaeus aztecus Ives) in Texas and Louisiana waters. FAO Fish. Rep.
57(3) :875-92.

, and Gordon E. Hall. 1963. Biological investigations of the St. Lucie es-
tuary (Florida) in connection with Lake Okeechobee discharges through the
St. Lucie canal. Gulf Res. Rep. 1(5) :189-307,

Lindner, Milton J. 1957. Survey of shrimp fisheries of Central and South America.
U. S. Fish Wildl. Scrv. Sp. Sci. Rep. Fish. 235 :1-166.

, and William W. Anderson, 1956. Growth, migrations, spawning, and

204

Gunter and McGraw

size distributions of the shrimp Penaeus setiferus. U. S. Fish Wildl. Serv.,
Fish. Bull. 106:555-646.

Lyles, Charles H. 1969. Fishery statistics of the United States: 1967. U. S. Gov-
ernment Printing Office. Washing^ton, D. C.

Say, Thomas. 1817. An account of the Crustacea of the United States (continued).

Journal of the Academy of Natural Sciences 1(6) :235-353.

Weymouth, F. W., M. J. Lindner, and W. W. Anderson. 1933. Preliminary report
on the life history of the common shrimp, Penaeus setiferus (Linnaeus).
U. S. Dept. Commerce, Bur. Fish. Bull. 14:1-26.

Gulf Research Reports

Volume 4 Issue 2

January 1973

Effect of Holothurin on Sarcoma 1 80 and B- 1 6 Melanoma Tumors in Mice

S.D. Cairns

Louisiana State University

C.A. Olmstead

Louisiana State University

DOI: 10.18785/grr.0402.07

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Biology Commons, and the Marine Biology Commons

Recommended Citation

Cairns, S. and C. Olmstead. 1973. Effect of Holothurin on Sarcoma 180 and B-16 Melanoma Tumors in Mice. Gulf Research Reports
4 (2): 205-213.

Retrieved from http://aquila.usm.edu/gcr/vol4/iss2/7

This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean
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EFFECT OF HOLOTHURIN ON SARCOMA 180 AND 8-16
MELANOMA TUMORS IN MICE^

by

S. D. CAIRNS 2 and C. A. OLMSTED
Department of Biological Sciences, Louisiana State
University in New Orleans, Louisiana 70122

ABSTRACT

Holothurin, a biotoxic principle from the Cuvierian glands of the
Bahamian sea-cucumber, Actinopyga agassizi, was studied as an anti-
tumor agent capable of retarding tumor grow’th and prolonging the
life of tumor-bearing mice. White Swdss mice injected with Sarcoma
180 had a mean survival time of 18,1 days wdth an average weight
gain representing tumor and ascites fluid accumulation amounting to
25.1 grams in 15 days. Of five white Swiss mice with Sarcoma 180 re-
ceiving 0.15 mg Holothurin every other day, one was alive at 57 days
and the average weight gain of the group was 10.4 grams in 15 days.
White Swiss mice with Sarcoma 180 which survived the lethal effects
of slightly higher doses of Holothurin also had prolonged survival
times and negligible tumor growth. C57-B1-6J mice with B-16 mela-
noma tumors did not show increased survival time using the same
doses of Holothurin that were effective in w'hite Swiss mice. Toxicity
tests indicated that the safe upper limit for intraperitoneal Holothurin
injection in white Swiss mice was 0.10 mg/day, 0.15 mg given every
other day, and up to 0.30 mg given in a single injection. Similar tests
with C57 black mice showed survival with as much as 0.60 mg Holo-
thurin in a single subcutaneous injection. Holothurin was found to be
250 to 500 times more effective in causing red blood cell hemolysis
than saponin and thus appears to have some action on living cells in
addition to its surfactant action.

INTRODUCTION

Cancer chemotherapy began in 1946 when nitrogen mustai’d was
used in treating leukemia patients. During the next 15 years only some
thirty drugs were used in cancer chemotherapy but screening of many
thousands of compounds was carried out each year (Clark 1961), Al-
though marine invertebrates provide a particularly rich source of
compound.^ with biological activity in mammalian species, less than

^ This work was supported in part by the Cancer Association of Greater New
Orleans, Inc.

2 Present address: Department of Biological Oceanography, the Rosenstiel
School of Marine and Atmospheric Sciences, University of Miami, Miami, Florida
33129 .

205

206

Cairns and Olmsted

1% of all marine invertebrates thought to possess biotoxic substances
have been investigated and of those, only a dozen have been thorough-
ly investigated pharmacologically (Halstead 1965).

Small concentrations of a crude water extract made from the
whole body of the Bahamian sea-cucumber, Actinopyga agassizi, was
found to be lethal to mice and fish and to have some tumor cell inhibi-
tory action in vitro (Nigrelli and Zahl 1952). The active principle of
this crude extract was found to be concentrated in the Cuvierian tub-
ules located in the sea-cucumber respiratory tree and was named Holo-
thurin (Chanley et al. 1955). Holothurin w’as the first known steroid
saponin of animal origin (Nigrelli et al. 1959). Chemical analysis of
Holothurin indicates that it is highly soluble in water, non-volatile,
heat stable, and exhibits surface-active properties (Nigrelli and Ja-
kowska 1960). Holothurin appears to consist of a few steroid agly-
cones that are bound individually to four monosaccharide molecules.
A provisional formula has been proposed (Alender and Russell 1966).
See Fig. 1.

Figure 1. Holothurin formula. Proposed structure for Holothurin mole-
cule, a steroid saponin of animal origin; the monosaccharide groups rep-
resented by Si-S. are, respectively, D-glucose, D-xylose, D-quinovose, and
3-o-methyl glucose.

Holothurin Effects on Sarcoma

207

Holothurin is highly toxic to many types of organisms in very
small concentrations. It retards onion root tip development (Nigrelli
and Jakowska 1960) and fruit fly pupation (Goldsmith* Osburg and
Nigrelli 1958) at 1000 ppm, alters regeneration of planarians at 100
ppm (Qiiaglio et al. 1957), is lethal to the “pearr’ fish Campus at
1 ppm and affects sea urchin development at 0.01 ppm (Ruggieri and
Nigrelli 1960). Holothurin exhibits a hemolytic effect on red blood
cells and reduces the size of a subcutaneously injected Sarcoma 180
tumor in white Swiss mice (Nigrelli 1952). Krebs-2 ascites tumors in
white Swiss mice are also inhibited by Holothurin (Sullivan, Ladue
and Nigrelli 1955).

The present study demonstrates inhibition of Sarcoma 180 tu-
mors in white Swiss mice but not of B-16 melanoma tumors in C57
Black 6J mice. Toxicity of Holothurin was estimated by measuring its
effects on red blood cell hemolysis and by lethal-dose measurements in
mice. It appears that the anti-tumor activity of Holothurin is not only
dosc-dcpcndcnt but that the effective dose is determined by the sensi-
tivity of the species to the lethal effects of Holothurin.

MATERIALS AND METHODS

Holothurin : The Holothurin referred to in this paper was a
“crude extract” obtained from Dr. Ross Nigrelli and was made from
the sun-dried, powdered Cuvierian glands of Bahamian sea-cucum-
bers, Actinopyoa agnssizi.

Red Blood Cell Hemolysis Tests Using Holothurin and Saponin :
Fresh red blood cells w^ere obtained from white Swiss mice just prior
to testing the hemolytic effects of Holothurin and saponin of plant
origin. The blood was obtained by cardiac puncture while the mice
were under light ether anesthesia. Approximately 5 ml of blood was
diluted in 100 ml of balanced-saline solution buffered at pH 7.4 (Olm-
sted 1967) containing calcium o.xalate (0.1% ) to prevent coagulation.
A serial dilution of Holothurin wa.s prepared ranging from 2 mg ml
to 0.001 mg/ml and a similar series of dilutions of saponin was made
for comparison.

One ml of diluted blood was added to 5 ml of each of several con-
centrations of both Holothurin and saponin. The test tube was centri-
fuged 1 minute after mixing the contents and the optical density of
the supernatant was measured using a spectrophotometer set at 540
mu. The degree of hemolysi.s was estimated by the concentration of
hemoglobin in the supernatant. These values were compared to 100%
hemolysis values obtained either by using water only as the diluent or
by using a high concentration of Holothurin or saponin.

Holothurin Toxicity in Mice: All Holothurin injections w^ere
given at dosages determined by previous tests done in this laboratory

208

Cairns and Olmsted

on normal, non-tumor-bearing mice. Tests to determine the lethal and
sub-lethal intraperitoneal doses of Holothurin in white Swiss mice
were carried out using several concentrations given daily, given on
alternate days, and for several concentrations given in a single injec-
tion. The dosages of Holothurin given to the white Swiss mice bearing
Sarcoma 180 tumors were based on these data. The dosages of Holo-
thurin given to the C67 black mice bearing B-16 melanoma tumors
was based on the effective doses of Holothurin given to tumor-bearing
white Swiss mice. In addition, a test of toxicity of a single subcuta-
neous injection of Holothurin was carried out using several concentra-
tions of Holothurin given to normal, non-tumor-bearing C57-B1-6J
mice.

Holotkm'in Administrotio^i to Mice Bearing Sarcoma ISO: Sar-
coma 180 cells and the white Swiss mice were obtained from the Gulf
South Research Institute in New Orleans. The mice were inoculated
intraperitoneally with 0.5 ml of Sarcoma 180 ascites fluid containing
11.6 X 10^ cells/ml. Groups of five mice received daily injections of
Holothurin at 0.01, 0.05, 0.10, and 0.15 mg each, and three groups re-
ceived injections of Holothurin every other day in the amounts of
0.10, 0.15 and 0.20 mg each. Twelve mice received Sarcoma 180 inocu-
lations, but no Holothurin. The survival time was recorded for each
mouse and the mice were weighed individually before inoculation with
tumor cells and at 5-day intervals during the course of the experi-
ments,

Holothurin Admmistration to Mice Bearing B~16 Melanoma: A
single C57-B1-6J mouse with a B-16 melanoma tumor was used as the
source for all of the B-16 melanoma used in this study. The tumor-
bearing donor mouse was obtained from the Department of Surgery
at Tulane Medical School. The C57-B1-6J mice used in the experi-
ments here were obtained from the Roscoe B. Jackson Memorial Lab-
oratory in Bar Harbor, Maine. The solid tumor was taken from the
donor mouse under light ether anesthesia and was minced with scis-
sors in a tissue culture nutrient medium containing serum (Olmsted
1967). Several .small pieces of the minced tumor were trochared sub-
cutaneously into the left .side of thirty-five recipient male mice. Thirty
of these mice were used for the experiments reported here. Daily in-
jections of 0.05, 0.10, and 0.15 mg Holothurin were given to each of
the five tumor-bearing mice in three groups and five tumor-bearing
mice in another group were each given 0.20 mg Holothurin every
other day. Five mice without melanoma, and ten mice with melanoma
tumors, were also maintained but not given Holothurin. All Holo-
thurin injections were given subcutaneously near the region of tumor
implantation in 0.10 ml volumes, and were given over a 13-day period.
The survival time was recorded for each mouse, and the mice were
weighed in groups of five before tumor implantation and at 6-day in-
tervals during the course of the experiments.

Holothurin Effects on Sarcoma

209

RESULTS

Hemolysis Tests: Minimum concentrations of Holothurin and
saponin that produced complete hemolysis were found to be 0.001
mg/ml of Holothurin and 0.25-0.50 mg/ml of saponin indicating that
Holothurin was 250 to 500 times more active as a hemolytic agent
than saponin.

Holothurin Toxicity in Mice: Holothurin treatment of tumor-
bearing white Swiss mice at the level of 0,15 mg/day was lethal at 3
to 8 days with a mean of 4.2 days. See Table 1. Toxicity tests on other

Table 1.

Effects of Holothurin on White Swiss Mice with Sarcoma 180

Body

Weight Change

No.

of

Mice

Sex

Dose

(mg)

Frequency

of

iniection

Survival

Time

(days)

5th

day

(8)

10th

day

(e)

15th

day

(e)

12

F

0

0

18.1

(15-21)

+2.3

+13.0

+25.1

5

F

0.01

daily

19.2

(15-22)

+2.0

+12.7

+27.6

5

F

0.05

daily

17.8

(16-23)

0

+13.8

+33.0

5

F

0.10

daily

16.3

(14-19)

-1.4

+ 8.2

+30.7

5

F

0.10

alternate

days

16.8

(15-20)

-1.1

+10.1

+29.4

5

F

0.15

alternate

days

26.4

(16-58)

-1.0

+ 2.9

+10.4

5

F

0.15

dally

4.2

(3-8)

died

■“ —

—

5

M

0.20

alternate

days

22.2

(10-47)

-1.8

0

0

groups of normal, non-tumor-bearing white Swiss mice indicated that
either 0.10 mg/day or 0.15 mg given on alternate days was a safe
upper limit for intraperitoneal Holothurin administration. Addition-
ally, 0.30 mg Holothurin in a single injection could be tolerated, but
more than this amount in a single injection was lethal. Toxicity tests

210

Cairns and Olmsted

OB C57-B1-6J mice without B-16 melanoma tumors indicated that
these mice could survive single subcutaneous injections of Holothurin
as high as 0.60 mg.

Effect of Holothurm o?i White Swiss Mice with Sarcoma 180:
Twelve tumor-bearing mice that were not treated with Holothurin
lived for an average of 18.1 days after tumor cell inoculation. These
mice gained an average of 25.1 g by the fifteenth day after tumor cell
inoculation. This increase in body weight represents the extent of in-
crease in tumor size and ascites fluid accumulation since body weight
increase due to growth of the animal during this time would have
been negligible. The tumor-bearing animals receiving 0,15 mg Holo-
thurin on alternate days had a mean survival time of 26.4 days repre-
senting an increase in survival time of 46%, and a mean w'eight gain
at 15 days of 10.4 g representing a 60 to 70% decrease in tumor
growth and ascites fluid accumulation. One of the mice in this high
Holothurin dose group had a weight gain of only 5.1 g and was still
alive after 58 days. Three of the mice receiving 0,20 mg Holothurin
every other day died in less than 12 days probably as a result of Holo-
thurin overdose. Of the other two mice in the 0.20 mg group, one
lived for 30 days and the other for more than 47 days and both had
negligible weight gain during this period of time indicating complete
tumor suppression. The four groups of tumor-bearing mice receiving
less than 0.15 mg of Holothurin per injection did not show any sig-
nificant increase in survival time or decrease in weight gain. See
Table 1,

Effects of Holothurin on C57~Bl-6J Mice with B-16 Melanoma:
Ten C57-B1-6J mice not injected with Holothurin lived for an average
time of 29.4 days after B-16 melanoma inoculation with a range of
18 to 43 days. The animals with malignant melanoma tumors de-
creased in total body weight. Normal, non-tumor-bearing C57-B1-6J
mice of comparable age to the tumor-bearing mice all gained w'eight
on the diet and the conditions' of this experiment. Measurement of
body weight w'as not a valid indicator of tumor growth or of Holo-
thurin effectiveness, as in the case of mice with Sarcoma 180, because
the melanoma tumor grows in a small solid mass and does not induce
ascites fluid accumulation. As the Holothurin dosages used here were
comparable to the effective doses for w^hite Swiss mice bearing Sar-
coma 180 tumors^ and were well below the toxic levels of Holothurin
for C57-B1-6J mice, none of the animals in this experiment showed
any increase in the survival time. See Table 2.

DISCUSSION

The crude extract of Holothurin used here was probably more
potent than purified preparations would be. Upon purification, the
anti-tumor principle is apparently lost or considerably reduced in con-

Holothurin Effects on Sarcoma

211

Table 2.

Effects of Holothurin on C57-B1-6J Mice with Melanoma

Body

Weight Change

No.

of

Mice

, Sex

1

Dose

(mg)

Frequency

of

injection

Survival
Time
: (days)

5 th
day

(fi)

! 10th
day

(8)

m

I 20 th
day
(8)

10

M

0

0

29.4

(18-43)

+0.4

-0.7

-0.8

-0.3

5

M

0.05

daily

25.6

(20-42)

-1.1

-3.4

-3.5

-2.5

5

M

0.10

daily

22.7

(19-26)

-4.1

-4.8

-5.4

-4.4

5

M

0.15

daily

1

i

24.4

(20-35)

-3.1

-3.5

!

-4.3

-4.5

5

M

0.20

alternate

days

29.6

(25-33)

-2.6

-3.5

-3.6

-3.3

centration (Chanley et al, 1960). One sample of purified Holothurin
showed a tenfold-loss in hemolytic activity, and no anti-tumor activ-
ity (Nigrelli and Jakowska 1960).

Although the crude extract of Holothurin is effective against var-
ious tumors (Nigrelli 1952, and Sullivan, Ladue and Nigrelli 1955),
the present study demonstrates that Holothurin is apparently not ef-
fective at the same dose levels in differing species of mice. Larionov
(1967) suggests that a natural biological product should be more prom-
ising as a chemotherapeutic agent than a synthetic compound. The
same author has stated that the maximum effective acceptable dosage
of an anti-tumor compound is close to the dosage which kills the
animal, whereas a decrease in dosage, even slight, leads to a sharp
drop in effectiveness. The narrow thre.shold is seen here in regard to
the ineffectiveness of the lower concentrations of Holothurin on the
Sarcoma 180 tumors, and the lethal effect of 0.20 mg given over a
period of several alternate days to the tumor-bearing mice. Due to the
narrow range of effective concentrations of Holothurin in relation to
its lethal dose, it is likely that the ineffectiveness of Holothurin in the
C57-B1-6J mice bearing B-16 melanomas was a reflection of the toler-
ance to higher doses of Holothurin in these animals. There is no in-
dication that Holothurin, or any other chemotherapeutic agent, would
have differing effects in male and female tumor-bearing mice although
profound metabolic differences between male and female rats and
mice are well-known (Olmsted 1969). However, the differences seen

212

Cairns and Olmsted

in this study could also be due to differences in the injection site or in
the types of tumor. While this consideration presents many variables,
the Sarcoma 180 tumor grows best in white Swiss mice and the B-16
melanoma grows best in C57-B1-6J mice. Further, it should be noted
that the mean survival time of the untreated mice with the two dif-
ferent types of tumor was about the same, i.e., about S to 4 weeks
after tumor inoculation. The hypothesis is further supported by the
finding that Holothurin is many times more effective in causing he-
molysis than saponin when living red blood cells are used in the test
system.

Other hemolysis experiments similar to those described here have
shown that between 0.04 and 0.10 mg of the crude extract of Holo-
thurin would cause complete hemolysis, while 0.08 to 0,10 mg of sa-
ponin was required for the same effect (Nigrelli and Jakowska 1960) .
The apparent 40 to 100-fold discrepancy between the findings of Nig-
relli and Jakowska and the findings of the present study could be ac-
counted for by the use of different dilutions of red blood cells, or by
differences in the age of red blood cells used, or by species effects.
Nigrelli and Jakowska's data suggest that Holothurin is only slightly
more effective in causing hemolysis than saponin, while in the present
study using red blood cells that were freshly obtained from mice just
prior to testing it would appear that Holothurin is some 250 to 500
times more effective than saponin in causing hemolysis. Since a num-
ber of studies have shown Holothurin to have profound biological ef-
fects in concentrations as low as 0.01 ppm it is not surprising that
this biotoxic principle is much more effective than saponin on fresh
red blood cells. It would therefore appear that the action of Holo-
thurin involves more than a surface tension lowering effect on living
cells.

Because Holothurin is an effective anti-tumor agent at concen-
trations only slightly lower than the lethal dose, it might be tenta-
tively hypothesized that Holothurin is acting on some process common
to all cells, not just tumor cells. However, no studies have been done
on the metabolic effects of Holothurin. The studies reported here indi-
cate that the mode of action of Holothurin in retarding tumor growth
and prolonging life of tumor-bearing mice is probably different from
the action of a surfactant such as saponin. It is likely that the energy
requirements of the white Swiss mice with Sarcoma 180 differed from
those of the C67-B1-6J mice bearing B-16 melanoma tumors. Other
studies from this laboratory on the C67-B1-6J mice with B-16 mela-
noma tumors have shown that the growth of melanoma tumor occurs
at the expense of the anabolic processes of the host mouse. Structural
phospholipids synthesized by the tumor were shown to be derived
from synthesis in the liver (Terranova, Hardy and Olmsted 1970).
Although several experiments have been reported in the literature
regarding .saponin, none have shown saponin to have any anti-tumor
activity in vivo. It is likly that saponin destroys cells simply by its

Holothurin Effects on Sarcoma

213

action to lower surface tension starting with the plasma membrane
and continuing its action on membranes of other cellular constituents
in proportion to the concentration of saponin used. While the action
of Holothurin has some yet unknown eilect other than, or in addition
to, its surfactant effect, it is likely that it causes an inhibition of an
active, or energy-dependent, physiological process.

LITERATURE CITED

Alender, C- B. and F. E, Russell. 1966. In Physiology of Echinoderms, R. A.

Boolootian (ed.), p. 531. Interscience, New York.

Chanley, J. D., S. K. Kohn, K. Nigrclli and H. Sobotka. 1955. Zoologica, 40:99.
Chanley, J. D., R. T.edeen, J, Wax, R. Nigrelli and H. Sobotka. 1959. J. Am. Chem.
Soc., 81:5180-3.

Chanley, J. D., J. Perlstein, R. Nigrelli and H. Sobotka. 1960. Ann. N. Y. Acad.
Sci,. 90:902-5.

Friess, S. L., F. G. Standaert, B. R. Whitcomb, R. Nigrelli, J. D. Chanley, and
II. Sobotka. 1959. J. Pharmacol., 126:323-9.

Clarke, R. L. 1961. Cancer Chemotherapy, Chas. C, Thomas, Springfield.
Goldsmith, B. D., H. B. Osburg and R. Nigrelli. 1958. Anat. Rec., 130:411-12.
Halstead, B. W. 1965. Poisonous and Venomous Marine Animals of the World,
U. S. Government Printing Office, Washington, D. C., 1 :994.

Larionov, L. F. 1967. Recent Results in Cancer Research, Vol. 8, P, Rentchnick
(ed.), Springer, New York.

Nigrelli, R. 1952. Zoologica, 37:89-90.

Nigrelli, R. and P. A. Zahl. 1952. Proc. Soc. Exp. Biol. Med.. 81:379-80.

Nigrelli, R., J. D. Chanley, S. K. Kohn and H. Sobotka. 1955. Zoologica, 40:47-8.
Nigrelli, R. and S. Jakowska. 1960. Ann. N. Y. Acad. Sci., 90:884-91.

Olmsted, C. A. 1967. J. Cell Biol., 48:283-99
Olmsted, C, A. 1969. Lipids., 4:401-12.

Quaglio, N. D„ S. F. Ndan, A. M. Veltri, P. M. Murray, S. Jakowska and R.

Nigrelli. 1957. Anat. Rec., 128:604-5.

Ruggieri, G. D. and R. Nigrelli, 1960. Zoologica, 46:1-16.

Sullivan, T. D., K. T. Ladue and R. Nigrelli. 1955. Zoologica, 40:49—52.
Terranova, J., K. Hardy and C. A. Olmsted. 1970. J. Am. Oil Chem. Soc., 47; 78 A.

Gulf Research Reports

Volume 4 Issue 2

January 1973

A Geochemical Study of a Marsh Environment

Thomas R Lytle

Gulf Coast Research Laboratory

Julia Sever Lytle

Louisiana State University

Patrick L. Parker

University of Texas

DOI: 10.18785/grr.0402.08

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Geochemistry Commons, and the Marine Biology Commons

Recommended Citation

Lytle; T. R, J. S. Lytle and R L. Parker. 1973. A Geochemical Study of a Marsh Environment. Gulf Research Reports 4 (2); 214-232.
Retrieved from http:/ / aquila.usm.edu/gcr/vol4/iss2/ 8

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A GEOCHEMICAL STUDY OF A MARSH ENVIRONMENT

by

THOMAS F. LYTLE
Gulf Coast Research Laboratory
Ocean Springs, Mississippi

JULIA SEVER LYTLE
Department of Chemistry
Louisiana State University
Baton Rouge, Louisiana

and

PATRICK L. PARKER
Marine Science Institute
The University of Texas
Port Aransas, Texas

INTRODUCTION

The study of the fate and distribution of carbon compounds and
their associated trace elements in contemporary environments as well
as ancient sediments has added information of importance to geology
and the ocean sciences. These recent advances have been spurred on
by such interests as the search for deposits of fossil fuels, the control
of pollution, the understanding of the origins of life, the analysis of
lunar samples, and the hopes for analyses of the surface of Mars.

The organic matter found in Holocene sediments is derived from
once living organisms. The complex molecules synthesized by orga-
nisms and their degradation products are the object of study of organ-
ic geochemistry. After the death of an organism^ most of the organic
matter in it is utilized by other organisms or oxidized to CO2. A small
amount of the organic matter is not destroyed but is trapped in the
top sediment. Biological activity gradually ceases, but slow chemical
and physical transformations are continuous. Since the carbon-carbon
bond is a strong bond, portions of the original molecules deposited in
sediments often survive for long periods of time. Detection of many
organic molecules in the concentrations found in nature was impos-
sible before recent improvements in instrumental methods of separa-
tion, characterization and quantitation. With the present tools of
mass spectrometry and gas chromatography the field of organic geo-
chemistry has undergone a tremendous surge of activity.

The trace metal components of sediments provide a rich area for
the study of geochemistry with such a diverse array of mechanisms
responsible for their inclusion in the sedimentary environment. The
biogenic source of trace metals in sediments is usually assumed but

214

Geochemical Marsh Study

215

rarely stimulates a great deal of interest. However, the biogenic com-
ponent may play a significant role in trace metal fixation in highly
productive areas. Because trace metals in a sediment may undergo
radical changes in concentration levels with only slight changes in the
overall chemistry of the sediment, they should be very sensitive indi-
cators of certain microscale and macroscale chemical and biological
factors in sedimentary environments. All organisms rely to a greater
or lesser extent upon a constant supply of trace metals for the func-
tioning of their metabolic processes. So profound is the need for cer-
tain trace metals in all organisms that they are termed essential trace
metals whereas others may form an essential ingredient in the nutri-
ents of only a few organisms. Others may perform no known func-
tion or be toxic to most organisms. Even so, plants and animals have
the capacity to concentrate most trace metals from their environ-
mental sources. When organisms die, most of the trace metals are re-
leased after decay, but a fraction may be retained in the organic
debris trapped in the sediments and be preserved as a remnant of the
pre-existing life forms. To study the trace metal transfer between
sediments and plants, analytical tools of high sensitivity are required
to cope with trace concentrations. Fortunately methods and instru-
ments — particularly atomic absorption spectrophotometry — having
the required sensitivity are available at modest costs to those inter-
ested in conducting trace metal analysis and have greatly facilitated
this type of study.

Harbor Island, located approximately 1 mile north of Port Aran-
sas, Texas (Fig. 1), was chosen as an environment to investigate in
an organic and trace metal geochemical study. Situated at the head of
Aransas Pass Channel, this tidal delta is characterized by a series of
salt marshes inundated by tidal creeks and tidal channels. Maximum
relief on the island is but 12 feet, but even slight variations in eleva-
tion are denoted by the variation of plant species dominating each
zone of elevation. The plants are limited to the more hardy herbaceous
species which withstand the extended periods of low rainfall and sedi-
ment dessication found in the intertidal and supratidal areas. Most of
the identified species of plants are also native to the salt marshes of
the off-shore islands of Mississippi. The fauna was restricted primar-
ily to members of the lower phyla of the animal kingdom.

Several factors made this island ideally suited for geochemical
investigations. The topography reveals rather simple and small scale
physiographic processes. The island is protected both from effects of
major oceanic disturbances by the barrier islands and from the effects
of man by its relative inaccessibility. The minimal number of external
parameters was helpful in facilitating the interpretation of the geo-
chemical data collected.

The goal of this study was twofold: 1) chemically to charac-
terize a specific salt marsh, the sediments and associated biota and 2)

216

Lytle, Lytle and Parker

Figure 1, Map of collecting area.

Geochemical Marsh Study

217

to establish clear relationships between the chemical substances re-
siding in the sediments and similar or identical substances occurring
in the biological specimens.

In this particular study it was felt that the hydrocarbons would
yield the most significant organic geochemical information. These
compounds are ubiquitous but minor components of all organisms.
Though their function is not entirely understood, it is known that they
are concentrated in the w^axy coatings of plants and mo.st likely aid in
the protective mechanisms of plants. Among the several classes of bio-
chemical materials, the hydrocarbons exhibit probably the greatest
resistance to biological and chemical degradation and therefore may
be preferentially concentrated and preserved in sedimentary environ-
ments. An extreme variet^^ of specific hydrocarbons occurs naturally
in plants and hence the possible combinations and distributions of
these hydrocarbons are limitless. The stability and unique distribu-
tions of hydrocarbons ranks them as a very important tool in the cor-
relation of biolipids and geolipids.

In choosing the trace metals to be studied, preference was given
to those that are known to be essential to all organisms. Those ele-
ments fitting this description were copper, zinc, molybdenum, man-
ganese and iron. In addition cobalt known to be essential at least to
blue-green algae was included. To monitor the biogenic contribution
of non-essential trace metals, nickel, cadmium and lead w'ere also
analyzed in the samples. With the diverse functions of these various
elements and occurrence at easily measurable levels in most organ-
isms, they should provide useful information about the plant-sediment
interplay in a salt marsh.

EXPERIMENTAL

CoUevlion and Handling

A shallow depression outlined by a band of Batis maritiina (Salt-
wort) and Salicomia bigelovii (Glass wort) was chosen as the site for
sediment and plant collection for organic analysis. The sediments
were black indicating a reducing environment and contained root mot-
tles and plant debris. Algal mats were forming at the edges of the de-
pression and benthic organisms could be seen as they fed on ditritus
present at the surface. Sediment samples chosen for trace metal analy-
sis were surface sediments from three intertidal zones of the island.
In addition one subtidal, subsurface .sample was taken from the lit-
toral zone of the island. This latter sample was taken to define the
trace element distribution in an area not influenced by the marsh
plants.

For the organic analysis, seven ‘marsh plants were collected,
washed with distilled water and extracted immediately after collec-

218

Lytle, Lytle and Parker

tion. After washing the whole plants with distilled water to remove
loose debris and epiphytes, the plants were minced in a Waring blen-
der with methanol. The material was filtered and the methanol saved.
The minced material was extracted ultrasonically for 15 minutes with
200 ml of chloroform with constant stirring. In addition to these seven
plants, seven more species were collected for trace metal analysis
(Table 1). Only herbaceous portions of the plants were saved. These

Table 1.

Marsh Plants Analyzed

HYDROCARBONS AND TRACE METALS

Limoniwn aavoli-nianim (Walt.) Britton (Sea lavender)
Batis mca^ltima L. (Saltwort)

Salioomia higelovii Torr. (Glasswort)

Lyoiym carolinianum Walt. (Christmas berry)

Sesuvium mapitimm. (Walt.) BSP (Sea Purslane)
Oenothera drummondii Hooker (Evening primrose)
Bovpiohia frutesoens (L.) DC. (Sea ox-eyes)

TRACE METALS

Spartina altevni flora Loisel (Cord grass)
Machaeranthera phyllooephala (DC.) Shinners
Spartina patens (Ait.) Muhl, (Salt-grass)
Monanthochloe littoralie Engelm. (Key-grass)
Hedyotis nigidoans (Lam. ) Fosberg
Avioennia germinans (L.) (Black mangrove)
Distiohlis spioata (L.) Greene (Marsh spike-grass)

portions were quickly rinsed in double distilled water and oven dried
at 60°C for 48 hrs. Following ashing in a muffle furnace at 450'’C
for 24 hrs., the samples were leached with 3N HCl and the filtrate
over glass fiber filters was saved.

Sediment samples for all analyses were hand collected, and all
necessary precautions were taken to eliminate contaminations and to
arrest bacterial action. The sediment samples for hydrocarbon analy-
sis were digested with dilute HCl to remove carbonates and w’ashed
to remove inorganic salts. Sediments for trace metal analysis were

Geochemical Marsh Study

219

oven dried at 60 °C, weighed, ashed at 450°C digested in 3N HCl, fil-
tered on glass wool and the filtrate saved.

The extracts from plants and sediments were further separated
as follows :

Hydrocarbon Separation and Characterization

Each lipid sample (plant and sediment) was saponified by reflux-
ing with 0.5N KOH-MeOH for 1 hour. Nonsaponifiable components
were removed by extracting the alkaline solution with benzene. After-
wards, the alkaline solution was acidified with dilute HCl to pH 3,
and the fatty acids were extracted into benzene. Methyl esters of the
fatty acids were prepared using BF.,-MeOH (Metcalfe and Schmitz
1961). Silica gel (Woelm, Grade 200, Act. I) was packed beneath 25
ml alumina (Woelm Neutral, Grade 100, Act. I) in a 43 cm X 2.5 cm
(o.d.) column. The nonsaponifiable residue was fractionated on the
column into four parts : n-hexane fraction contained aliphatic hydro-
carbons; benzene fraction contained aromatic hydrocarbons; chloro-
form-methanol (4:1, v/v) fraction contained alcohols; methanol frac-
tion contained glycerides and the polar lipids.

The aliphatic hydrocarbons were identified and measured by gas
chromatography on columns of SE-30, Apiezon L, and FFAP (Va-
rian). Standard hydrocarbons were used to calibrate the Perkin-
Elmer 880 gas chromatograph equipped with hydrogen flame ioniza-
tion detectors. The columns were 8 feet by 1/8 inch o.d. copper tubing.
The support was 80/100 mesh Chromosorb G, acid washed, dichloro-
di methyl silane treated (Johns Manville). Fatty acid methyl esters
were identified using the same in.strument but using columns with
FFAP, SE-30, and diethylene glycol succinate (DEGS). Linear-log
plots of the retention times yielded straight lines for both hydrocar-
bons and acids and were useful for identification when standards for
each carbon number were unavailable. Coinjection of standards were
used to clarify some identifications. All hydrocarbon samples were run
at programmed temperatures from 100° to 260° at 6®C per minute
holding at 265°C. All fatty acid methyl esters were run from 150° to
265'' at 6°C per minute holding at 265°C. Mass spectra were obtained
with a modified Consolidated Electrodynamics Corp., Model 21-1 03C
mass spectrometer. The spectra w’ere run at 70eV.

Additional sample treatment helped in some identifications. Urea
adduction enriched the branched from the nonbranched components.
Sulfuric acid and/or bromine-CCb treatment (Morrison and Boyd
1969) identified unsaturated component peaks.

The organic carbon content of the sediment was determined by a
combustion technique using a Leco gasometric carbon analyzer.

220

Lytle, Lytle and Parker

Trace Metal Separation and Determination

Iron was removed from the sediment filtrate by extraction of
the iron-chloro complex from an 8N HCl solution of the total trace
metals with isopropyl ether. Separation of all the trace elements of
interest from matrix materials except Mo and Fe w'as effected by ad-
justing: the pH to ca 3, adding sufficient ammonium pyrrolidine dithio-
carbamate (APDC) to make the final solution 0.5% in APDC and ex-
tracting the APDC-metal complexes with methyl isobutyl ketone
(MIBK). Molybdenum w'as extracted from a fresh batch of filtrate
by adding KCNS and SnCla to make a final IN HCl solution 2% in
both. The molybdenum-CNS complex was extracted with isopropyl
ether.

All metals but molybdenum were analyzed using the Perkin-
Elmer 303 atomic absorption spectrophotometer. The fuel was acety-
lene and the oxidant, compressed air. The MIBK solution of cobalt,
cadmium, copper, manganese, nickel and zinc and the aqueous solution
of iron were aspirated directly using the following wavelength set-
tings (in nm) : Cd-229, Co-241, Cu-325, Fe-248, Mn-279, Ni-232,
Pb-283 and Zn-214. The color intensity of the Mo-CNS complex was
measured at 460 nm on the Beckman Model DU spectrophotometer to
determine molybdenum concentrations. Appropriate standards and
blanks were prepared for all determinations and any necessary cor-
rections were applied.

RESULTS

Organic

The hydrocarbons from the marsh plants and the blue-green algal
mats were assumed to be the main source of the hydrocarbons for the
ecosystem studied. Microorganisms were present, but their contribu-
tion is taken to be small in an organic-rich sink. Table 2 summarizes
the analytical results.

Normal straight-chain hydrocarbons ranging from C15 to C33
were identified in the sediments. Pristane and phytane were present
in low concentrations. Phytane was detected only in trace amounts.
The branched hydrocarbons were only a very small percentage of the
total hydrocarbon fraction. These branched hydrocarbons were al-
most completely olefinic.

The sediment hydrocarbon distribution was bimodal with maxi-
mums at Cl7 and C29 (Fig. 2) . However, the largest concentration of
hydrocarbons was in the C27 to C31 range. This reflects the hydro-
carbon pattern found in the two prominent contributors to the organic
matter, blue-green algae and higher plant life. Normal alkanes iso-
lated from the marsh plants showed an odd-carbon number predomi-

Geochemical Marsh Study

Table 2.

Marsh Plant and Sediment Analytical Analyses

221

Marine Plant

%

lipid
dry wt

%

HC

dry wt

%

HC

lipid wt

Ma j or
Compo-
nent

Limon'iion carolinianum

2.02

0.057

2.8

n-C29

Batis mavitima

2.31

0.008

0.37

n-C27

Salicomia higelovii

2.14

0.005

0.26

n-C31

Lyaium aavolinianum

3.41

0.085

2.3

I1-C29

Sesuvium maritirnwn

1.91

0.005

0.32

n-C25

Oenothera dmmiond'i'i

4.72

0.11

2.3

n-C29

Bond chi a frutesoens

4.91

0.043

0.75

n-C29

Sediment

0.15

0.0011

0.75

n-C29

Figure 2. Histogram for normal hydrocarbons of Harbor Island sedi-
ments.

222

Lytle, Lytle and Parker

nance with C27, C29, and C31 the most prominent alkanes while C17
was the most prominent alkane in blue-green algae.

The urea adducted fraction of the hydrocarbons indicated that
the cluster of peaks in the CIS range were not isoprenoid alkanes, but
were branched-hydrocarbons with their branching points near the end
of the chain. The large number of peaks in the C19 to C20 range were
completely absent from the urea adducted fraction indicating that
they were more highly branched. These peaks disappeared almost
completely after treatment with H.SO, indicating that they w^ere ole-
fmic.

Infrared spectra of urea adducted and urea nonadducted frac-
tions confirmed the evidence that the olefins present in the sediment
hj'drocarbons are primarily multibranched olefins, in conclusion, the
following generalizations can be made about the hydrocarbon results
for the sediments analyzed :

(1) All samples e.xhibitcd an odd-carbon preference.

(2) A bimodal distribution was exhibited in the hydrocarbon
patterns of sediments whose organic matter was derived
from both blue-green algae and terrestrial plant sources.

(3) The amount of extracted lipid material in plants was on the
order of 25 times as great as that extracted from the sedi-
ments.

(4) Pristane and phytane were found in all samples.

Normal alkane distributions in the plants were demonstrated be-
tween Cl 3 and C35 but for the most part ranged between C23 and
C31. Isoprenoid hydrocarbons, farnesane, pristane, and phytane, were
identified, but this fraction represented less than 2% of the total frac-
tions. Branched-chain hydrocarbons, iso-C27, C29 and C31, anteiso-
C26, C28 and C30, were tentatively identified. There were some un-
identified peaks in each chromatogram, but those peaks represented
less than 10% of the total hydrocarbons. Borrickia fr'utescens con-
tained a large number of branched and oleftnic peaks in the C15 to
C18 molecular weight range, none of w'hich was identified (approxi-
mately 21% of the total hydrocarbons fell in this Cl 5 to C18 range).
In all other plants analyzed the lower molecular weight hydrocarbons
represented less than 5% of the total weight.

There was a definite odd-carbon preference in every plant sample.
The largest component in four of the species was C29. The distribu-
tions were slightly different for each species. Gas chromatographic
analyses, tabulated in terms of individual normal hydrocarbons from
C15 to 033, are presented in Fig. 3.

Olefins have been reported in the hydrocarbons of many plants
(Stransky and Streibl 1969), but there was little evidence of olefins

PERCENT Caf/POSITIDN

Geochemical Marsh Study

223

HYDROCARBON

Figure 3. Histograms of normal hydrocarbons in marsh plants.

224

Lytle, Lytle and Parker

being present in greater than trace amounts in any sample except
Borrichia frutescens. Microhydrogenation of olefins in this particular
sample yielded normal alkanes. However, some peaks in this sample
were not changed by hydrogenation; nor were they enriched in the
urea iionadducted fractions. Hence, they were tentatively identified as
branched hydrocarbons having their branching points near the end
of the carbon chain.

Gas chromatograms are shown for two plants which contained
unidentified peaks (Fig. 4). Tentative identification for the iso- and
cwiteiso-alkanes as was given in Fig. 5 was based on their response to
urea adduction, their inertness to HoSO^, and the absence of func-
tional groups in the IR spectra. Plots of retention time vs. carbon num-
ber for the three series, normal alkanes, ABC, and XYZ yields three
parallel lines which indicates that the latter series are two distinct,
homologous alkane families.

Trace Metals

In the sediments selected for trace metal analysis it was assumed
that primary sources of biogenic material were the marsh plants. By
choosing intertidal and supratidal sediments a large contribution from
blue-green algae was hopefully excluded. A summary of the trace ele-
ment data obtained from three surface and one subsurface sediments
is contained in Table 3* Included are the results of organic car-
bon determinations. Overall the levels of trace metals are quite low
when compared to an “average” sediment from the near-shore area
such as provided by Chester (i965). The distribution among the sedi-
ments of any one element when compared to another shows similar
trends so that it appears that no highly significant variations in trace
metal fixations exist among the four sediments. However in inter-
sediment comparisons it can be seen that significant differences do
exist among the individual sediments, indicating degree rather than
type as the more important aspect in investigating the effects respon-
sible for trace metal fixation in these sediments.

The fourteen marsh plants have a trace metal distribution which
is shown in Table 4. In Fig. 5 are displayed the ranges of trace metals
in the marsh plants and some results of other workers in summarizing
available data on terrestrial and on marine plants. Fitting certain
characteristics of both groups, it might be expected that the marsh
plants would assume some sort of “middle ground”. These marsh
plants appear to be slightly depicted in most elements except for
molybdenum with respect to the terrestrial-marine range. With the
principal source of heavy metals coming from the soil solution, it is not
surprising from viewing the trace metal array in the sediments that
these relatively low levels would be reflected in the biota of the
regions.

BATIS MARITIMA

SESUVUM MARITINUM

TIME MIN

Figure 4. Chromatograms of Batis maritima and Sesuvium maritinum hydro

carbon extract:

Operating Coyiditions

Column coating: FFAP

Temperature program: 150°-

Column dimensions: 150 ft.

210"^ at 4°C/min

X 0.02 in

Helium flow: 4 cc/min

X-tso-C25

Key

A-anteiso-C26

Y-iso-C27

B-anteiso-C2S

Z-iso-C29

C-a7itei8o-CS0

226

Lytle, Lytle and Parker

M n
Fe
Co
Ni
Cu
2n
Mo
Cd
Pb

0.05 01 0.5 1.0 5,0 10.0 500 lOO.O 500.0 1000.0

Concentration (;jg/6 dry wt.)

Figure 5. Ranges of trace metals in marsh plants — The range of each
metal among the plants is indicated by the solid bar for that metal ex-
tending from the minimum value to the maximum value- Also included are
bars extending from averages of terrestrial (T) to those of marine (M)
plants reported by Bowen (1966).

1

1 1 r-

f

1

M

“1

—i

T

T

M

T M

M T

T M

M T

M T

T .

M

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1

1 —

1

1 L_

DISCUSSION

Hydrocarbons

The odd-carbon predominance of the hydrocarbons in the Harbor
Island sediments reflected the hydrocarbon composition of the marine
plants, in particular the distribution in the C25-C31 range. The large
concentration of CT7 reflected the blue-green algal contribution to the
sediment hydrocarbons (Winter.s, Parker and Van Baalen 1969, Gelpi
et al. 1970). Thus, the correlation between biological and geological
lipids was found to exist, the sediments retaining the biological infor-
mation needed in order to recognize what type of life dominated their
surroundings. This correlation between biological and geological hy-
drocarbons can be extended to include the isoprenoid hydrocarbons.
Isoprenoids are abundant in the marine environment (Blumer 1965,
Clark and Blumer 1967, Blumer, Mullin and Thomas 1964, Blumer
and Thomas 1965). Pristane in marine algae was reported by Clark
(1966). Pristane and phytane were found in photosynthetic and non-
photosynthetic bacteria by Han et al. (1968). Isoprenoid hydrocar-

Distribution of Trace Metals in Harbor Island Sediments

Geochemical Marsh Study

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The standard deviations for metal analysis by plant are added and then averaged to arrive
at the Average Standard Deviation.

Geochemical Marsh Study

229

bons are found in petroleum and crude oils in very high concentra-
tions (Bendoraitis, Brown and Hepner 1962, Han and Calvin 1969).
The presence of only trace amounts of pristane and phytane in Har-
bor Island sediments indicates lack of pollution from these sources.
This simple model should be useful. It could be of economic signifi-
cance by its use in understanding the origin of fossil fuels. It could be
of social significance by its use in understanding the fate of organic
matter. With man’s environment being polluted at a rapid pace, it is
essential that we understand the fate of organic matter whether it be
in the geosphere, atmosphere or hydrosphere if we are to gain control
over our environment.

T race Metals

In an attempt to emphasize the fine differences in trace metal
levels in the Harbor Island Sediments a plot of trace metal vs organic
carbon was made and shown in Fig. 6. This selection of X-Y variables
was made assuming the percentage organic carbon is a measure of bio-
genic input to the sediments. The trends and apparent deviations from
these trends were observed rather than absolute levels to establish the
correlation of organic matter and trace metal levels.

Since a slightly different selection of marsh plants prevail in each
sediment area, a semi-quantitative estimate was made of the actual
biomass of plant material in a 25-foot radius of the three surface sedi-
ments and the ratios of the various plants in those areas revealed the
following plant contributors in decreasing order of importance:

Sediment No. 1 — Salicornia higel-ovii, Batis maritima, Svartina
alterniflora, Lycmm carolinianurri (Christmas berry), Li-
monium caroliniamim, Borrichia friitescens (Sea ox-eyes)
and Oenothera drummondii.

Sediment No. 2--Bpartina alterniffora, Avicennia germinans
(Black mangrove), Borrichia f^nttescens, Ma^haeranthera
phyllocephala, Monanthochloe littoralis (Key-grass), Hedy-
otis nig't'icans, Distichlis spicata (Marsh spike-grass) and
Spartma patens (Salt-grass) .

Sediment No. 3 — Salicornia bigelovii and Batis maritima.

Defining the relationship of individual plants or groups of plants
and the sediments for specific heavy metals was, in this limited study,
impossible. However, by comparing the plant data to the data in Fig.
6 and observing the sediment-plant association groups some rather in-
teresting information may be gained.

With respect to all elements but molybdenum (not shown in Fig.
6) a relative slump is seen in Sediment No. 4, the subsurface sample
with little contribution from marsh plants. Still other effects such as

ABUNDANCE 0

230

Lytle, Lytle and Parker

Figure 6. Metals organic carbon in Harbor Island sediments — Num-
bers 1-4 on the organic carbon axis correspond to the Harbor Island soil
sediment numbers from Table 3. Both ordinate and abscissa values are
on a linear but arbitrary scale. Only the ordinate uses the origin as a
value of zero. Organic carbon values: 1 — 0.66%, 2 — ^0.31%, 3 — ^0.22%,
4—0.41%.

Geochemical Marsh Study

231

migration of trace elements to the surface and concentration there is
a known phenomenon and might account for the surface enrichment.
It might be expected that this effect would be noticeable even in the
absence of the marsh plants. To a degree the marsh plants still could
contribute a significant portion of the trace metals. For any release by
dissolution by bacterial action of trace metals beneath the surface of
an intertidal or supratidal zone would certainly be impeded by the con-
tinual percolation downward of rainwater and receding tidal waters.

Most elements show a positive correlation with organic matter.
This previously has been deduced by several workers (Jenne 1968).
However, as Jenne pointed out, this contribution by organic matter acts
really as a secondary effect. In sediments not having an overwhelming
load of organic matter the organic matter with its trace metal load
can establish the proper conditions for inclusion of trace metals in
ferromanganese oxides. These oxides present in most oxidizing sedi-
ments are known to act as very effective trace metal accumulators.
The marsh plants may not complete the fixation of trace metals in
sediments but at least may start the process by acting as a source and
sediment conditioner.

Iron appears quite enriched in Sediments No. 1 and 3 relative to
Sediments No. 2 or 4. It seems that any explanation based on a physio-
graphical or geological basis would also alter the distribution of other
elements more so than observed, It could very well be that the marsh
plants are more active in the transport of this element than the
others. Neither Sediment No. 2 nor 4 received any sizeable contribu-
tion from Salieornia higelovU or Batis maritinum or others having
the highest concentrations of iron. Baas Becking and Moore (1959)
concluded that the majority of iron in marine sediments exists as com-
plexed iron so that it is possible that these two plants supply substan-
tial amounts of complexed iron in their plant litter.

Though molybdenum is extremely depleted in the surface sedi-
ments (Table 3), this metal maintains a normal level in the subsur-
face sediment. In the marsh plants the levels of molybdenum are the
only ranges above and including those predicted from averages of ter-
restrial and marine plants (Fig. 5). It is known that in surface, oxi-
dizing sediments molybdenum may be lost through its conversion to
the soluble molybdate ion whereas in subsurface reducing sediments
molybdenum as well as copper, iron, zinc et al. may be retained as
their insoluble sulfides. The marsh plants then must speed the deple-
tion of molybdenum from surface samples which after decay of the
plants is either leached out of the sediments or transported to reduc-
ing layers deeper in the sediments.

Lead is greatly enriched in Sediment No. 2. Some of the plants
in this area contain the highest levels of lead though their relative en-
richment over those species in other areas could not fully account for

232

Lytle, Lytle and Parker

this high level. The possibility of some lead or other metal contamina-
tion cannot be entirely dismissed in view^ of the debris washed in by
tidal action.

The marsh plants as a whole have exhibited an ability about as
strong as terrestrial and marine plants in accumulating a variety of
trace metals from their environment. In some cases the marsh plants
may act as sources for trace metal enrichment in the sediments and in
other instances, notably molybdenum, as an active depleting agent. The
coastal marsh environment widespread along the Gulf and Atlantic
coastlines will undoubtedly continue to be a primary target of heavy
metal pollution. The varying degrees of enrichment of the various ele-
ments both essential and nonessential in marsh plants should make
them of value in establishing base-line evaluations of heavy metal in-
ventories in a coastal area.

LITERATURE CITED

Baas Becking, L. G. M. and D. Moore. 1959. The relation between iron and or-
ganic matter in sediments. J, Sediment. Petrol. 29:454.

Bendoraitis, J. G., B. L. Brown and L. S. Hepner. 1962, Isoprenoid hydrocarbons
in petroleum. Anal. Chem. 34:4.

Blumer, M. 1965. Organic pigments; their long-term fate. Science 149:722.
Blumer, M., M. M. Mullin and D. W. Thomas, l.%4. Pri.stane in the marine en-
vironment, Helgolander wis.s. Meeresunters 10:187.

Blumer, M. and D, W. Thomas- 1965. Phytadienes in zooplankton. Science 147:-
1148.

Bowen, H. J. M. 1966. In Trace Elements in Biochemistry, Academic Press, New
York.

Chester, R. 1965. Elemental geochemistry of marine sediments. In Chemical
Oceanography Vol. 2, (editors, J. P. Riley and G. Skirrow), Academic Press.
Clark, R, C., Jr. 1966, Master's thesis, Woods Hole Oceanography Institution,
Woods Hole, Mass.

Clark, R. C. and M. Blumer. 1967. Distribution of paraffins in marine organisms
and sediment. Limnology and Oceanography 12:79.

Gelpi, E., H. Schneider, J. Mann and J. Oro. 1970. Hydrocarbons of geochemical
significance in microscopic algae. Phytochemistry 9:603.

Han, J. and M. Calvin. 1969. Occurrence of Ci> 2 -C 2 fl isoprenoids in Bell Creek
crude oil, Geochim. Cosmochim. Acta 33:733.

Han, J., E. D. McCarthy, M. Calvin and M. H. Benn. 1968. Hydrocarbon con-
stituents of the blue-green algae Nostoc muscorum, Anacystis nidulans, Phor-
midiivm Luriduvi and Cklorogloea fitsekii. J. Chem. Soc. (Sect. C), 2785.
Jenne, E, A. 1968. Controls on Mn, Fe, Co, Ni, Cu and Zn concentrations in soils
and water: the significant role of hydrous Mn and Fe oxides. In Trace Inor-
ganics in Water, American Chemical Society, Washington.

Metcalfe, L. D. and A. A. Schmitz. 1961. The rapid preparation of fatty acid
esters for gas chromatographic analysis. Anal. Chem. 33:363.

Morrison, T. R. and R. N. Boyd. 1969. In Organic Chemistry, 2nd Ed, Allyn and
Bacon, Inc., Boston.

Stransky, K. and M. .Streibl- 1969. On natural waxes, Xll. Composition of hydro-
carbons in morphologically different plants. Collection Czschoslov. Chem.
Commun. 34:103.

Winters, K., P. L. Parker and C. Van Baalen. 1969. Hydrocarbons of blue-green
algae: geochemieal significance. Science 163:467.

Gulf Research Reports

Volume 4 Issue 2

January 1973

An Examination of Legislation for the Protection of the Wetlands of the Atlantic and Gulf
Coast States

Anthony J. Haueisen

University of Mississippi

DOI: 10.18785/grr.0402.09

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Environmental Law Commons, and the Marine Biology Commons

Recommended Citation

Haueisen, A. J. 1973. An Examination of Legislation for the Protection of the Wetlands of the Atlantic and Gulf Coast States. Gulf
Research Reports 4 (2): 233-263.

Retrieved from http:// aquila.usm.edu/gcr/vol4/iss2/9

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AN EXAMINATION OF LEGISLATION FOR THE
PROTECTION OF THE WETLANDS OF THE ATLANTIC
AND GULF COAST STATES

by

ANTHONY J. HAUEISEN, J.D.

School of Law

The University of Mississippi
Oxford, Mississippi

INTRODUCTION

The most useful aquatic areas in the world are in serious danger
of destruction. The estuaries, where fresh water, land and sea meet
in a dynamic and highly productive zone, are today gravely threatened
through unwise and unplanned excessive use of their valuable but
finite capacities. Estuaries include the coastal zone which is affected
by both the run-off of fresh water from the land and the salt w^ater
from the sea. This zone includes tidal rivers, marshes, bays and river
mouths. The value of these estuarine regions has been well established
by biologists. However, this value is fully appreciated by only a hand-
ful of people. The intense uses to w^hich the coastal zone is being placed
are so expan.sive, so competitive and potentially so destructive. Much
shipping for industrial and military purposes begins and ends in es-
tuaries. The waste products of the industries which crowd the coastal
zone and of one-third of the population of this country are daily being
poured into these waters. Estuaries are directly linked to suitable con-
ditions needed for the development of three-fourths of the fish and
shellfish taken for food production and recreational fishing. In addi-
tion, the use of these so important coastal regions is ever increasing
by our growing population for aesthetic and recreational purposes.

Each of the above uses is an important human use. However, each
of these uses is potentially destructive. 'Even aesthetic uses, which
have heretofore been above reproach, can irretrievably destroy this
fragile ecosystem when vast areas are developed for housing projects
by dredging and filling in the land. In fact, this action may destroy
the very rea.son why people wish to move to the shore. All of these
uses, and others, have developed without sufficient comprehension of
their effects and interactions and totally without planning for an
optimal balance for present and future human uses. Present and po-
tential human uses involve vast and complicated economic problems,
political and geographic difficulties, and grave social and legal com-
plications. It is with the latter, the legal problems of wetlands pre-
servation and utilization, that this paper is concerned.

It is proper that I should here acknowledge the work of Mr.
Lionel Eleuterius of the Gulf Coast Research Laboratory who first

233

234

Haueisen

visualized the need for a collection and examination of the laws of the
various coastal states in regard to the preservation of their wetlands
and who personally assembled the raw materials out of which this
paper is formed. Furthermore, I should acknowledge Dr. Thomas
Lytle, also of the Gulf Coast Research Laboratory, who recommended
this project to me and guided my research.

This paper will be divided into several distinct sections. It is
initially important that we define just what we are examining; there-
fore, the first part of this paper will describe what wetlands are and
attempt to explain something of the ecology of this region. The next
part of the paper will examine the laws of the coastal states to deter-
mine what, if anything, they have done to preserve and utilize their
wetlands. The last part of the paper will examine what states may do
to protect their wetlands, contain a legislative model for a wetlands
act, and discuss possible implementation of this proposed legislation.

WHAT ARE WETLANDS?

Wetlands is a general term used to describe the water-land inter-
face, whether ocean and shore or river and bank. However, in this
paper we will be more concerned with the ocean-shore interface be-
cause of the extreme richness of this particular area and its vulner-
ability. Wetlands may occur as a dividing fringe along shorelines in-
terposed between permanent dry land and open surface water expanses
of rivers, estuaries, etc. They may also exist in another configuration ;
as rather extensive tracts continuing for hundreds of miles and acres
in area.

There are numerous local names by which wetlands are known.
Some of these are salt marsh, tidal marsh, marshland, tideland, sub-
merged land, swamp, slough, bog, mud flats, wet meadow, or flood plain.
People tend sometimes to use these terms interchangeably, although
ecologists differentiate between many of these terms. The following
clas.sification system is derived from an extensive analysis of wetlands
in Maryland.^ The classification system briefly describe.^ each type of
wetland and outlines very generally the more important physical and
ecological characteristics distinguishing each. In examining wetlands,
we are concerned with two major distinct groupings: 1.) those wet-
lands which occur in inland, freshwater areas, and 2.) those which
occur in coastal areas and are our especial area of interest.

I. Inland, freshivater wetlands.

In this category of freshwater, non-tidal wetlands there are seven
types of wetlands included. They are :

1 Maryland State Planning Department, Draft Report-W etlands In Maryland,
January, 1970, p. V3.

Wetlands Legislation

235

1. Seasonally flooded basins and flats

2. Inland fresh meadows

3. Inland shallow fresh marsh

4. Inland open fresh water — ponds, lakes, etc.

5. Shrub swamp — found alonj? sluggish streams.

6. Wooded .swamp — often occurs on poorly drained uplands.

7. Bogs — usually waterlogged and having a spongy covering of
mosses.

II. Coastal Wetlands

This category can be further subdivided into two distinct areas.
These are fresh and saline marshes, which are distinguished by differ-
ences in shoreline elevation and the consequent varied influence of the
tide. This tidal activity partially accounts for the high productivity
of the coastal wetland as will be described later. There are seven types
of coastal wetland, three of which are in freshwater areas and four of
which are in saline areas.” They are:

A. Fresh Areas

1. Coastal shallow fresh marsh — average mean high tide
may cover this area with up to 6 inches of water.

2. Coastal deep fresh marsh — average mean high tide may
cover this area with from 6 inches to 3 feet of water.^

3. Coastal open fresh marsh — these are usually more or less
enclosed tidal ponds, and vegetation is usually scarce or
lacking because of the turbidity of the water due to the
tidal cycles and currents which keep sediment and detri-
tus in suspension.

B. Saline Areas

1. Coastal salt meadow — this soil is always waterlogged;
but because of its elevation, it is rarely covered by tidal
waters.

2. Irregularly flooded salt marsh — the soil in this area is
covered by a few inches of water at irregular intervals.

3. Regularly flooded salt marshes — ^the soil is covered at
mean high tide by one-half foot or more of water. This
is the area with which we are specifically interested. This
area is important as nesting areas for gulls and rails, as
feeding areas for herons, as habitat for mussels, snails,
and crabs and is used by fish and crustaceans.

4. Sounds and bays — this type consists of submerged land
under the open waters of sounds or bays. Vegetation is

2 Ibid., p. V-7.

3 The difference in depth of saltwater inundation determines the types of vege-
tation and animal life found in these areas.

236

Haueisen

usually scarce. However, this area is important because
it is the habitat of fishes, oysters, mussels, shrimp, clams,
crabs, and many other invertebrates upon which these
species feed.

This necessarily brief description of the terms used in defining wet-
lands will serve to illustrate just how minute and complex terminolo-
gies can become when dealing with any involved and intricate natural
system.

BASIC WETLANDS ECOLOGY

The term ecology has very recently come into vogue. However,
the language and techniques of ecology and ecological research are
largely unknown to the public. Ecology may be defined as the study
of the interrelationships of organisms to one another and to their en-
vironment. It is possible to divide a study of an ecological system into
two distinct parts: 1.) the structure of the ecosystem itself, including
quantity and distribution of plants and animals and the physical char-
acteristics such as temperature, light, pH, salinity, dissolved oxygen,
etc.; and 2.) the function of the ecosystem, including the rate and
amount of biomass production'* and the cycling of nutrients within
the biotic community. Without a knowledge of these ecological princi-
ples and other information, it would be impossible for well-intentioned
individuals to formulate sound natural resources management policies.
Therefore, I will briefly describe some general principles of ecology
and some concepts concerning the marsh ecosystem.

Since tidal marshes are ecological formations resulting from the
invasion of shallow water by land vegetation, it would perhaps be best
to examine those areas best suited to this invasion. The most obvious
place for land vegetation to invade the shallow sea is along the edges
of the vast coastal plain which makes up a part of the continental
shelf. This coastal plain extends along most of the Eastern and Gulf
coasts of the United States but is largely lacking along the Pacific
Coast. In early geological history, the present continental shelf was,
for the most part, also the coastal plain. Today, the continental land
mass is less expansive and much of the former coastal zone is sub-
merged. Some submergence is still occurring, at a rate of about one
foot per century; but the coastal area is just about holding its own
due to the seaward transport and deposition by rivers of sediments
from inland. However, the levee-building activity of the Army Corps
of Engineers is diverting this sediment from the coastal zone and ac-
tually forcing it out to the edge of the continental shelf where it is
lost for human use for all practical purposes. In some coastal areas,

4 Biomuss may be defined as the total quantity at a given time of living orga-
nisms of one or more species per unit of space (species biomass), or of all the
species in a community (community biomass).

238

Haueisen

marshes. Among marine species only a relatively few can adjust to
the rapid salinity changes which occur with each tide. Those few
species that can endure the tidal marsh conditions, however, are rela-
tively free of the kinds of competitors and enemies that harass related
species in nearby waters.

As in other ecosystems, the same general relationships and com-
ponents exist in a tidal marsh. Plants utilize sunlight energy in their
growd:h and reproduction and are the primary producer component of
the ecosystem. They are a source of food for grazing animals. These ani-
mals in turn*furnish food for carnivores. Bacteria and fungi comprise
the component which reduces or decomposes the dead organisms to
inorganic levels, returning such nutrients as phosphates and nitrates
to the marsh system where they are re-used by the plants in the man-
ufacture of more plant tissue. In conjunction with the re-use of the
nutrients and the one-way flow and ultimate release of the energy first
stored by the plant life, it is necessary to add to this simple structure
other natural trends and cycles which affect the marsh ecosystem. The
tides redistribute nutrients and sediments throughout the tidal marsh.
They also affect the overall primary productivity by decreasing or in-
creasing exposure of the microscopic algae and marsh plants or the
quantity of phytoplankton that is favorably exposed to sunlight as
the volume of water over the marsh changes.

It has been claimed that “tidal marshes and estuaries . . . are
among the most fertile areas in the world in terms of energy, calories,
proteins, carbohydrates and vitamins.’’ ^ Ecological studies have
shown that “gross income from the best market farms . . . runs as
high as $2,000 per acre per year. On a comparative basis, the major
marshlands are producing $4,000 worth of nutrients per acre per
year.” ^ Fertility of estuaries also results from year-round primary
production. Wetlands ecosystems tend to maintain a constant rate of
production during seasonal environments. For example, marsh grass
produces at least tw'o crops per year as compared with wheat which
grows only a few months with zero growth for many months. The an-
nual production of a marsh, or an estuary as a whole, may be double
or triple that of ordinary agricultural land simply because it produces
two or three times as long each year.

Now that we have very briefly examined some basics of marsh
ecology and its importance, it is time to turn to the next major area
of this paper, an examination of the various coastal states with re-
gard to those laws which have been enacted to protect their wetlands.

® Robert L. Dow, Maine*s Coastal Marshlands: Their Values, Present and Fu-
ture (Augusta: Maine Department of Sea and Shore Fisheries, 1962), p. 3.

® Robert L. Dow, Economic Yields of So^ne Maine Coastal Wetlands (Augusta:
Maine Department of Sea and Shore Fisheries, 1966) , p. 1.

Wetlands Legislation

239

LEGAL APPROACHES TO THE PROTECTION OF WETLANDS

In this chapter, I shall examine statutes from the states on the
Atlantic Coast and on the Gulf Coast of the United States and shall
emphasize those which act to protect wetlands. First, I will examine
the Federal laws relative to this ai'ea; and then, I will examine the
laws of the states in geographical order from Maine to Texas.

A. FEDERAL LEGISLATION

The English Common Law has come into all of the United States,
with the exception of Louisiana, and has evolved through case inter-
pretation by the Supreme Courts of these various cases. The essential
common law principle is that title to soil under navigable waters is in
the sovereign except as far as private rights have been acquired in an
expressed grant from the original sovereign. During the time of the
original thirteen colonies, the tidelands and the submerged lands were
under the complete control of the Crown and they were held by the
Crown in trust for the people as a whole. The riparian owner has own-
ership ordinarily only to the mean high water mark.

After the Revolutionary War, the American states succeeded the
Crown as Sovereign. The States continued to hold the tidelands and
submerged lands in a sovereign capacity in trust for the people sub-
ject to the public purposes of navigation, commerce, fishing, boating,
recreation and enjoyment free from obstruction and interference.

State ow'nership, and in some cases private ownership, of tide-
lands and submerged lands are subject to the paramount right of con-
trol by the Federal Government under the U.S. Constitution for com-
merce and navigation. As stated, all real property w^as originally in
the Sovereign. It w'as then granted in one w^ay or another to private
individuals. There are three types of grants from the Sovereign: 1.)
the first, was by the Lords Proprietors j 2.) by the Crown itself up
until the time of the Revolutionary War; and 3.) from the Revolu-
tionary War to the present in the form of State Grants,

The States of the Atlantic and the Gulf Coast have absolute title
to submerged lands. The state has prima facie title (i.e. it goes into
the court with a presumption of title) to the tidelands, i.e., the area
betw'een the mean high water mark and the mean low water mark. A
claimant to private owmership of tideland.s must come into court with
a chain of title, tracing hi.s title back to the original grant from the
Sovereign. This is sometimes difficult to do due to the destruction of
the intervening records. A claimant mu.st produce an original grant
which Is then presented to the court as a question of law as a con-
struction of that grant.

The Federal Government has jurisdiction over these tidelands,
submerged lands and navigable waters under the Navigation and

240

Haueisen

Commerce Clauses of the Federal Constitution. There is joint or con-
current jurisdiction between the Federal Government and the Gov-
ernment of the particular State over the navigable waters of the
United States which are also navigable waters of that particular
State. The navigable waters of a State include all of the navigable
waters of that State, whereas, the Federal Government only has juris-
diction over a certain percentage of tho.se waters which have been
classified, either by the Federal Congress or by some rule and regula-
tion, as being Federal navigable waters. These w'aters are usually
waters that are in continuous connection between different states or
between the state and the open ocean.

The other State waters are under the complete jurisdiction of
that particular State. The law, as far as the Federal Government is
concerned, was relatively stable up until 1947 w'hen after a period of
time the U.S. Department of Interior through the U.S. Attorney Gen-
eral’s Office began litigation first against the State of California and
later involving decisions in 1950 against the States of Texas and Lou-
isiana attempting to take over actual Federal ownership of all tide-
lands and submerged lands. The decisions of the U.S. Supreme Court
in effect completely reversed all prior law and stated that the Federal
Government was the absolute owner of these areas. The Congress
passed the Submerged Lands Act of 1953 w'hereby the Federal Gov-
ernment, in effect, abandoned the title to the area lying below the
mean high water line and left the actual ownership in the hands of the
State as determined by the State law. At the present time, the only
interest the Federal Government has in these areas is this easement
or servitude on the part of the Federal Government for the control of
commerce and navigation which goes up to the mean high water mark.
Congress also has an interest, of course, in any of these areas which
affect National Defense, international affairs, flood control and elec-
tric power production.

Some of these Federal Agencies with various powers of regula-
tion are the Federal Power Commission, U.S. Navy, U.S. Army Corps
of Engineers (which has essential control over interstate navigation
and the intracoa.stal w^aterway), and the U.S. Coast Guard wdth the
enforcement of revenue laws and boating laws.

In 1953, the Federal Government also passed the Outer Conti-
nental Shelf Lands Act which stated that the Federal Government had
absolute jurisdiction betw^een the outer limits of the state boundary,
which is usually taken to be three miles, but in the case of Texas and
Florida extends out to three marine leagues (which is ten and a half
miles) outside of the state's mainland area. The Federal Government
is supreme between the state boundary and the extent of the Conti-
nental Shelf, extending out in some areas 120 to 150 miles.

The State jurisdiction over the tidelands, submerged lands, and
navigable waters are co-extensive with the Federal Government in-

Wetlands Legislation

241

volving both the state navigable waters and the Federal navigable
waters. The State has exclusive jurisdiction over the state navigable
waters. Within each State there are a number of agencies and depart-
ments that have overlapping jurisdiction, regulation and control of the
tidelands and submerged lands.

The focus of the Federal Government with regard to the protec-
tion of the wetlands has been directed in two major areas: pollution
control and preservation of migratory bird nesting and feeding areas.
Both of these approaches may be helpful in protecting marshlands.
First, let us examine the role of the Federal Government in pollution
control.

The Federal Government entered into the area of water pollution
control in 1899 with the enactment of the Rivers and Harbors Act.'
This statute made it a criminal offense to deposit any refuse matter of
any kind or description into any navigable water of the United States.
In 1966. the U.S. Supreme Court held that “refuse" was not limited
to materials of no value but included any substance, whether or not
usable by industrial standards, which has a deleterious effect on navi-
gable waters,®

The Oil Pollution Act of 1924 was an enactment to deal with the
problem of oil discharges from vessels into coastal waters damaging
aquatic life, harbors and dock and recreational facilities.® The pro-
hibitions in this Act proved to be quite difficult to police as a practical
matter.

The Water Pollution Control Act of 1948 was also the really first
modern identifiable Federal program for water pollution control.^® It
stated that the pollution of interstate waters which endangered the
health or welfare of persons of another state is to be considered a
public nuisance subject to abatement and that the States were recog-
nized to have primary responsibility and the right to control water
pollution while the Federal Government had jurisdiction over the na-
tion’s interstate waterw'ays and their tributaries.*^ The Surgeon Gen-
eral was directed to coordinate and encourage cooperation among all
levels of government involved in pollution control and to engage in
joint activities with State and interstate agencies.*® The Act also en-
couraged the adoption of uniform State law’s and the creation of inter-
state pollution control compacts.*^ The Surgeon General was autho-
rized to prepare and adopt comprehensive programs to eliminate

7 Rivers and Harbors Act of 1899, 38 U.S.C, §§401-413 (1899).

s United StaUR v. Standard Oil Co., 384 U.S. 223 (1966) .

» Oil Pollution Act, 33 U.S.C. §§431-437 (1924).

10 Water Pollution Control Act, 62 Stat. 1155, 33 U.S.C. §§466 (1948).

** Ibid.

i2/6wi., at §466(b) (1952).

i3/6id., at §466(c) (1964).

242

Haueisen

pollution in waterways and support and aid technical research to de-
vise and perfect methods of treatment of industrial wastes.^^

In 1956, amendments to the original 1948 Act were passed which
provided for a much more intensive and well-organized Federal pollu-
tion abatement program than did the earlier law.'*^ They emphasized
the basic policy that water pollution problems were best solved at the
local level. Grants to States and to interstate agenciexS were authorized
for administration of water pollution control programs including com-
prehensive river basin programs involving control, research, and en-
forcement. It provided for technical assistance, the encouragement of
interstate compacts and uniform State laws, the appointment of a Fed-
eral Water Quality Advisory Board, and a cooperative program for
the control of pollution from Federal installations.

In 1961, the Federal Water Pollution Control Act Amendments
were enacted. These amendments extended Federal pollution abate-
ment authority to all interstate or navigable waters, increased con-
struction grants and authorized research facilities in various parts of
the country and the conducting of water quality studies in the Great
Lakes. It should also be noted that the amendments transferred the
administrative responsibilities for the program from the Surgeon
General to the Secretary of Health, Education and Welfare.^"^

In 1965, the Federal Water Quality Act was signed into law.^^
Briefly, this Act provided for the adoption and enforcement of water
quality standards for interstate waters and set up the Federal Water
Pollution Control Administration in the Department of Health, Edu-
cation and Welfare. It increased construction grants and authorized
research and development grants for preventing discharge of untreat-
ed wastes from storm sewers or combination storm-sanitary sewers.

In 1966, the Federal Clean Water Restoration Act became law.^®
This Act provided incentives to the States to adopt water quality
standards for pollution control and provided for Federal reimburse-
ment for qualified construction projects commenced at a time when
Federal grant funds were not sufficient to pay the full Federal share.
The purpose of this provision was to encourage a State prepared to
move ahead with sewage treatment plant projects to start building
without waiting for a Federal grant. Under this program, if the proj-
ect is Federally approved, the local governments may advance the
Federal share of the project themselves and be reimbursed with funds
as they become available. It authorized the use of Federal enforce-

Ibid.

Water Pollution Control Act Amendments of 1956, 70 Stat. 499, 33 U.S.C.
§466-466(k) (1964).

16 Ibid.

17 Ibid.

18 Water Quality Act of 1965, 79 Stat. 903 (1965).

i*> Clean Water Eestoration Act of 1966, 80 Stat. 1246 (1966).

Wetlands Legislation

243

merit equipment with relation to international boundary waters and
transferred responsibility for administration of the Oil Pollution Act
to the Secretary of Interior and changed that Act to include inland
waters.

In 1970, the Water Quality Improvement Act was enacted.^o This
Act forbade oil discharges into navigable waters, adjoining shorelines
and the waters of the contiguous zone and requires a National Con-
tingency Plan for removal of any spills. This Act provides that the
owner of a polluting facility can be fined up to eight million dollars
for the clean-up costs or more if there is willful negligence or willful
misconduct.

The second focus of Federal programs is concerned with migra-
tory wildfowl. The Migratory Bird Treaty Act of 1918 assigned to the
Federal Government primary jurisdiction over the protection of mig-
ratory birds, including wildfowl. This was followed in 1929 by the
Migratory Bird Conservation Act which provided the authority and
funds for the establishment of Migratory Bird Refuges. The first wa-
terfowl hunting stamp (at a cost of $1) was required by the Migra-
tory Bird Hunting Stamp Act of 1934. This Act helped finance the
refuge program.

The Federal Aid in Wildlife Restoration Act (Pittman-Robert-
son) of 1937 made it possible for many states to initiate programs of
wetlands acquisition and development. In 1949, an amendment to the
Hunting Stamp Act raised the price of the “duck stamp'’ to $2; an-
other in 1958 further increased it to ^3. The latter amendment also
specified that duck stamp funds be used exclusively for the purchase
of waterfowl production areas and suitable areas for migratory bird
refuges; also that as much as 40% of a refuge may be opened to hunt-
ing of migratory birds.

In spite of these efforts, loss of w^etlands continues at an alarm-
ing rate. In recognition of the growing problem, Congress in 1961
passed a bill which has become known as the Accelerated Wetlands
Acquisition Act (Public Law 87-383, approved Oct. 4, 1961). This Act
makes possible the stepped-up purchase of essential wetlands now,
while they still exist. In effect, Congress could loan the conservation
movement $105 million which, plus the revenues from duck stamp
sales, could be used for this purpose. This money is to be used to pro-
mote the conservation of migratory waterfowl and to offset or prevent
the serious loss of important wetlands and waterfowl habitat essential
to the preservation of such waterfowl. At the end of 7 years, 75 %
of each year’s duck stamp revenues must be used for repayment of the
loan. The task of this legislation is obvious — to save at least an essen-
tial minimum of the nation’s wetlands.

“t* Water Quality Improvement Act of 1970, 84 Stat. 91 (1970).

244

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B, STATE LEGISLATION

1. Maine

Maine. Revised Statutes Annotated, Title 12, ch. 421, § §4701—4709
( Additional Supp. 1967 )

No person, agency or municipality may remove, fill, dredge or
drain sanitary sewage into, or otherwise alter any swamp, marsh, bog,
beach, flat or other wetland bordering coastal waters, or fill, dredge or
drain sanitary sewage into such waters within such area, without fil-
ing written notice to do so, with plans, to the m\micipal officers and
the Wetlands Control Board. A public hearing is held after receipt of
such notice. After the hearing, a permit is issued if the Wetlands Con-
trol Board approves. Approval may be conditioned upon the applicant
amending his plans. Approval may be withheld by the municipality or
the Board if either body finds the plans damaging to fish life. An ap-
peal is provided for. Anyone violating any provision of this statute is
subject to a maximum fine of $100. A continuing violation of this
statute may be enjoined.

Maine Revised Statutes Annotated, Title 30, ch. 229, ^4001 (Addi-
tional Supp. 1967 ).

Wetlands may be ‘*taken” by a municipality with the consent of
the owner and payment of compensation.

Maine Revised Statutes Annotated, Title 30, ch. 229, §300J (Addi-
tional supp. 1967 ).

Any municipality may receive wetlands as devises or gifts.

2. New Hampshire

Wetlands regulation had a modest beginning in New Hampshire
in 1955 when the legislature made it illegal to create land by filling in
great ponds — lakes or ponds over 10 acres in size — ^without permission
of the Governor and Council. Before that time, apparently a person
could convert the state’s water into his own land by the simple ex-
pedient of filling it in.

This statute dealt with public inland waters, not wetlands as such
and was the extent of any regulatory effort until 1965. Then, in an act
regulating sewage disposal systems on islands, it was provided that no
one could fill in a marsh bordering on or adjacent to a great pond of
the State for building purposes without approval of the sewage dis-

21 New Hampshire Revised Statutes Annotated 482:41-a to 41-d.

Wetlands Legislation

245

posal system in accordance with local zoning ordinances of the mu-
nicipality, or in absence of same, the Water Supply and Pollution Con-
trol Commission. The emphasis of this statute, though, was on the
prevention of pollution and not the preservation of wetlands*

In 1967, New Hampshire got its first full-fiedged wetland regula-
tions — three dredge and fill laws. The first prohibited any person,
firm or corporation from excavating or dredging any bank, flat, marsh,
swamp or lake bed that lies below the natural mean high water mark
of any fresh public waters of the state without petitioning the Water
Resources Board. The second replaced the 1955 law regulating plac-
ing fill in fresh public waters with a more up-to-date version. The
third prohibited persons from excavating, removing filling or dredg-
ing any bank, flat, marsh or swamp in and adjacent to tidal waters
without approval of the New Hampshire Port Authority.®''^

In 1971, Bill No. 228 was introduced into the New Hampshire
House of Representatives and dealt with excavating, filling, mining
and construction in the inland waters of the state and established an
inland wetlands authority to which anyone seeking to alter any of the
existing interior wetlands of the state would have to make petition for
a permit to do so.^®

3. Massachusetts

Massachusetts General Laws ch. 131, §40 (1967) entitled: Protec-
tion of Flood Plains.

No person shall remove, fill or dredge any bank, flat, marsh, mea-
dow, or swamp bordering on any inland waters without filing notice of
hia intention to do so, with plans of the proposed activity, wuth the
local authority and with State departments of public works and nat-
ural resources, A public hearing is provided for. The local authority
may recommend protective measures in the public interest, which are
submitted to the Commissioner of Natural Resources. If the area
where the w’ork is to be done is essential to proper flood control, the
Commissioner may impose conditions necessary to protect the public
interest, which must be complied with. A continuing violation of this
section may be enjoined.

Massachusetts General Laws ch. 130^ ^105 (Additional supp. 1966)

Because of the urgent necessity of protecting coastal w’etlands,
the Commissioner of Natural Resources was given the power to adopt,

22 New Hampshire Revised Statutes Annotated 149 :4^^.

23 New Hampshire Revised Statutes Annotated 488-A (1967).

24 New Hampshire Revised Statutes Annotated 482:41-e to 41-i.

25 New Hampshire Revised Statutes Annotated 483-A, §1-5.

2« New Hampshire Revised Statutes Annotated 483-B, §1-26.

246

Haueisen

amend, modify or repeal orders regulating, restricting or prohibiting
dredging, filling, removing or otherwise altering, or polluting coastal
wetlands.Violations of the Commissioner’s order is punishable by a
fine of between $10 and $60 and/or maximum imprisonment of one
month.

However, if the Commissioner’s order so restricts the use of the
property as to deprive the owner of its practical use, a court may de-
cree that the order does not apply to that owner’s land. In such a case,
the Department of Natural Resources may take the land for the State
by eminent domain.

Massachusetts General Laws ch. 40 ^8(c) (Additional supp. 1968)

This section empowers a city or town to acquire by gift, pur-
chase, grant, bequest, devise, lease, or otherwise the fee or any lesser
interest in wetlands and open spaces. It also empowers a city or town
to take land by eminent domain for conservation purposes.

Massachusetts General Laws ch. 132 A, §11 (Additional supp. 1968)

The State may reimburse a city or town up to 60% of the cost of
acquiring land for conservation purposes pursuant to §8(c) of ch. 40.

Massachusetts General Laws ch. 130, §2 7 A (Additional supp, 1966)

No person shall remove, fill or dredge any bank, flat, marsh,
meadow or swamp bordering on coastal waters without filing written
notice of his intention to do so with the local licensing authority, the
State Department of Public Works, and the Director of Marine Fish-
eries. Restrictions may be placed on such work and such work must
be done subject to these restrictions. Violations of this section are
punishable by a maximum fine of $100, imprisonment for not more
than 6 months, or both. A continuing violation of this section may be
enjoined.

A case concerning this section held that it does not authorize an
absolute prohibition against the filling in of a privately owned marsh-
land if the result would be that the owner would be so deprived of the
practical uses of his land so as to amount to a taking of his land with-
out compensation.

Whether there had been such a deprivation of the practical uses
of the marshland as to be equivalent to a taking without compensation
depended upon the uses to which the marshland could be put without
violating the statutory prohibition against the marsh. Since the evi-
dence at trial on this issue was lacking, the injunctive decree of the

27 Commisaioner of Natural Resources v. S. Volpe & Co., 206 N.E. 2d 666 (Mass.
1966).

Wetlands Legislation

247

trial court was reversed and the case was remanded for additional
findings on the above issue.

4. Rhode Island

Rhode Island General Laws Annotated ^^2-1-13 to 2-1-17 (Addi-
tional supp. 1967 )

These acts established a public policy of preserving the coastal
wetlands of the State. The Department of Natural Resources may,
after public hearing, designate coastal wetlands or parts thereof, the
ecology of which shall not be disturbed. This designation will be re-
corded in the registry of deeds in each city or town where the land is
located. The right of appeal is allowed for 2 years after recordation.
Provision is made for award of damages.

Rhode Island General Laws Annotated ^^11 -46. 1-1 (Additional
supp, 1967)

Anyone who dumps or deposits mud, dirt or rubbish upon, or who
excavates and disturbs the ecology of intertidal saltmarshes, without
first obtaining a permit therefor issued by the Department of Natural
Resources shall be fined for each offense $100: $50 to the State and
$50 to the complainant. The Director of Natural Resources shall re-
fuse to issue such a permit if in his judgment the dumping or deposit-
ing of mud, dirt or rubbish or excavation would disturb the ecology of
intertidal saltmarshes.

5. Connecticut

Conn. Gen. Stat. Ann. Title 25, ch. 473, §26-17a. (1967)

The State Board of Fisheries and Game is empowered to do the
following:

(1) Acquire wetlands, or any easements, interests or rights
therein, by purchase, exchange, condemnation, gift, devise,
lease or otherwise.

(2) Enter into agreements with owners of wetlands to conserve
wetlands.

(3) Enter into leases with an option to purchase wetlands, pro-
vided :

a. approval of the Commissioners of Agriculture and Natu-
ral Resources is obtained, and

b. the lease does not exceed 10 years.

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Haijeisen

(4) Take wetlands by eminent domain.

(5) Secure title to wetlands by paying to a municipality the
amount of the municipality's tax liens on such wetlands,
where the municipality's property tax on such wetlands is
unpaid for 6 years.

Conn. Gen, Stat. Ann. Title 25, ch. 473, ^25-10 to §25-77 (1967)

These statutes provide for the dredging of sand and gravel from
lands under tidal and coastal waters. This is regulated by the Water
Resources Commission, supplemented by a member designated by the
Shellfish Commission. Public hearings must be held. Shore erosion,
navigation, and living resources must be considered.

Local zoning for marshland protection has been unsuccessfully
attempted in Connecticut.

6. New York

Long Island Wetlands AcC^

This Act permits the State to enter into cooperative agreement
with towns, villages, or counties for the purpose of preserving and
maintaining wetlands on Long Lsland which have been dedicated for
conservation purposes. The State is empowered to provide one-half
of the cost of maintaining such areas.

New York Conservation Law, §429a-g (1967 )

These sections require the issuance of a permit by the Water Re-
sources Commission before anyone may alter the waters of the State.
The Commission may issue the permit subject to conditions upon
which the work must be done. The following activities require a per-
mit: altering the channel of a stream, removing materials from a
stream, excavating or filling in navigable waters, erecting an im-
poundment structure, dock or wharf in or across a natural stream or
watercourse. A violation of this statute constitutes a misdemeanor,
punishable by a maximum fine of $500, maximum imprisonment of 1
year, or both.

New York Public Lands Law, art. 2, §3.5 (Additional supp. 1967)

The Commissioner of General Services may license and regulate
the business of taking sand, gravel or other materials in or upon lands
under water and may prescribe the terms and conditions under which

Dooley v. Town Zoning Commission, 157 Conn. 304, 197 A.2d 770 (1964).

29 New York Conservation Law, §394 (1967).

Wetlands Legislation

249

the same may be taken. After adoption of regulations by the Com-
missioner, it shall be unlawful to take or remove from lands of the
State under water any sand, gravel, or other material, without a
license.

!\etv York Public Lands Law^ art. 6, §§75-7^ (Additional supp.
1967)

Empowers the Commissioner of General Services to grant lands
under water to a county, city, town or village for conservation and
other purposes*

7. New Jersey

The W^etlands Act of 1970^^

This Act recognizes the ecological importance of the coastal wet-
lands, and states that it is necessary to prevent the deterioration and
destruction of these lands in order to preserve the ecological balance
of the coastal area.

The Act directs the Commissioner of the Department of Environ-
mental Protection to make an inventory and maps of the wetlands.
The Act authorizes the Commissioner to make regulations restricting
or prohibiting dredging, filling, removing, or otherwise altering or
polluting the coastal wetlands. The Act requires the Commissioner to
hold a public hearing before adopting any regulations concerning the
wetlands.

The Act prohibits any regulated activity from being carried on
without a permit from the Department of Environmental Protection.
The Commissioner must consider the ecological effect of the work to
be performed before issuing a permit. The Act provides that any per-
son having a recorded interest in land affected by the Commissioner's
regulations may file a complaint in the Superior Court to determine
if the regulations deprive him of the practical use of his land. The Act
provides a fine of $1,000 to be levied against violators of the regula-
tions promulgated by the Commissioner, and also makes violators
liable for the cost of restoration of the affected coastal wetland to its
condition prior to the violation in so far as is possible.

iS.J. ST AT. AI\I\. 12:5-3 to 12:5-3 (1914)

The Board of Commerce and Navigation must pass on all plans
for development of waterfront which involves the construction or al-
teration of a dock, wharf, pier, bulkhead, bridge, pipeline, or any

1 could find no other citation. In the official legislative copy, all that is said
is that it may be cited as the Wetlands Act of 1970.

250

Haueisen

other similar or dissimilar waterfront development. Public hearings
may be held. No provision is made for eminent domain. No reference
is made to protection of natural resources.

NJ. ST AT. ANN. 13:8A-I to 13:8A-18 (1961 )

This is New Jersey’s Green Acres Land Acquisition Act of 1961.
The Act provides for purchase of lands for public recreation and con-
servation of natural resources. A sum of $60 million was made avail-
able by a Green Acres Bond referendum. The acquisition program is
under the direction of the Commissioner of Conservation and Econom-
ic Development. Of the total amount available, $20 million was for the
purpose of supporting local acquisition. In addition to fee simple ac-
quisitions, acquisition of conservation easements is permitted.

8. Delaware

Delaware Coastal Zone AcC^

This Act states that the coastal areas of Delaware are the most
critical areas for the future of the State in terms of the quality of life
in the State. It, therefore, declared that it is the public policy of the
State of Delaware to control the location^ extent and type of industrial
development in Delaw^are's coastal area. In so doing, it is thought that
the State can better protect the natural environment of its bay and
coastal areas and safeguard their use primarily for recreation and
tourism. Specifically, this chapter seeks to prohibit entirely the con-
struction of new' heavy industry in its coastal areas, which industry is
determined to be incompatible with the protection of that natural en-
vironment in those areas. While it is the declared public policy of the
State to encourage the introduction of new industry into Delaware,
the protection of the environment natural beauty and recreation po-
tential of the State is also of great concern. In order to strike the cor-
rect balance between these two policies, careful planning based on a
thorough understanding of Delaware's potential and her needs is re-
quired. Therefore, control of industrial development other than that
of heavy industry in the Coastal Zone of Delaware through a permit
system at the State level is called for. It is further determined that
offshore bulk product transfer facilities represent a significant danger
of pollution of the Coastal Zone and generate pressure for the con-
struction of industrial plants in the Coastal Zone, which construction
is declared to be against public policy.

This Act, as seen from the above, is quite a radical departure
from the historic and traditional policy of every state to encourage
any industry it can to build and operate from that state. Since the Act

•■^1 DEL. CODE ch. 70, §7001-7014.

Wetlands Legislation

251

is so new, it cannot as yet be determined what economic effect this
Act will have on the State of Delaware; but it appears to be a serious
and determined effort to protect the coastal zone.

9. Maryland

ANN, CODE of MD. §§715-731 (1967 Replacement Vol.)

This Act declares that in many areas of the State much of the
wetlands have been lost or despoiled by unregulated dredging, dump-
ing, filling, etc. and that the remaining wetlands of the State are in
jeopardy of being lost. It declares that it is the public policy of the
State to preserve the wetlands and to prevent their despoliation and
destruction, §719 defines **state wetlands” and ‘'private wetlands”.
§721 declares that it is unlawful for anyone to dredge or fill in State
wetlands without a license to do so by the Board of Public Works. Any-
one violating the provisions of this section is deemed guilty of a mis-
demeanor and may be fined not less than $500 and not moro than
$1,000. Each violation shall be a separate and distinct offense, and in
the case of a continuing violation, each day’s continuance thereof will
be deemed to be a separate and distinct offense. Any land created in
violation of this section will be the property of the State.

§723 states that the Secretary of Natural Resources may from
time to time promulgate rules and regulations governing dredging,
filling, removing or otherwise altering or polluting private wetlands.
Provisions are made for a permit to carry out any of the above activi-
ties on private wetlands. Appeal may be taken to the Board of Review
of the Department of Natural Re.sources and from there to a circuit
court in the county in which the land is located. If the court rules that
the restrictions under the Secretary’s rules and regulations constitute
a taking of land without compensation, the Secretary of Natural Re-
sources may proceed to condemn the land or interests therein and take
it by eminent domain.

10. Virginia

Virginia has no specific statutes in the Code relative to coastal
wetlands protection. There are some statutes and a constitutional ar-
ticle that do provide some measure of protection to some of the wet-
lands. Two statutes and the constitutional provision listed below
pertaining to ownership of lands are particularly pertinent. The Con-
stitution of Virginia states that:

The natural oyster beds, rocks and shoals, in the waters of
this State shall not be leased, rented or sold but shall be held
in trust for the benefit of the people of this State, subject to

252

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such regulations and restrictions as the General Assembly
may prescribe, but the General Assembly may, from time to
time, define and determine such natural beds, rocks or shoals
by surveys or otherwise.^^

Some wetlands and shallows are included within the area thus
protected.

The Code states that:

All of the beds of the bays, rivers, creeks and the shores of
the sea within the jurisdiction of this Commonwealth, and not
conveyed by special grant or compact according to law, shall
continue and remain the property of the Commonwealth of
Virginia, and may be used as a common by all the people of
the State for the purpose of fishing and fowling, and of tak-
ing and catching oysters and other shellfish . .

The Code further states that;

All unappropriated marsh or meadow lands lying on the
Eastern Shore of Virginia, which have remained ungranted,
and which have been used as a common by the people of this
State, shall continue as such common, and remain ungranted,
and no land warrant shall be located upon the same . .

However, an accurate designation of those lands used as a com-
mon has been lost over the years and only now are state historians
trying to relocate these long obscured lands from old deeds and grants.
This work could take several years. Title 62.1, §62.1-3, at first glance,
seems to offer protection to coastal wetlands. Closer reading, however,
shows that to a large extent the Marine Kesources Commission does
not have discretionary power relative to filling or over construction of
private docks and landings for non-commercial use. The Commission
does not have any authority over dredging by a riparian owner.

Altogether, Virginia's statutory provisions are highly inadequate
for protecting the coastal wetlands.

11. North Carolina

The North Carolina Department of Conservation and Develop-
ment shall pass on all excavations and filling proposals. If any state
agency raises an objection to action of the Department, a meeting of
a Review Board composed of the Directors of other state agencies may
be held. The Review Board may affirm, modify, or overrule the action
of the Department of Conservation. Provisions for appeal to the

32 VA. CONST, art. 175.

33 CODE of VA. tit. 62, §62-1.

34 CODE OF VA. tit. 41, §41-81.

Wetlands Legislation

253

courts are provided. No provision is made for taking of any land by
eminent domain.^®

A limited acquisition program is in effect funded by part of the
State Motor Vehicle Tax Fund.^**

]2. South Carolina

The State of South Carolina in seeking to protect its coastal wet-
lands has taken the approach of defining its jurisdiction and own-
ership of the tidelands, submerged lands and waters located in the
coastal region of the State. The State of South Carolina has declared
that it has absolute title to Submerged Lands (the area below the
mean low water mark) in the navigable waters of the State. The State
has prima facie title to Tidelands (marshes) (the area between the
mean high water mark and the mean low water mark), in and ad-
jacent to the navigable waters of the State. The State holds the Tide-
lands, Submerged Lands and Navigable Waters in trust for and sub-
ject to public purposes and rights of navigation, commerce, fishing,
bathing, recreation or enjojmient, and other public and useful pur-
poses, or such other rights as are incident to public waters at common
law, free from obstructions and interference by private persons.

On the basis of this general interpretation of the State’s definition
of tidelands and tideland ownership, there has arisen a legal conflict
concerning which lands are actually owned by the State and held in
public trust for the people, and which lands are actually owned and/or
operated by private individuals. With the exception of isolated cases
brought to settle legal disputes to title of specific acreages of marsh-
lands, there has, up to the present, been little or no effort on the part
of the State to inventory on behalf of the people of South Carolina the
extent of the State’s claim to ownership of lands held in trust. The
continued lack of an applied formula concerning legal interpretation
of ownership of these tidelands has led to an ever increasing number
of conflicts between the State and private individuals. In a pilot proj-
ect in one of South Carolina’s counties, an evaluation of ownership
claims has pointed out that approximately 90 % of tidelands, marshes,
and coastal w'aterw'ays are now claimed by private individuals. These
areas so claimed have in many cases undergone extensive improve-
ments relating to water control and management to provide recre-
ational use and development, channel construction, and dredge and
fill operations for private real estate development. The South Carolina
Water Resources Commission on behalf of the State has collected and
evaluated information relating to the tidelands ownership question.
This and additional information is available to the State for considera-
tion in resolving the tideland’s ownership question.

35 1969 Adv, Legislative Service No, 7, §113-229 (effective January 1, 1970).

N.C. gen. STAT. ch 105, §446.2 (1967 amended 1969).

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§1-357 of the Code of Laws of South Carolina, 1962 embodies the
principles expressed above, It states, in part, that the State Budget and
Control Board is charged with the complete control, regulation and
leasing of all State lands and all public trust properties. For the pur-
pose of leasing oyster and clam rights, it shall use as its agent and
advisor the Wildlife Resources Commission and funds derived from
the Commission shall be used by the Marine Resources Division of the
Wildlife Resources Department for research and management of its
marine resources. No person or agency, public or private, will con-
struct or in any manner place upon or within tidelands or submerged
lands any pier, w'harf, or other structure of any nature or excavate,
dig or in any manner create a dock, ditch, canal, or w’atercourse of
any nature within or upon such tidelands and submerged lands or
place any material upon or change in any manner the natural condi-
tions of such lands without first obtaining a permit from the Board.
Public notice shall be made by the Board of all permits issued.

13. Georgia

An attempt to establish a Coastal Wetland Protective Board
failed to pass the 1969 Georgia General Assembly. Otherwise only the
usual fish and game laws and water pollution laws are germane to
wetland protection.

14. Florida

Chapter 67-393 of the General Acts of 1967

This Act amended the Florida Bulkhead Act of 1957 (Florida
Statutes, §253.12 et seq.).

§253.12. Vests title in tidal lands in the trustees of the Internal
Improvement Trust Fund. The trustees may sell such lands, provided
that they determine that the sale would not adversely affect public
interests, including the preservation of fish and other natural re-
sources. Notice of the sale must be published in the county newspaper.
If objections are filed to the sale, a public hearing must be held. If it
appears as a result of the hearing that the public interest w’ould be ad-
versely affected by the sale, then the trustees must withdraw the land
from sale. A biological and an ecological survey of the land to be sold
must be made in determining whether the sale of the land would ad-
versely affect the public interest.

§253.122. The Board of County Commissioners of each county or
governing body of any municipality, after obtaining a biological and
an ecological survey, may locate and fix bulkhead lines. Any extension
of land outward into the waters of the county is deemed an interfer-
ence with navigation and the conservation of natural resources.

Wetlands Legislation

255

§253.123 The removal of sand, rock or earth from the navigable
waters of the State and the submerged bottoms thereof lying channel-
ward of bulkhead lines is not permitted. Certain exceptions are pro-
vided for.

§253.124. Anyone desiring to add to existing land bordering on
the navigable waters of the State must apply for a permit to do so. A
permit will be issued only if a biological and an ecological survey re-
veal that the public interest will not be adversely affected. The permit
may be revoked for non-compliance with its terms. Anyone who vio-
lates this section is guilty of a misdemeanor and upon conviction shall
be fined a maximum of $500, imprisoned for not more than 6 months,
or both. The trustees have the authority to require the person to re-
move the fill.

Zahel V. Pmellas County Water and Nav. Control AuthoritVy 171
So.2d 376 (1965), on remand 179 So.2d 370, dealt with §§253.122 and
253.124. In this case the local authority lost in its attempt to prevent
owners of bottom land from filling approximately 11.5 acres of land
to be used as a trailer park. The court held that denial of permission
to fill the land amounted to a taking of property without just com-
pensation, because it was not established that granting the permit
would materially and adversely affect the public interest.

Florida Statutes Annotated ch. .175 ( Additional supp. 1966)

This chapter, entitled “Outdoor Recreation and Conservation,”
empowers the trustees of the Internal Improvement Fund to acquire
wetlands and floodlands by purchase, lease-purchase agreements, or
otherwise, with funds from the Land Acquisition Trust Fund.

15. Alabama

The State of Alabama has taken the position, as has South Caro-
lina and other States, that the state does own absolutely the submerged
lands or the area lying below the mean low water mark, which would
include the navigable waters. They have also taken the position that
the State owns the tidelands, the area between the mean high water
mark and the mean low water mark. They are in the process of litiga-
tion, as in all other states. The claimants are now going into court
and are attempting to prove by grants, in their chain of title, that
their private ownership extends to the mean low water mark, because
this tidelands area is now valuable.

The State has entered into litigation involving cases w'here the
tidelands have been filled up by natural accretion. Other cases involve
the action of the water in the reliction, or the washing away, of for-
merly high and dry lands, or formerly tidelands, that are now com-
pletely submerged. Ordinarily where there is accretion or reliction,

1

256

Haueisen

the boundary line of the fast land owner changes with this change in
the mean high water line.

ALA, CODE tit. 8, §§ 232-252 (1932)

The Director of Conservation is vested with authority to develop
State-owned swamplands. These laws are designed to encourage ex-
ploitation.

ALA. CODE tit. 38, 19-122. (1932)

These statutes set forth the right of riparian owners. These au-
thorize and encourage riparian owners to develop lands abutting on
tidelands owned by the State by filling and improving these tideldnds.

The Department of Conservation is authorized to acquire lands
in connection with fish and game programs.

16. Mississippi

MISS. CODE A!\!\. ^^549.7-01 and 7605-09 (1942)

These sections give Port Commissioners or County Port Authori-
ties, respectively, full jurisdiction and control over lands below mean
high tide, including filling and dredging operations. The title to oil
and gas remains in the State. These statutes are designed to encourage
development of the submerged lands.

At the regular session of the 1972 Mississippi Legislature, a bill
was proposed in the House entitled the ^'Mississippi Coastal Wetlands
Protection Act.” This Act recognizes that the coastal wetlands are of
great ecological importance and states that it is necessary to prevent
the deterioration and destruction of these lands in order to preserve
the ecological balance in the coastal area. It declared that the remain-
ing coastal wetlands of the State are in jeopardy of being lost or de-
spoiled by unwise and unplanned activities; that such loss or despolia-
tion will adversely affect, if not entirely eliminate, the value of such
wetlands as sources of nutrients to finfish, Crustacea and shellfish of
significant economic value; that such loss or despoliation will destroy
the ecological system of such wetlands as habitats for plants and ani-
mals of significant economic value and will eliminate or substantially
reduce marine commerce, recreation and aesthetic enjoyment; and
that such loss or despoliation will, in most cases, disturb the natural
ability of tidal wetlands to reduce flood damage and adversely affect
the public health and welfare; that such loss or despoliation will sub-
stantially reduce the capacity of such wetlands to absorb silt and will
thus result in the increased silting of channels and harbor areas to the
detriment of free navigation. Therefore, it was declared to be the
public policy of the State of Mississippi, taking into account varying

Wetlands Legislation

257

scientific, ecological, economic, developmental, recreational and aesthet-
ic values, to preserve the natural state of coastal wetlands and to pre-
vent the despoliation and destruction of these wetlands.

The Act prohibits any regulated activity from being carried on
without a permit. It is the duty of any person proposing to perform
any regulated activity to ascertain whether such work affects wet-
lands.

Any person who violates the provisions of this Act will be civilly
liable to the State for the restoration of the affected wetlands to their
condition prior to such violation, insofar as restoration is possible. In
addition to civil liability under this Act, a violation of this Act is a mis-
demeanor and will be punished by a fine of not less than $500 and not
more than $1,000 or by imprisonment of not more than 30 days, or
both. The Mississippi State Legislature enacted an amended “Missis-
sippi Coastal Wetlands Protection Act” during its February 1973
session.

17. Louisiana

In addition to general water pollution control legislation and leg-
islation for control of fishing, legislation relative to mineral leasing
(oil wells) is the only pertinent legislation in Louisiana.

18. Texas

REV. CIV. ST AT. TEXAS arts. 4051 through 4056a

These statutes give the Texas Parks and Wildlife Commi.s.sion
management control over marl, sand, gravel and shell deposits in the
navigable streams, bays, bayous, and the Gulf of Mexico within the
jurisdiction of the State. Prior to issuing dredging permits, the Com-
mission must consider possible damage to oysters, oyster beds and fish.

CONCLUSIONS

Several states have statutory provisions relating to wetland,
marsh and submerged lands and flood plains. A general categorization
of the legal approaches various states have taken based upon the
above examination of these statutes indicates the following:

(1) legislation which enables a state to acquire wetlands or any
easement, interest or rights therein by the following means:
eminent domain, purchase, exchange, gift, devise, lease, lease
with an option to purchase, payment of unpaid tax liens on
the land.

(2) legislation which prohibits certain activities in wetlands
areas.

258

Haueisen

(a) many statutes provide that a project which involves fill-
ing, dredging, obstructing or altering the course of wa-
ters in wetlands areas may not commence without ob-
taining a permit therefor; any conditions placed upon
the work in the permit must be complied with. Many of
these statutes provide for fines and imprisonment, and
violations are subject to injunction or abatement.

(b) a few' statutes prohibit uses of wetlands inconsistent
with conservation by zoning wetlands for conservation
purposes.

(c) one statute prohibits the use of earth-moving equip-
ment in wetland areas, unless such equipment is regis-
tered with the Department of Water Resources.

(3) the Long Island (New York) Wetlands Act is unique. It pro-
vides that the State may enter into cooperative agreements
with counties to maintain wetlands and may furnish one-half
the cost of maintenance.

(4) legislation not directly related to wetlands, but affecting
flood plains, has been enacted by some states. Such legisla-
tion requires that a county zone its flood plains to prevent
encroachment and consequent damages.

I would call special attention to legislation of the following States :

1. Connecticut — wetlands acquisition

2. Florida — sale of tidal lands

3. Maine — regulation of dredging and filling

4. Massachusetts — regulation of dredging and filling

5. New Hampshire — regulation of dredging and filling

6. New York — cooperative management agreements

7. Rhode Island — wetlands zoning and regulation of dredging
and filling

Recent court decisions in Massachusetts suggest that a legal basis
exists for State regulation of marshland use in instances where private
ownership and use rights exist w’hich may be in conflict with the pub-
lic purpose of protection of marine resources. Commissioner of Natu-
ral Resources v. 5. Volpe and Co., 206 N. E. 2d 666 (Mass. 1965) Reg-
ulation, through the police powers of the State, is the most direct legal
approach to control of use of wetland, marsh and submerged land.
This is short of outright acquisition or control of development rights.
Therefore, the significance of the Massachusetts court decision cannot
be ignored in analyzing other states^ laws and policies which clearly
support the doctrine that protection of marine re.sources is a public
purpose.

Wetlands Legislation

259

A LEGISLATIVE PROPOSAL FOR PROTECTION OF THE

WETLANDS

An act to provide for the orderly preservation and development
of the coastal wetlands; to provide procedures for obtaining permits
to alter wetlands; to provide penalties for violation of this act; and
for related purposes.

Be it enacted by the legislature of the State of :

SECTION 1. This act is to be known as the “Coastal Wetlands
Protection Act'" and may be so cited.

SECTION 2. It is declared that much of the coastal wetlands
of the State of have been lost or despoiled by unregu-

lated dredging, dumping, filling and the like activities, and that the
remaining coastal wetlands of this State are in jeopardy of being lost
or despoiled by these and other activities; that such loss or despolia-
tion will adversely affect, if not entirely eliminate, the value of such
wetlands as sources of nutrients to finfish, Crustacea and shellfish of
significant economic value; that such loss or despoliation will destroy
the ecological system of such wetlands as habitats for plants and ani-
mals of significant economic value and will eliminate or substantially
reduce marine commerce, recreation and aesthetic enjoyment; and
that such loss or despoliation will, in most cases, disturb the natural
ability of tidal wetlands to reduce flooding and adversely affect the
public health and welfare; that such loss or despoliation will substan-
tially reduce the capacity of such wetlands to absorb silt and will thus
result in the increase silting of channels and harbor areas to the det-
riment of free navigation. Therefore, it is declared to be the policy of
this State, taking into account varying scientific, ecological, economic,
developmental, recreational and aesthetic values, to preserve the nat-
ural state of coastal wetlands and to prevent the despoliation and de-
struction thereof.

SECTION 3. For purposes of this act:

(a) “Coastal wetlands,’* “tidal wetlands,” or “wetlands” shall
mean those areas w'hich border on or lie beneath tidal waters, such as
but not limited to banks, bogs, salt marsh, swamps, meadows, flats or
other lowlands subject to tidal action, including those areas now or
formerly connected to tidal waters, and the surface of which is at or
below an elevation of one (1) foot above local extreme high w^ater.

(b) “Regulated activity” means any of the following: draining,
dredging, excavation or removal of soil, mud, sand, gravel, aggregate
of any kind, or rubbish from any w^ctland or the dumping, filling or
depositing thereon of any soil, stones, sand, gravel, mud, aggregate of
any kind, rubbish or similar material either directly or otherwise, and
the erection of structures, driving of pilings, or placing of obstruc-

260

Haueisen

tions, whether or not changing the tidal ebb and flow. Notwithstana-
ing the foregoing, “regulated activity” shall not include the construc-
tion or maintenance of aids to navigation which are authorized by
governmental authority; the accomplishment of emergency decrees of
any duly appointed health officer of a municipality acting to protect
the public health; conservation of soil, vegetation, water, fish, shellfish
and wildlife performed by duly authorized governmental agencies; or
trapping, hunting, fishing and shellfishing where otherwise legally
permitted.

(c) “Dredging" means the removal or displacement by any means
of soil, sand, gravel, shell or other material ; whether of intrinsic value
or not, from wetlands.

(d) “Filling” means either the displacement of waters by the
deposition into wetlands of soil, sand, gravel, shell or other material ;
or the artificial alteration of water levels by physical structures, drain-
age ditches or otherwise.

(e) “Person” means any natural person, partnership, joint stock
company, unincorporated association or society, or the State and any
agency thereof, or municipal or political subdivisions or other corpora-
tion of any character whatsoever,

(f) “Commission” shall mean the Natural Resources and Conser-
vation Commission, the director of said commission or his duly au-
thorized representative.

SECTION 4. No regulated activity shall be conducted upon any
wetland without a permit. Any person proposing to conduct or cause
to be conducted a regulated activity upon any wetland shall file an ap-
plication for a permit with the commission, in such form and with
such information as the commission may prescribe. Such application
shall include a detailed description of the proposed work and a map
showing the area of wetland directly affected, with the location of the
proposed work thereon, together with the names of the owners of
record of adjacent land and known claimants of water rights in or ad-
jacent to the wetland of whom the applicant has notice. The commis-
sion shall cause a copy of such application to be mailed to the chief
administrative officer in the town or towns where the proposed work,
or any part thereof, is located, and to the Director of the State Game
and Fish Commission, the county attorney of the county or counties
in which any part of such proposed work may occur or which may be
affected by such work, the district attorney of any such county or coun-
ties, the boards of supervisors of any such county or counties, and the
Marine Resources and Fisheries Conservation Commi.ssion, No sooner
than thirty (30) days and not later than sixty (60) days from the
receipt of such application, the commission shall hold a public hearing
on such application. The following shall be notified of the hearing by
mail not less than fifteen (16) days prior to the date set for the hear-

Wetlands Legislation

261

ing: all of those persons and agencies who are entitled to receive a
copy of such application in accordance with the terms hereof and all
owners of record of adjacent land and known claimants to water
rights in or adjacent to the wetlands of whom the applicant has notice.
The commission shall cause notice of such hearing to be published at
least once not more than thirty (30) days and not fewer than ten (10)
days before the date set for the hearing in the newspaper having a
general circulation in each county where the proposed work, or any
part thereof, is located. All applications and maps and documents re-
lating thereto shall be open for public inspection at the office of the
commission. At such hearing any person or persons may appear and
be heard.

It shall be the duty of any person proposing to perform any reg-
ulated activity, including the performance of any contract with any
state agency for dredging, sale or removal of shells, gravel, sand or
other such materials, to ascertain whether such work affects wetlands.

SECTION 5. In granting, denying or limiting any permit the
commission shall consider the effect of the proposed work with refer-
ence to the public health and welfare, marine fisheries, shell-fisheries,
wildlife, the protection of life and property from flood, hurricane and
other natural disasters, and the public policy set forth in Section 1 of
thisAct. The commission shall require a bond in an amount and with
surety and satisfactory conditions securing to the state compliance
with the conditions and limitations set forth in the permit. The com-
mission may suspend or revoke a permit if the commission finds that
the applicant has not complied with any of the conditions or limita-
tions set forth in the permit or has exceeded the scope of the work as
set forth in the application. The commission may suspend a permit if
the applicant fails to comply with the terms and conditions set forth
in the application. The commission shall state, upon his record, his
findings and reasons for all actions taken pursuant to this section. The
commission shall cause notice of any order in issuance, denial, revoca-
tion or suspension of a permit to be published in a daily newspaper
having a circulation in the county or counties wherein the wetland lies.

SECTION 6. (a) An appeal may be taken by the applicant or
any person or corporation, municipal corporation or interested com-
munity group other than the applicant who has been aggrieved by such
order from the denial, suspension or revocation of a permit or the is-
suance of a permit or conditional permit within thirty (30) days after
publication of such issuance, denial, suspension or revocation of any
such permit to the court of any county having jurisdiction over the
property which may be affected by any such proposed work authorized
by such permit.

If the court finds that the action appealed from is an unreason-
able exercise of the police power, it may set aside the order. If the

262

Haueisen

court so finds that the action appealed from constitutes the equivalent
of a taking without compensation, and the land so regulated otherwise
meets the interests and objectives of Section 1 of thisAct, it may, at
the election of the commission, (1) set aside the order or (2) proceed
to award damages as provided by Section 9 of thisAct.

(b) Such appeal shall be brought by a complaint in writing, stat-
ing fully the reasons therefor, with a proper citation, signed by a com-
petent authority, and shall be served at least twelve (12) days before
the return date upon the commission and upon all parties having an
interest adverse to the appellant. Such appeals shall be brought to the
next return day of the court after the filing of such appeal. The com-
mission shall forthwith, after service of notice of any appeal, prepare
and file in said court a copy of such portions of the record of the case
from which such appeal has been taken as may appear to the commis-
sion to be pertinent to such appeal, with such additions as may be
claimed by any party of interest to be essential thereto, certified by
the commission. The court, upon such appeal in making its determina-
tions as provided in subsection (a) of this section, shall review, upon
the record so certified, the proceedings of the commission and examine
the question of the legality of the action of the commission and the
propriety of said action. If, upon hearing such appeal, it appears to
the court that any testimony has been improperly excluded by the com-
mission or that the facts disclosed by the record are insufficient for the
equitable disposition of the appeal, it shall refer the case back to the
commission to take such evidence as it may direct and report the same
to the court, with the commission’s finding of fact and conclusions of
law. Such appeal shall have precedence in the order of trial.

SECTION 7. In determining the propriety of issuing permits
for any regulated activity under thisAct, the commission and courts
are to interpret broadly the provisions of thisAct in favor of the pre-
servation of wetlands as opposed to any alteration of the character of
such wetlands, and to favor the best public interest as opposed to pri-
vate or corporate pecuniary interest.

SECTION 8. The Attorney General, district attorney or county
attorney having jurisdiction may institute civil and/or criminal ac-
tions or proceedings against any person believed to be in violation of
thisAct. Such action shall be brought in the court of any county in
which the alleged violation occurs or in which property affected by
such alleged violation is located in the manner of other proceedings.

SECTION 9. Any person who violates the provisions of thisAct
shall be civilly liable to the State for the restoration of the affected
wetland to its condition prior to such violation, insofar as restoration
is possible. The appropriate court shall specify a reasonable time for
the completion of restoration.

In addition to civil liability under thisAct, a violation of thisAct

Wetlands Legislation

263

is a misdemeanor and shall be punished by a fine of not less than Five
Hundred Dollars ($500.00) and not more than One Thousand Dollars
($1,000.00) or by imprisonment of not more than thirty (30) days, or
both.

In the case of continuing violations, each day shall constitute a
separate charge ; however, separate violations under this Act need not
be severed for trial when an identity of parties and location exists.

SECTION 10. If any clause, sentence, paragraph or part of this
Act shall for any reason be adjudged by any court of competent juris-
diction to be invalid, such judgment shall not affect, impair or invali-
date the remainder of thisAct, but shall be confined in its operation to
the clause, sentence, paragraph or part thereof directly involved in
the controversy in which judgment shall have been rendered.

SECTION 11. This Act shall take effect and be in force from and
after its passage.

Gulf Research Reports

Volume 4 Issue 2

January 1973

The Seasonal Occurrence and Abundance of Chaetognatha in Mississippi Sound

Mohammed Saeed Mulkana

Gulf Coast Research Laboratory

Thomas D. Mcllwain

Gulf Coast Research Laboratory

DOI; 10.18785/grr.0402.10

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Recommended Citation

Mulkana, M. S. and T. D. Mcllwain. 1973. The Seasonal Occurrence and Abundance of Chaetognatha in Mississippi Sound. Gulf
Research Reports 4 (2): 264-271.

Retrieved from http:// aquila.usm.edu/gcr/vol4/iss2/ 10

This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean
Research by an authorized administrator of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

THE SEASONAL OCCURRENCE AND ABUNDANCE
OF CHAETOGNATHA [N MISSISSIPPI SOUND

by

MOHAMMED SAEED MULKANA^
and

THOMAS D. McILWAIN
Gulf Coast Research Laboratory
Ocean Springs, Mississippi

INTRODUCTION

The Chaetognatha as “biological indicators" have attracted atten-
tion from various fields of oceanography and fisheries biology because
various species of Chaetognatha are associated with different water
masses, and their distribution and numerical abundance are used to
infer biological productivity and movements of water with their in-
herent temperature, salinity, sediment load pH, and other variables
(Legare and Zoppi 1961, Redfield 1950). The Chaetognatha popula-
tions, therefore, tend to fluctuate following the seasonal changes and
the influx of water masses from different sources.

The present work is a study of seasonal changes in the numerical
abundance of the Chaetognatha species during a yearly cycle in Mis-
sissippi Sound. A comparison is made with the seasonal abundance of
calanoid copepods, an important zooplankton component, for a possi-
ble correlation.

MATERIAL AND METHODS

Samples were collected at one station near the middle of Missis-
sippi Sound at approximately 30° IT N and 80° 45' W. The area re-
ceives open Gulf waters through the Dog Keys Pass and brackish
water from the Bay of Biloxi.

Plankton samples were collected at monthly intervals with
Clarke-Bumpus plankton samplers, equipped with silk bolting nets
:^2 and #10. The nets were towed 15 rain, for each haul, and the
amount of water filtered was recorded by means of calibrated flow
meters attached to the samplers. Both types of nets were used at the
surface and at a depth of 10 ft. which is near the bottom. Occasionally
the efficiency of the plankton nets decreased due to large amounts of
suspended clay particles, phytoplankton blooms, ctenophores and med-
usoid coelenterates.

Data on temperature, salinity and transparency were recorded at
each sampling. Surface temperatures were taken with a precision
mercury thermometer (Fig. IL Water transparency was recorded by

1 Present address, Jackson County Campus, Mississippi Gulf Coast Junior
College, Gautier, Mississippi 39553.

264

Chaetognatha in Mississippi Sound

265

30

Figure 1. Seasonal variations in temperature, salinity and numerical
abundance of chaetognaths in Mississippi Sound from January to Decem-
ber 1965.

266

Mulkana and McIlwain

means of a Secchi disc. Salinity was determined from hydrometer
readings.

The identification of Chaetognalha is difficult when body sizes
overlap, and morphological characters vary with age and size. The
important characters used in classification were size, extent of collar-
ette, number of hooks, number of anterior and posterior teeth, size
and shape of the seminal vesicles, presence or absence of anterior fin,
T.C. ratio (the ratio of posterior fin along the trunk to the posterior
fins along the caudal segment multiplied by 100) the completeness of
the fins, the ventral ganglion and presence or absence of the gut diver-
ticulum (Cf. Grant 1963, Legare and Zoppi 1961, Pierce 1947, and
Thomson 1947).

RESULTS AND DISCUSSIONS

Eight species of Chaetognatha belonging to two genera, Sagitta
and Krohnitta, were identified in the samples (Table 1).

Pierce (1947) reported the occurrence of Sagitta tenuis, S. his-
pida, S, enfUita, S, hdenae and Krohnitta pacifica off the west coast
of Florida. All these species were present in our samples. Sagitta
tennis and S. hispida were most abundant and formed the major part
of the chaetognath populations (Tables 1 and 2). Although both of
these species are considered warm water neritic forms, they also occur
in the Atlantic, but their northern limits have not been determined.
S. tenuis is found as far north as Delaware Bay and S. hispida has
been reported to occur as far north as Cape Hatteras in the Atlantic
(Deevey 1960) . Sagitta enflata and S. bipimctata were comparatively
less abundant, Sagitta enflata is an epiplanktonic, warm-water form,
but occurs north of Cape Hatteras, where it seems to be a stray form
from the Gulf stream. Sagitta enflata is abundant in Florida waters
(Grant 1963) . Sagitta bipimctata is an oceanic form and its presence
indicates mixed oceanic and neritic waters. Grant (personal communi-
cation) has suggested that the Gulf of Mexico chaetognath fauna in-
cludes moat, if not all, the forms found in the central and equatorial
Atlantic. Sagitta helenae has restricted distribution and is limited to
the waters off the Atlantic coast of the United States and the Gulf of
Mexico. Sagitta hexaptera, S. serratodentata and Krohnitta paciflea
were found in fewer number and probably occur as stray forms.

Surface and bottom samples were analyzed separately to see if
there was any variation in these shallow depths. A significant numeri-
cal abundance was observed only during the summer period (Table 2) .
In June and August a greater number of organisms occurred in the
bottom samples, while in July a comparatively higher number was
found in the surface samples. In June and Augu.st transparency was
low (McIlwain 1968), apparently because of pronounced mixing of

S

C

03

3

ec

B

o

««-i

A

X

03

C

5£ -a

5 C

3

“•a

tw fil
® ■«
« .2

§ J
is
i-s

> »ri
(£)

ss^

I

•a w

03 Q

4)

U

cs

c3

XJ

e

s

4»

W

S

O)

3

u

u

o

Krohnitta

paoifica

II 1 1 1 1 v£> 1 1 1 1

Sagitta

serrato-

dentata

II 1 1 1 1 in 1 1 I 1

+i \

B Ss
« oa 01
to 'Ki

II 1 I v£> 1 1 1 1 1 1

y 1

+!>

§ «

Co rQ 4^

I

i 119

i 208

t

Sagitta
he lenae

1 —1 1 cvj I 1 CTi <N 1 1

Sagitta

enflata

295

192

9

1

Sagitta

hispida

1

186

6

673

2089

1687

375

2

1

Sagitta

tenuis

126

5632

796

262

1

51

WATER
FILTERED
IN CU M

coco oococn'-i-<rcM'^mt-<
C\JCMCSlfnCN(N<— ICOlA

DATE

. . . . (U {>^ . 4_l . . .

C^>-lM{>>c:»HW3eU4J>U

tocucvjp^coppDcutjoaj

»nP*^S<J^'-)'-o<!coOZQ

267

268

Mulkana and McIlwain

Table 2.

Variations in Seasonal Abundance of Chaeiognatha Found at Surface
and at Bottom, 10 Feet Deep

NUMBER IN SAMPLES

DATE

Surface

Bottom

Total

January

0

0

0

February

March

2

0

2

April

0

186

186

May

8

0

8

June

763

48

811

July

78

8048

8126

August

2501

440

2941

September

195

500

695

October

4

0

4

November

1

1

2

December

0

0

0

Bay and incoming Gulf waters resulting in a turbid condition in the
Sound, Such a condition would stir up the normally negatively photo-
tropic plankton which would then be distributed in the water column.
In calm waters with greater light penetration chaetognaths, like most
plankton forms, would tend to be close to the bottom, which seems
true of the samples collected in July (Table 2). During the rest of the
year numerical abundance of chaetognaths in the plankton samples
was low, and no conclusions could be made about depth distribution.

SEASONAL VARIATIONS

The chaetognath populations indicated marked fluctuations, with
changes brought about by the climatological factors and surface drift.
The temperature increased from 16.5°C in January to 29.3'^C in Au-
gust (Fig. 1), and decreased rapidly to 12.5®C in December. Salinity
showed a rapid decrease from 25.1 g/kg in January to 16.0 g/kg in
May 1965. A sharp increase was noticed again beginning in May,
reaching a peak of 30.5 g/kg in July. The salinity then decreased,
with a sharp decline in August and September 1965.

According to available information, the surface drift in the

Chaetognatha in Mississippi Sound

269

northern Gulf of Mexico predominates as a northeast flow from the
area east of the Mississippi Delta from March to June and from north-
east to southeast from July to August (Drennan 1963) .

The chaetognath populations showed a sharp rise and decline
following the general pattern of salinity and temperature contours
(Fig. 1). Although with warming climatological conditions the repro-
ductive activities of most organisms ensue and the populations in-
crease, the maximum numerical abundance is not noticed at the high-
est temperature at any one time (Richards 1963, Mulkana 1964). No
studies related to the maximum abundance of chaetognath popula-
tions at maximum salinity are available, but during the summer,
under dry conditions, salinity tends to follow a rising trend and the
greatest abundance at the station studied clearly came at a period of
high temperature and high salinity.

Life studies of Chaetognatha are now well known. Redfield (1940)
has indicated that fluctuations of Chaetognatha will be most marked
if the adult does not survive its first breeding, and less pronounced if
the adult survives through several breeding seasons. The sharp rise
and fall of chaetognath populations in the present study seems to in-
dicate the former condition (Fig. 1). Bigelow and Sears (1939), how-
ever, have pointed out that marked occurrence of S. serratodentata
and periodic occurrence of S. enftata in the Cape Cod area appear to
depend upon environmental conditions rather than the biology of the
reproductive cycle.

Russell (1952), Hardy (1963) and other workers have termed
arrow-worms as “biological indicators" because of their usual pres-
ence in great abundance when productivity of other organisms is at
its maximum. Mcllwain (1968) has also shown that the calanoid cope-
pods which form a large part of the zooplankton component in Missis-
sippi Sound were at their maximum during the greatest abundance of
Chaetognatha (Table 3) .

Other workers attribute the seasonal abundance and fluctuation
to hydrographical factors. Redfield (1940) considered that in regions
which do not develop permanent eddies, populations of chaetognaths
come by immigration from the adjoining areas. In such areas seasonal
fluctuations become greater and little time is required to wash away
the dense populations. The populations are, therefore, pseudo-endemic
and depend upon immigration. The surface drift in Mississippi Sound
water changes periodically (Drennan 1963) and perhaps it contrib-
utes to the pronounced seasonal fluctuations by immigration.

270

Mulkana and McIlwain

Table 3.

Comparison of Seasonal Abundance of Chaetognatha and Calanoid
Copepods from January to December 1965 in Mississippi Sound

DATE

WATER

FILTERED

IN CU M

NUMBER OF
COPEPODS

PER CU M

NUMBER OF
CHAETOGNATHS

PER CU M

January

13

56

0

February

March

43

20

.05

April

28

106

6.7

May

23

83

! .3

June

23

178

35.3

July

31

385

262

August

24

314

123

September

22

53

31.6

October

11

165

.4

November

38

100

.05

December

51

2

0

REGENERATION

During the course of identifications some specimens were found
with a missing head region. Close examination showed the heads were
cut off, hut the cut ends indicated signs of regeneration. Apparently,
some chaetognaths can survive at least for a while after accidental
removal of the head region. One other such observation has been re-
ported (Pierce 1947) .

Plankton samples collected by Mr, J. Y. Christmas and Mr. C. B.
Subrahmanyam in 1966 also showed occasional specimens of S. enflata
and Sagitta sp. with regenerating head regions.

LITERATURE CITED

Bigelow, H. B. and M. Sears. 1939. Studies of the waters of the continental shelf,
Cape Cod to Chesapeake Bay. HI. A volumetric study of the zooplankton.
Mem. Mus. Comp. Zool,, Harvard University. 44:183-379.

Deevey, Georgina B. 1960. The zooplankton of the surface waters of the Delaware
Bay region. Bull. Bingham Oceanogr. Coll., Yale University. 17(2):l-53.
Drennan, K. L. 1963. Surface circulation in the northeastern Gulf of Mexico. Gulf
Coast Res. Lab. Techn. Report No. 1.

Grant, George C. 1963. Investigations of inner continental shelf waters of Lower

Chaetognatha in Mississippi Sound

271

Chesapeake Bay. Part IV. Descriptions of the Chaetognatha and a key to
their identification. Chesapeake Sci. Vol. 4:107-19.

Hardy, Alister. 1963. Continuous plankton records: Contribution towards a plank-
ton atlas of the North Atlantic and the North Sea. Vol, VI, Part VIII:
Chaetognatha, pp. 40-.'51.

Legare, J. E. and E. Zoppi. 1961. Chaetognatha of eastern Venezuelan waters,
with notes on their abundance and distribution. Bol. Insti. Oceanografico. 1:
Numero 1. p. 25,

Mcllwain, T. D. 1968. Seasonal occurrence of the pelagic Copepoda in the Missis-
sippi Sound. Gulf Res. Kept., 2(3) :257-70.

Mulkana, M. S. 1964. The growth and feeding habits of juvenile fishes in two
Rhode Island Estuaries. Gulf Res. Reports. 2:97-168.

Pierce, E. L. 1947. The Chaetognatha of the west coast of Florida. Biol. Bull.
100:206-28.

Redficld, A. C. 1940. Factors determining the distribution of populations of Chae-
tognatha in the Gulf of Main. Biol. Bull. 79:459-87.

Richards, S. W. 1963. The demersal fish populations of Long Island Sound. Bull.
Bingham Oceanogr. Coll,, 18(2) : 1-101.

Russell, F. S. 1952, The relation of plankton research to fisheries hydrography.

Cons. Internat. Explor. de la Mer. 131 :28-33,

Thomson, J. M. 1947. The Chaetognatha of Southeastern Australia. Council Sci.
Ind. Res. (Aust), Bull. 222:1-43.

Gulf Research Reports

Volume 4 Issue 2

January 1973

Studies on the Toxicity of Mirex to the Estuarine Grass Shrimp Palaemonetes pugio

Greg Redmann

Gulf Coast Research Laboratory

DOI: 10.18785/grr.0402.11

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Part of the Marine Biology Commons

Recommended Citation

Redmann, G. 1973. Studies on the Toxicity of Mirex to the Estuarine Grass Shrimp, Palaemonetes pugio. Gulf Research Reports 4 (2) :
272-277.

Retrieved from http:/ / aquila.usm.edu/ gcr/vol4/iss2/ 1 1

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Research by an authorized administrator of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

STUDIES ON THE TOXICITY OF MIREX TO THE
ESTUARINE GRASS SHRIMP, Palaemonetes pugio

by

GREG REDMANN
Gulf Coast Research Laboratory
Ocean Springs, Mississippi

INTRODUCTION

Limited information exists concerning the effects of the poly-
cyclic chlorinated insecticide Mirex (Dodecachlorooctahydro-1, 3, 4-
metheno-2H-cyc]obuta [<?d] pentalene) on non-target organisms. Mi-
rex is the active ingredient in a bait presently being used by the U.S.
Department of Agriculture in the Southeastern states to kill the im-
ported fire ant, Solerwpsis saevissima richteri (Coon and Fleet 1970).
The bait consists of 84.7% corn-cob grits, 15% soybean oil, and 0.3%
Mirex. The approved application rate is 1.25 lbs, per acre per treat-
ment. The use of this persistent pesticide is causing some concern,
particularly in coastal areas dependent on estuarine productivity.

Mirex causes delayed mortality in blue crabs and pink shrimp.
After a 96-hr. exposure to 0.1 parts per million Mirex solution, mor-
tality in these animals reached 100% after a 3-week period (Lowe,
Davison and Wilson 1970). Mahood et al. (1970) surveyed pesticide
residues of blue crabs collected throughout the year in North Caro-
lina, South Carolina, Georgia and Florida and reported that Mirex
was second only to DDT in frequency of occurrence. Mirex exposure
caused mass mortality and alteration of the developmental stages in the
larvae of two crabs, Rhithropanopeus harrisii and Menippe mercemaria
(Bookhout et al. 1972). None exposed to 10 parts per billion Mirex
survived the megalops stage. Mortality approached 100% in crawfish
in 5 days following a 144^hr. exposure to a 1 ppb Mirex solution
(Ludke, Finley and Lusk 1971). Crawfish suffered a high mortality
from granular Mirex through leaching and feeding. These crawfish
accumulated residues 16,860-fold greater than the surrounding water
into which the Mirex had leached.

Lowe et al. (1971) reported the toxicity of Mirex to brown
shrimp, Pemeus aztecus; pink shrimp, Penaeus duoranim\ blue crabs,
Callinectes sapidus; fiddler crabs, Uca pugilator, and grass shrimp,
Palaemonetes pugio. Adult fiddler crabs exposed to levels of Mirex
bait equivalent to a field application had 73% mortality within 2
weeks. Juvenile brown shrimp and blue crabs died after exposure to
one particle of 0,3% Mirex bait per animal. Using 0.8% bait on Palae-
monetes pugio, Lowe et al. (1971) induced 70% mortality in 96 hrs.

272

Mirex Toxicity in Grass Shrimp

273

with one particle of bait per shrimp and 100% mortality in 48 hrs.
with five particles per shrimp.

Bodies of grass shrimp killed from exposure to one granule of
0.3% Mirex bait and then fed to blue crabs caused 100% mortality
to the crabs within 2 weeks (Lowe et al. 1971), High residues in these
crustaceans and mortality in crawfish fed Mirex-exposed Gambima
(Ludke, Finley and Tiusk 1971) suggest the potential harmful side
effects of the fire ant program to natural estuarine communities.

Estuarine grass shrimp (Palaemonetes) are common along the
Gulf and Atlantic coasts (Faxon 1879, Gunter 1950, Williams 1965,
Fleming 1967). These small decapods are found ranging through
freshw'ater areas, brackish estuaries and into inshore saline waters
where they are occasionally the predominant animal (Gunter 1950).

In view of the trophic significance of these shrimp, static system
bioassays were made to determine the sensitivity of Palaemonetes
puf/io to Mirex in water and to 0.16% Mirex bait granules, since the
USDA is continuing the fire ant control program but has plans in the
future to substitute 0.15% Mirex bait for the 0.3% bait presently in
use (J. I. Lowe 1972, personal communication).

MATERIALS AND METHODS

Grass shrimp, P. pugio, were netted in early fall from the dock
area of the Gulf Coast Research Laboratory, Ocean Springs, Missis-
sippi. All grass shrimp were held in a stock tank with 20 ppt salinity
filtered and aerated sea water prior to the bioassays. Mortality was
recorded and dead grass .shrimp removed at 12-hr. intervals. Paralyzed
shrimp were counted as dead, since they would be subject to abnormal
predation in their natural environment. Approximately 10% were
ovigerous ; the average weight was 0.2 g.

Test solutions consisted of either filtered seawater or distilled
water prepared with Rila marine salts, at 20 ppt salinity. Test vessels
were 1-gallon glas.s jars, each containing 1 liter of solution and five
grass shrimp. Technical grade Mirex was prepared in 0.1% acetone
solution and serially diluted for experiments. The Mirex bait used was
obtained from the USDA distribution center in Harrison County, Mis-
sissippi and was produced by Allied Chemical Company, Morristown,
N. J. Solutions were constantly aerated through glass disposable pip-
ettes, All tests were run at room temperatures of 20 ± 1°C,

Experimental procedures included the determination of the toxi-
city to grass shrimp of Mirex leached from bait granules (0.15% ac-
tive ingredient) in water and through exposure to various concentra-
tions of technical Mirex solution.

274

Redmann

EXPERIMENTS AND RESULTS

Toxicity of Technical Grade Mir ex

Samples of five shrimp were placed in each jar for three sets of
nine jars with concentrations of .01 ppm, .1 ppm and 1 ppm Mirex.
Samples of fifteen individuals were exposed for 48, 96, and 144 hrs. at
each concentration and were then transferred to clean sea water. Con-
trols w’ere set up with an equivalent amount of acetone, but had no
mortality throughout the test period. Rostrum-telson length of the
shrimp averaged 26 mm.

Mortality began between 24 and 72 hrs. after the beginning of
exposures to 1 ppm and .01 ppm respectively, with .1 ppm being inter-
mediate in time to beginning of mortality. Exposure for 48 hrs. to .01
ppm caused 40% mortality during a 12-day period, including ex-
posure time. A 100% mortality occurred in 6 days following a 96-hr.
exposure to .1 ppm, and total death prevailed during a 96-hr. ex-
posure to 1 ppm (Table 1). Symptoms exhibited by shrimp prior to
death were irritability, uncoordinated movement, loss of equilibrium
and spasmodic paralysis.

Toxicity from Mirex Granules

Juvenile (10-20 mm rostrum-telson length, avg. 15mm) and
adult (avg. length 26 mm) Palaemonetes pugio were exposed to vary-

Table 1.

Percentage Mortality of Palaemonetes Pugio
(N = 15, avg. length 26 mm) after initiation of exposure to various
concentrations of technical Mirex

Mirex

concen-

tration

Exposure

time

(hours)

%

Paralysis
or death

Days after
exposure
initiation

.01 ppm

48

40%

12

.01 ppm

96

60%

12

.01 ppm

144

83%

12

0 , 1 ppm

48

91%

10

0 . 1 ppm

96

100%

10

0.1 ppm

132

100%

5.5

1 ppm

48

100%

5

1 ppm

96

100%

4

0 ppm

0

0%

12

Mirex Toxicity in Grass Shrimp

275

Table 2.

Percentage Mortality of Grass Shrimp
(juveniles and adults during different exposure times to varying
amounts of Mirex bait — 0.15% active ingredient)

Granules

per

shrimp

Length

(rostrmn-

telson)

Days

of

exposure

Paralysis

or

death

N

1

20-35 mm

10

43%

30

2

10-20 mm*

8

80%

15

2

20-35 mm

8

50%

15

5

10-20 mm*

8

100%

15

5

20-35 mm

8

67%

15

0

10-20 mm*

8

0%

10

0

20-35 mm

10

0%

15

* Juveniles, avg length = 15 mm
Adult avg length = 26 mm

ing amounts of 0.15% active ingredient Mirex bait (Table 2). Jars
were set up as described above, and five to twenty-five granules of
bait (avg. wt. = per granule of 1.2 mg) were added to each jar, equal-
ling one to five granules per individual.

Exposure to one granule caused 43% mortality in 10 days to adult
shrimp, two granules induced 50% mortality in 8 days, and five gran-
ules 67% mortality in 8 days. Juveniles had 80%^ mortality during an
8-day exposure to two granules, and 100% mortality during an 8-day
exposure to five granules of 0.15% Mirex bait. Mortality began 36 hrs.
after beginning of exposure, There was no mortality in the controls.

DISCUSSION AND CONCLUSION

Grass shrimp were found to be fairly sensitive to Mirex through
direct and indirect exposure. Mortality increased with time and Mirex
concentration and appeared to be correlated inversely with animal size.
Exposure to all concentrations (.01 ppm-1 ppm) caused Pa/aewoweies
pugio to suffer continuous delayed mortality for an extended period of
time after the end of the exposure period (Fig. 1). Mortalities from ex-
posure to technical Mirex solutions ranged from 40% in 12 days from a
48-hr. exposure to .01 ppm, to 100% during a 96-hr. exposure to 1 ppm.

Lowe, Davison and Wilson (1970) demonstrated that crabs suf-

276

Redmann

Hxposure Days

period

Figure 1. Mortality in grass shrimp caused by a 48-hour exposure to 1
ppm, 0.1 ppm and 0,01 ppm Mirex solutions. Mortality continued well after
the end of the exposure period.

fered mortality in clean water after a 96-hr. Mirex exposure. This de-
layed continuous mortality was also found in crawfish (Ludke, Finley
and Lusk 1971) and penaeid shrimp (Lowe, Davison and Wilson

1970) .

Extensive mortality occurred in test populations exposed to
0.15% Mirex bait. Exposure to fewer granules decreased the amount
of mortality in a given time period. Though mortality was not as
great or as rapid as in tests using 0.3% bait on P. pugio (Lowe et al.

1971) , it appears that significant toxicity is exhibited by the 0.15%
Mirex fire ant bait in spite of its reduction in insecticide content.

Water contamination by chlorinated hydrocarbon insecticides
through runoff from treated areas in Mississippi (Finley, Ferguson
and Ludke 1970), the resistance of Mirex to degradation, and its ten-
dency to move into water by leaching would indicate the high prob-
ability of contamination in treated watersheds. Upon reaching Gulf
coastal areas (with possible mortality to fresh water shrimp on the
way) Mirex-contaminated water could prove destructive to grass
shrimp populations since their habitat is associated with estuarine
shorelines. The abundance of grass shrimp in coastal shallows (Gun-
ter 1950) indicates their ecological significance.

Mirex-induced mortality in estuarine Palaemonetes could prove
detrimental to the estuarine eco.system. This might affect the Gulf's
commercial fisheries, since estuaries are the nurseries of many eco-
nomically important marine animals and estuarine species make up
about 97,5% of the total commercial fisheries catch of the Gulf States
(Gunter 1967).

Mirex Toxicity in Grass Shrimp

277

In addition, the toxicity of Mirex through trophic levels (Ludkc,
Finley and Lusk 1971, Lowe et al, 1971), its persistence in the tissues
of fish (Lowe, Davison and Wilson 1970) and the usually high toxicity
of chlorinated hydrocarbons to copepods, shrimp and crabs (Butler
1964) should be considered in evaluating the potential importance of
Mirex as a pollutant in the estuarine ecosystem. The present indica-
tions are that it is extremely harmful to crustaceans of all kinds.

ACKNOWLEDGEMENTS

1 would like to thank W. David Burke, Dr. Gordon Gunter and
other personnel of the Gulf Coast Research Laboratory, Ocean
Springs, Mississippi for their aid and assistance. Doctor Gunter iden-
tified the shrimp. The author is currently an undergraduate at Har-
vard University. This work was done at the Gulf Coast Research Lab-
oratory in the summer of 1972.

LITERATURE CITED

Bookhout, C. G., A. J. Wilson, T. W. Duke and J. E. Lowe. 1972. Effects of Mirex
on the larval development of two crabs. Water, Air, and Soil Pollution 1:165-
80.

Butler, Phillip A. 1964. Commercial fishery investigations, Pesticide-wildlife stud-
ies, 1963. U, S. Fish Wildlife Circ. 199:5-28.

Coon, D. W. and R. R. Fleet. 1970. The ant war. Environment 12(10) :18-38.

Faxon, W. 1879. On the development of Palaemonetes vulgaris. Bull. Mus. Comp.
Zook, Harvard College. 5:303-37.

Finley, M. T., D. E- Ferguson and J. L. Ludke. 1970. Possible selective mecha-
nisms in the development of insecticide-resistant fish. Pest. Monit. J. 3:212-
18.

Fleming, L. E. 1967. Use of male external genitalia details as taxonomic charac-
ters in epigean species of Palaenionetes (Decopoda, Palaemonidae) of the
Gulf Coastal Plain of the United States and its coastal waters, M, S. Thesis,
Miss. State Univ., Dept, of Zook

Gunter, G. 1950. Seasonal population changes and distributions as related to
salinity of certain invertebrates of the Texas Coast, including the commercial
shrimp, Pubk Inst. Marine Sck, Univ. of Texas, 1:7-51,

1967. Some relations of estuaries to the fisheries of the Gulf of Mexico.

Estuaries, Amer. Ass. Adv. Sci. 83:621-38.

Lowe, J. L., R. B. Davison and P. D. Wilson. 1970. Effects of Mirex on crabs,
shrimp and fish. BCF Progress Report, 1969. Bur. Com. Fish Circ. 335:22-23.

Lowe, J. I., P. R. Parrish, A. J. Wilson, P. W. Wilson and T. W. Duke. 1971. Ef-
fects of Mirex on selected estuarine organisms. 36th N. Amer. Wild!, and
Natural Resource.^ Conference, pp. 171-86.

Ludke, J. L., M. T. Finley and C. Lusk. 1971. Toxicity of Mirex to crayfish, Pro-
camburis blandingi. Bulk Environ. Contain, and Toxicol. 6:89-96.

Mahood, R. K., M, D. McKenzie, D, P. Middaugh, S. J. Bollar, and J. R. Davis.
1970. A report on the cooperative blue crab study — South Atlantic States.
Georgia Game and Fish Commission,

Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas. Fish. Bulk
65:1-298.

Gulf Research Reports

Volume 4 Issue 2

January 1973

A Study on the Growth Rate of Brown Shrimp (Penaeus aztecus aztecus Ives^ 1891) from
the Coasts of Veracruz and Tamaulipas^ Mexico

Ernesto A. Chavez

Escuda Nacional de Ciencias Biologicas, Mexico

DOI: 10.18785/grr.0402.12

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Marine Biology Commons

Recommended Citation

Chavez, E. A. 1973. A Study on the Growth Rate of Brown Shrimp (Penaeus aztecus aztecus Ives, 1891) from the Coasts ofVeracruz
and Tamaulipas, Mexico. Gulf Research Reports 4 (2): 278-299.

Retrieved from http:// aquila.usm.edu/ gcr/vol4/iss2/ 12

This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean
Research by an authorized administrator of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

A STUDY ON THE GROWTH RATE OF BROWN SHRIMP
(Penaeus aztecus aztecus IVES, 189] ) FROM THE COASTS <.)F
VERACRUZ AND TAMAULIPAS, MEXICO

by

ERNESTO A. CHAVEZ
Departamento de Zoologi'a
Escuela Nacional de Ciencias Biologicas (I.P.N.)

Mexico 17, D.F.

EXTRACTO

Se hizo un estudio biometrico y del crecimiento individual prome-
dio del camaron cafe {Permeus aztecus aztec%is Ives, 1891). Las mues-
tras se tomaron en las empaca doras de Tampico, Mexico desde junio
de 1967 hasta marzo de 1969, y una muestra complementaria hecha
en Julio de 1971. Los resultados del analisis consisten en regresiones
logaritmicas que establecen las relaciones longitud abdominal-longitud
total, y longitud-peso para cada sexo y para toda la muestra nobla-
cional en conjunto. Con los datos mensuales de frecuencia de tarnanos
se determ inar on las clases de edad mediante el uso del papel de pro-
babilidad y posteriormente se hizo el adjuste de los datos resultantes,
en longitud y peso, al modelo de crecimiento de von Bertalanlfy.

INTRODUCTION

The present study was undertaken with the purpose of getting
the proper basis for exploiting brown shrimp resources in a rational
manner, by introducing control regulations of fishing activities. The
growth rate models are necessary for the other parameters of popu-
lation dynamics of the stocks of this species, whose annual catch, Os-
born, Maghan and Drummond (1969), averages 54 million pounds of
tail weight, representing 52% of the total shrimp caught in the Gulf
of Mexico. This shows that Penaeus aztecus aztecus is the most im-
portant of shrimp species exploited in this region. Nevertheless, the
brown shrimp is the least well known of the three most common
species.

The growth rate was determined by fitting the von Bertalanlfy
growth curve to offshore populations, on which according to Cook and
Lindner (1970), there are no published data, although an unsuccess-
ful attempt was made by Klima (1964). Published data are only from
inshore waters (Williams 1955, Loesch 1965, Joyce and St. Amant
et al„ from Cook and Lindner 1970) .

Statistical calculations of length-weight and total length-tail

278

Growth of Penaeiis azteciis aztecus 279

length ratios were required before available raw data could be used
in growth analyses. Tesch (1968), points out that age data, along
with length and weight measurements represent the basis for infor-
mation on growth, composition of stocks, maturity age, life span, mor-
tality and production.

ACKNOWLEDGEMENTS

The writer feels deeply indebted to Miss Rocio Gonzalez for her
aid in the translation of this paper into English. I would like also to
thank Ren6 Marquez, formerly jefe de la Estacion de Biologia Pes-
quera de Tampico, who provided access to the data studied here. The
final revision of the manuscript was made by Dr. Gordon Gunter and
Mr. J. Y. Christmas. I appreciate very much their time invested on
the subject and the corrections made to this paper.

MATERIALS AND METHODS

The original information consists of samplings made in canner-
ies, where offshore shrimp are landed in Tampico, Mexico. The sam-
plings were made by biologist Jesus Macias and an assistant, Mr. Fran-
cisco Robles, both from the Estacion de Biologia Pesquera, in that port.
The frequency of samplings was twice or thrice a week from June 1967
until March 1969 and consisted of abdominal length records, later
grouped to give them monthly representation by means of size-fre-
quency curves with 3 mm intervals for each sex.

The resultant length-frequency curves often show polymodal dis-
tributions, and by using probability paper and Cassie’s method
(1954), it was possible to locate the overlapping values of adjacent
modal groups, such as in Fig. 1 where one analyzed month is shown
as an example. The mean values gotten for each month were considered
as age classes ; in such a way a series of data was obtained, every one
corresponding to the medium size of the respective generations pre-
sented in each monthly sample, averaged in order to fit von Berta-
lanffy's growth model.

To complete the von Bertalanffy growth model it was first neces-
sary to know the corresponding total length of the original tail length
data ; consequently, a new sampling was made in July 1971, consksting
of total length, tail length, and weight data of five hundred individ-
uals of both sexes. Analyses of these data as logarithmic regressions
of the form Y^aX^* gave constants to establish the relationship be-
tween both variables and at the same time gave the necessary infor-
mation on the growd:h constants not only in terms of length but also in
weight.

280

Chavez

Figure 1. Size-frequency of tail length of brown shrimp plotted on prob-
ability paper. The data are from females sampled in January 1969.

RESULTS

Biometric analysis. The biometric study of samples was under-
taken first for each sex separately, and later with both sexes together,
in order to obtain the parameters of the analyzed population as a
whole.

Figure 2 shows the scatter diagram of the total length-tail length
data and the regression line describing the relationship between 251
pairs of values in males. The correlation coefficient was r =0.999 and
the equation with the constants is Y = it corresponds to

an almost straight line which is not rare in the relatively narrow range
of sizes of the sampled shrimps.

In the case of females (Fig. 3), a straighter relationship than in
males can be shown, getting the same value for the correlation factor,
as in the former analysis, of r = 0.999. The values of constants found
are as follows: a = 1.531 and b = 1.001. The number of females ex-
mined was 247.

After the data for each sex were studied, the regression con-
stants for the whole population were calculated. The results are shown
in Fig. 4. They indicate that the parameters found are values lying
between those of the regressions formerly analyzed. They are as fol-
lows: r = 0.998; a = 1.621 ; and b = 0.987.

The length-weight ratios were determined in the way mentioned
before, and for the same reason the results are analogous. The results

Growth of Penaeus aztecus aztecus

281

70 100 130 mm

ABDOMINAL LENGTH

Figure 2. Regression analysis of total length-abdominal length in males
of brown shrimp.

TOTAL LENGTH (mm)

180 H

I40H

100 -

/

60-

' — I — T — r T — f-- -p - , r , —

60 80 100 120 140
ABDOMINAL LENGTH (mm)

Figure 3. Regression analysis of total length-abdominal length in females
of brown shrimp.

282

Figure 4. Regression analysis of total length-abdominal length of brown
shrimp (both sexes combined).

284

Chavez

gotten for each sex are slightly different, but the calculated constants
of the complete samples are intermediate, as follows:

Males: W = 0.000214 r ^ o.968

Females: W “ 0.000010 ^ ^ o.997

Both sexes: W = 0.000023 1^ r = 0.986

It is pertinent to point out that in the first case (Fig, 5), the

value of the exponent seems to be underestimated. According to data
published by Chin (1960), values above three are expected. The ex-
ponent for female length (Fig. 6), and the one of both sexes together
(Fig. 7), are closer to those expected. Growth is supposed to be iso-
metric (at least in this group of crustaceans), and for this reason, the
theoretical expected value for that parameter is three.

One of the reasons the formerly obtained result is not satisfac-
tory is the narrow range of sizes of the male shrimp sampled for
biometric study (in spite of which, the total length-tail length ra-
tio found in this sex seems to be quite admissible). From the data
obtained, their narrow size range limited the range of conclusions, and
consequently the necessity of getting more sample increases, since at
present the probabilities of erroneous results are very large, due also
in part to the limited accuracy of measurement equipment. Also, Cook
and Lindner (1970) point out that old shrimp are proportionally
heavier than young. Nevertheless, the correlation coefficient was quite
high in all cases, particularly in females.

Groivth, For growth studies, the 22 continuous monthly samp-
lings were used. The .samples were taken in a cannery at Tampico.
The monthly data were grouped in 3-mm size classes and represented
by the Petersen method, that is by length-frequency curves. The cum-
ulative frequencies of each month were plotted on probability paper in
the same way as the example in Fig. 1. With this graphic method it
was possible to determine how many age classes were present in each
monthly sample, and at the same time to show some anomalies in these
data, because it was evident that in some months there were more age
classes than in others. Those anomalies are attributable, apparently,
to involuntary sampling errors, which were not completely random.
Nevertheless, they were very useful for inferences about growth of
individuals in the population.

The mean values of monthly age classes found were grouped in a
partially subjective way, for this was the only way possible to elimi-
nate some of the sources of error included, since the sampling time,
the intrinsic variations of population, and those due to ecological fac-
tors, such as temperature, that modify not only growth rate, but the
velocity of physiological processes in general and their effects are also
reflected among others in the growth.

The selection of age classes by this method has the convenience

WEIGHT (g)

TOTAL LENGTH (mm)

Figure 6. Regression analysis of length-weight in females of brown
shrimp.

WEIGHT (g)

Growth of Penaeiis aztecm aztemis

287

Figure 7. Regression analysis of length-weight of brown shrimp (both
sexes combined).

288

Chavez

of reducing the variations in growth caused by the presence of in-
termediate age classes, by seasonal changes and by other ecological
factors, from which a series of size classes result whose values rep-
resent the average growth rate of the analyzed population. For this
reason in Fig. 8 by following the monthly grow'th of each age class,
identified by letters, if some discrepancies are noticed, it is because it
was necessary to make some adjustments to minimize extreme varia-
tions.

The mean values of selected age classes are indicated in Table 1.
With respective averages obtained for each sex a Ford-Walford plot
was done (Fig. 9), graphic method to estimate the average maximum
lengrth of individuals in the population studied, finding L cc = 117 and
150 mm of abdominal length for males and females. With all these data,
those of age classes and Loo, transformations were made to determine
their corresponding values in total length; this was done by using the
regression formulas previously calculated, and with them the von Ber-
talanffy grow'th equation was fitted, as follows:

J — Lee [1 —

where :

1 = Length at age t in mm

L 00 = Average maximum length

k = Constant, proportional to catabolic rate
t = Age, in months in this case

to = Theoretical adjustment parameter, which expresses the
age when the length is zero.

For additional details about this model, refer to Beverton and
Holt (1957), Ricker (1958), Cushing (1968), Gulland (1969) and
similar texts.

On following the curve as far as its origin, it was found to be
reasonably well fitting in very small sizes, and it could be seen on the
average, that the smallest age class fitted should be 4 months old, nine
the elder in males, and ten in females.

The constants found in this case are shown in the following
formulas, and the curves that they describe are also shown in Fig. 10.

Males: 1 z= 178.1 [1 — e 1 - 4 - 0 . 2388 )]

Females: 1 = 236 [1 — e - 0162 ( 1 + 0 . 759 )]

With the growth model it is possible to express growrth not only
in terms of length but also in weight. This can be accomplished by
transforming the values of Leo into its corresponding weight and rais-
ing the rest of the member to cube; the constants k and t„ do not
modify. The result is a sigmoid curve whose inflection point is found
at the third part of the value of asymptotic w^eight or W^c of that
population. The transformation of length into weight was done with

Cfc )

Growth of Penaens aztemis aztecus

289

MALES

.hAN\

C H 0 T H (tTvfn>

L C N C I H ifTsm)

Figure 8. Monthly tail length-frequency curves of males and females of
brown shrimp on which all the sampling period is represented. Mean
length values of successive generations are identified by letters. The
corresponding total length scale (Lj) is also shown below.

290

Chavez

Table 1.

Monthly age classes determined from the analysis of frequencies
made with probability paper. They are represented as abdominal
length, in ram. Each generation is identified by letters. (These values
are also shown in Figure 12)

Generation

Mean size values of monthly age classes

I

II

III

IV

V

VI

A

103.5

108.0

B

88.5

94.5

94.5

99.0

C

82.0

87.0

91.5

96.0

105.0

D

93.0

E

81.0

91.5

94.5

F

81.0

88.0

96.0

G

79.0

87.0

96.0

H

79.0

87.0

94.5

105.0

I

79.0

87.0

91.5

96.0

100.0

J

78.0

K

77.0

88.0

L

89.5

95.5

106.5

M

78.0

88.5

97.5

102.0

N

87.0

94.5

103.5

0

78.0

87.0

93.0

96.0

103.5

P

85.5

Q

87.0

96.0

102.0

R

87.0

93.0

S

85.0

Average

79.2

87.3

94.1

98.4

102.8

108.0

the regressions previously determined. But in the case of males and
with the considerations formerly made, it seemed pertinent to take
the results of males and females computed together instead of those
of males only in order to reduce the risk of working with wrong data,
mainly when there is the antecedent of no marked differences between
the sexes (Chin 1960). Therefore, the results of growth in weight are
shown in the following expressions :

Males: w = 46 [1 — e-o-2567(t+o.238a)]3
Females: w =:^113 [1 — e—^. 162 ( 1 + 0 . 750 )] s

The expressions seem to be similar to those of the length growth.

Growth of Penaev.s aztecus aztecus

291

Figure 9. Ford-Walford graph of abdominal length in mm at age t
against length at age t+1 for males and females of brown shrimp.

Nevertheless, the differences existing between Loo of each sex are
equivalent to quite distinct weights of specimens of the same age but
different sex, variations that increase with age. This can be seen in
Fig, 11 where the growth in weight is represented by sex.

The sizes and weights of each monthly age class calculated by
•means of the von Bertalanffy growth model are shown in Table 2,
whose variations explain well the differences in growth found by
Williams (1955), Loesch (1965), and St. Amant et al, and Joyce
(both after Cook and Lindner 1970) upon juvenile brown shrimp
populations, somewhat larger growth rates than those found during
the present analysis for similar size ranges.

Finally, bearing in mind that only global data are frequently
available, for example, the Gulf Coast Shrimp Data, in which it is not

292

Chavez

Figure 10. Von Bertalanffy growth curves, in length, found for each sex
of brown shrimp. Observed values of age classes are shown by small
circles.

possible to separate the differences due to sex, it was decided to cal-
culate growth parameters of the population as a whole, which was
done in the following way : with the values of Leo and W co determined
for each sex, new average values were obtained and were considered
as Loc and W oc of the population, and the age classes were those de-
termined by making an average of the resulting pairs of values form-
erly analyzed after the growth rate was considered for each sex sep-
arately. Therefore, the latest obtained results are as follows :

Loo = 207 mm Woo = 70 g
k = 0.1904 to = —0.872

The graphic representation of individual growth rate of the Pe-
naeiis aztecus aztecus population studied is shown in Fig. 12, and in
Table 3 the length and weight values for each month of age are also
indicated.

294

Chavez

Table 2.

Growth rate in length and weight of Penaens aztecus aztecus Ives
calculated for each sex and month during the first
fourteen months of age

MALES

Age

(Months)

Length

(mm)

Increase

(mm)

Weight

(g)

Increase

1

48.5

37.9

0.9

0.9

2

75.7

27.2

3.7

2.8

3

100.6

24.9

8.3

4.6

4

118.3

17.7

13.4

5,1

5

131.8

13.5

18.6

5.2

6

142.0

10.2

23.4

4.8

7

150.4

8.4

27.7

4.3

8

156.6

6.2

31.3

3.6

9

161.4

4.8

34.3

3.0

10

165.3

36.7

2.4

11

168.2

38.7

2.0

12

170.4

40.3

1.6

13

172.1

1.7

41.5

1.2

14

173.5

1.4

42.5

1.0

FEMALES

1

58.3

30.9

1.7

1.5

2

85.0

26.7

5.3

3.6

3

107.6

22.6

10.7

5.4

4

127,0

19.4

17.5

6.8

5

143.3

16.3

25.2

7.7

6

157.1

13.8

33.3

8.1

7

169.0

1 11.9

41.4

8.1

8

179.0

10.0

49.2

7.8

9

188,0

9.0

56.6

7.4

10

195.0

7.0

63.4

6.8

11

200.9

5.9

69.7

6.3

12

205,0

4.1

75.3

5.6

13

210.6

5.6

80.3

5.0

14

214.4

3.8

84.7

4.4

Growth of Penaeus azteciis aztecun

295

AGE, MONTHS

Figure 12. V^on Bertalanffy growth curves found for all brown shrimp
population sampled (both sexes combined). The curves show the longi-
tudinal and ponderal average growth of individuals.

DISCUSSION OF RESULTS AND CONCLUSIONS

As a corollary of biometric study of the analyzed population, it
is concluded that the total length-abdominal length ratio shows a
mostly straight relationship in both sexes, and that the observed dif-
ferences in each sex are very slight. For this reason, it is supposed
that the calculated population formula is representative of such a
transformation, a characteristic that on the other hand means that
the shrimp growth is isometric respectively to these two factors. The
calculated, relationship was made on a basis of the need of knowing
the total length of specimens formerly sampled, and it was necessary
to make a reference calculated in terms of total lengths of the shrimp.

Concerning the length-weight relationship, it is pertinent to
point out that the exponent of length usually is a value which fluctu-
ates about three, and in this case described an isometric growth^ char-
acterized from Ricker (1958), because the specific gravity and the
body form remain constant, regardless of the size of the organism.
Former experiences with shrimp (Hall 1962, Butler 1970, Chin 1960,

296

Chavez

Table 3.

Monthly growth rate in length and weight of brown shrimp calcu-
lated for all population samples (both sexes combined)

Age

(Months)

Length

(mm)

Increase

(ram)

Weight

(g)

Increase

(g)

1

61.8

i

30.1

1.9

1.7

2

87.0

25.2

; 5.3

3.4

3

105.5

18.5

9.9

4.6

4

121.1

15.6

15.5

5.6

5

139.6

18.5

21.3

5.8

6

151,1

11.5

27.2

5.9

7

160.8

9.7

32.8

5.6

8

168.5

7.7

37.9

5.1

9

175.2

6.7

42.6

4.7

10

181.0

5.8

46.7

4.1

11

185.4

4.4

50.3

3.6

12

189.1

3.7

53.4

3.1

13

192.2

3.1

56.1

2.7

14

195.0

2.8

58.3

2.2

Kutkuhn 1966, George 1970a and 1970b, Nikolic' and Garcia 1970,
Cruz-Morejon and Cadima 1970, Angelescu and Boschi 1959, and
Chavez and Rodriguez-de la Cruz (1971), suggest that at least in
this group, the growth shows a remarkable tendency to isometry,
where the calculated exponents fluctuate around three, ranging from
2.65 to 3.25. These variations are probably due to insufficient data,
small number of samples, to variations in condition coefficient of
shrimp, or to the influence of other physiologically responsible factors.

Gulland (1969) points out that there is a wide number of growth
equations, but none is entirely satisfying in all possible situations.
Dickie (from Tesch 1968), asserts that growth curves are valuable
for descriptive purposes, but the biological interpretation of models
and their parameters still present great diflficulties. The von Bcrta-
lanffy growd^h model in the author's opinion, has been shown to be
quite satisfactory; besides, it should be kept in mind that this model
is the best known and the most widely used in production studies on
species of economic value ; it satisfies reasonably the two most impor-
tant criteria : it fits most of the observed data on growth, and can be
readily incorporated into stock assessment models.

The growth analysis developed in the present paper seems to offer

Growth of Penaens aztecus azteciis

297

a good outlook for further application to the penaeid shrimp group.
This group presents serious difficulties for the determination of age
and growth of the species belonging to it, because of its variability
due to environmental changes. There is also the impossibility of re-
ferring the age classes to growing marks to check the analytical infer-
ences made from the information acquired with the samples.

With the comparative study of size-frequency curves, it is possi-
ble to figure out that in analyzed samples there is a small percentage
of shrimp whose age ranges are from 2 to 13 months old in males,
approximately, with lengths ranging from 93 to 172 mm in each case;
in females, all seems to show that age classes ranging from 2 to 15
months old are present (the number of extreme size classes is also
small), whose lengths lie between 91 and 216 mm. It is supposed that
the presence of larger shrimp is a random occurrence rather than the
result of an ecological factor. These large sizes were found from June
through August in both sexes, from January until March in males,
and from November to March in females. The smaller shrimp, approx-
imately 2 months old, were observed quite well chronologically located.
In spite of the fact that Gulf of Mexico shrimps viiTually repro-
duce the year-round, the presence of the smallest shrimp is neces-
sarily interpreted as a maximum of reproduction existing at least 2
months before their recruitment. If recruitment of such tiny shrimp
occurs mainly from June through August, it is supposed-that they be-
long to a generation born from March through May (it is necessary
to add 15 days to the apparent age, because of the larval stage dura-
tion). In part, this confirms the opinion of Kutkuhn (1962) about the
increased spawning activity during March- April and September-
October.

The results obtained by the use of probability paper to determine
the size classes suggests more objectivity than the use of the size-
frequency curves only (Chavez and Rodriguez-de la Cruz 1971), es-
pecially when the number of samples is small. For instance, either
one of these methods may be profitable, but the accuracy of the results
is determined by the experience of the analyst.

It is necessary to keep in mind that the present study was under-
taken with the idea of determining the average individual growth
rate, regardless of the seasonal valuations, due to the fluctuating
changes of ecological factors determining the habitat of P. azteciis
azteciis populations. The study was made in that way because on in-
corporating growth parameters into stock assessment models, extreme
variations capable of modifying the results are discarded.

SUMMARY

A study of brown shrimp {Penaeus aztecus aztecus Ives, 1891)
was undertaken. The data were obtained from samplings made twice

298

Chavez

or thrice a week in Tampico (Mexico) canneries, during the period
from June 1967 to March 1969. The data consist of abdominal length
records of 20,003 specimens of both sexes, 6,879 males and 13,124
females. An additional sampling of 500 specimens was made in July
1971 in which total length, tail length and weight were recorded.
These data were analyzed as logarilhmical regressions establishing
the corresponding relationships, of which the formulas are as follows :

Total length (Y) — abdominal length (X) ratio:

Males: Y = 2.05 r = 0.999

Females: Y = 1.531 r = 0.999

Both sexes: Y = 1.621 ^ ^ o.998

Weight (W) — length (1) ratio:

Males: W = 0.000214 r = 0.968

Females: W = 0.000010 r = 0.997

Both sexes: W 0.000023 j, ^ o.986

Tail length data were grouped and represented as monthly
length-frequency curves, to which mean values of age classes found
in each month were incorporated. These values were obtained after
analyzing each monthly group of data on probability paper, Once the
mean values of age classes were obtained, a Ford-Walford plot was
made and a von Bertalanffy growth curve was fitted. The constants
are a.s follows:

Males :

Loo

= 178.1 mm

Woo

^ 46 g

k

= 0.2567

to

^ —0.2388

Females :

Loc

= 236 mm

Woo

= 113 g

k

= 0.162

to

^ —0.759

Both sexes;

Loo

= 207 mm

Woo

= 70 g

k

= 0.1904

to

= —0.872

LITERATURE CITED

Angelescu, V. and E. E. Boschi. 1959. Estudio biologico pesquero del langoatino
del Mar del Plata en conexion con la operacion nivel medio. Servicio de Hi-
drografla Naval, Argentina, 1017:1-135.

Beverton, R. and S. Holt. 1957. On the dynamics of exploited fish populations.
Min. Agr. and Fish., Fish. Invest. Lond., Series 2, 10.

Butler, T. H. 1970. Synopsis of biological data on the prawn Pandalus platyceros
Brandt, 1851. FAO Fish. Rep. (57)4:1280-315.

Cassie, R, M. 19.54. Some uses of probability paper in the analysis of size fre-
quency distributions. Australian J. Marine and Freshwater Res., 5:513-22.

Chavez, E. A. and M. C. Rodriguez-de la Cruz. 1971. Estudio sobre el crecimiento

Growth of Penaeus azteciis azteciis

299

del camaron cafe (Penaeus callfondensis Holmes) del Golfo de California.
Rev. Soc. Mex. Hist. Nat. 32:111-27.

Chin, E. 1.960. The bait shrimp fishery of Galveston Bay, Texas. Trans. Amer.
Fish. Soc., 89(2) :135-4l.

Cook, H. L. and M. J. Lindner. 1970. Synopsis of biological data on the brown
shrimp Penaeus aztecus aztecus lyes, 1891. FAQ Fish, Rep. 57(4): 1471-97.

Cruz-Morejon and E. Cadima. 1970. Relaciones entre largos y pesos de camarones
capturados en la plataforma cubana. FAO Fish. Rep. (57)2:539-48.

Cushing, D. H. 1968. Fisheries biology. The Uniyersity of Wisconsin Press.

George, M. J. 1970a. Synopsis of biological data on the penaeid prawn MeAuve-
naeus monoceros (Fabricius 1798). FAO Fish. Rep. (57)4:1539—57.

1970b. Synopsis of biological data on the penaeid prawn Metapenaeus

brevicornis (II. Milne Edwards, 1837). FAO Fish. Rep. (57)4:1559-73.

Gulland, J. A. 1969. Manual of methods for fish stock assessment. Part I. Fish
population analysis. FAO Man. Fish, Sci.

Hall, D, N, F. 1962. Obseryations on the taxonomy and biology of some Indo-
west Pacific Penaeidae (Crustacea, Decapoda). Fish. Pubis. Colon. Off., 17:-
1-229.

Klima, E. F. 1964. Mark recapture experiments with brown and white shrimp in
the northern Gulf of Mexico. Proc. Gulf. Carib. Fish. Inst, 16:52-64,

Kutkuhn, J, H. 1962. Gulf of Mexico commercial shrimp populations, trends and
characteristics, 1956-59. U. S. Fish Wildl. Sery., Fishery Bull. 62(212) :343-
402.

1966. Dynamics of a penaeid .shrimp population and management im-
plications. U, S. Fish. Wildl. Sery., Fishery Bull. 65(2) : 313-38.

Loesch, H. 1965. Distribution and growth of penaeid shrimp in Mobile Bay, Ala-
bama, Publ. Inst. Mar. Sci. (Texas), 10:41-58.

Nikolic’, H. and R. Garcia. 1970. Diez viajes de exploraci6n con el barco Camaron
I en la plataforma sudoriental de Cuba. FAO Fish. Rep. (57)2:571-87.

Osborn, K. W., B. W. Maghan, and S, B. Drummond. 1969. Gulf of Mexico shrimp
atlas. U. S. Dept. Int. Bur Com. Fish. Circular 312:1-20.

Ricker, W. E, 1958. Handbook of computations for biological statistics of fish
populations. Bur. Fish. Res. Bd. Canada (119) : 1-300.

Tesch, F. W. 1968. Age and growth. In Methods for assessment of fish production
in fresh waters. W. E. Ricker (Ed.). I. B. P. Handbook No. 3:93-123. Black-
well.

Williams, A. B. 1955. A contribution to the life histories of commercial shrimps
(Penaeidae) in North Carolina. Bull. Mar. Sci, Gulf, Caribb. 5:116-46.

Gulf Research Reports

Volume 4 Issue 2

January 1973

Nutritional Components of the Standing Plankton Crop in Mississippi Sound

Mohammed Saeed Mulkana

Mississippi Gulf Coast Junior College

Walter Abbott

Gulf Coast Research Laboratory

DOI: 10.18785/grr.0402.13

Follow this and additional works at: http:/ / aquila.usm.edu/ gcr

Part of the Marine Biology Commons

Recommended Citation

Mulkana, M. S. and W. Abbott. 1973. Nutritional Components of the Standing Plankton Crop in Mississippi Sound. Gulf Research
Reports 4 (2): 300-317.

Retrieved from http:// aquila.usm.edu/gcr/vol4/iss2/ 13

This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean
Research by an authorized administrator of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

NUTRITIONAL COMPONENTS OF THE STANDING
PLANKTON CROP IN MISSISSIPPI SOUND^

by

MOHAMMED SAEED MULKANA^
and

WALTER ABBOTT
Gulf Coast Research Laboratory
Ocean Springs, Mississippi 39564

ABSTRACT

A study on seasonal changes in the nutritional components of
standing plankton biomass was made from 20 April 1965 to 6 Septem-
ber 1966. Plankton were separated into net plankton and nannoplank-
ton fractions. Nannoplankton'* standing biomass exceeded by 72 times
that of net plankton.

Although protein and carbohydrate levels were higher in net
plankton on a unit weight basis, total nutrients available from nan-
noplankton were substantially higher. Some seasonal trends were ap-
parent as changes in the standing biomass in net plankton. Nanno-
plankton exhibited no seasonal trends. Lipid and caloric values of net
plankton showed seasonal trends similar to those shown by dry
weight.

INTRODUCTION

Early studies on marine plankton were directed primarily toward
species composition, geographical distribution and seasonal succes-
sions. The importance of plankton to fisheries led to studies of plank-
ton biomass, with considerable recent emphasis on plankton produc-
tion as indicated by primary photosynthetic productivity. There are
relatively few data, however, on specific nutritional components —
carbohydrate, protein, and lipid — available from plankton and on
quantitative seasonal changes in these components. Furthermore, al-
though nannoplankton comprises the major fraction of the total plank-
ton and exceeds the net plankton manyfold (Banse 1964, Pomeroy
and Johannes 1966), those nutritional component data available relate
only to net plankton.

1 This work was supported in part by National Science Foundation Grant No.
BG-3452, and is extracted from a Ph.D. Thesis submitted to Department of Zool-
ogy, Mississippi State University.

2 Jackson County Campus, Miss. Gulf Coast Jr. College, Gautier, Mississippi
39563.

3 It should be noted that the authors did not Separate the nannoplankton from
what has been called the nannodetritus and both components are included in the
word nannoplankton, as used in this paper. Ed.

300

Mississippi Plankton Crop

301

Mississippi Sound is regarded as a highly productive body of
water (Gunter 1962, 1963, Christmas, Gunter, and Musgrave 1966)
because of the influence of nutrient rich waters from Mississippi
River (Riley 1937). This report is concerned with measurements of
net plankton and nannoplankton dry weight, protein, and carbohy-
drate, and with lipid and caloric content of net plankton during a yearly
cycle in Mississippi Sound.

MATERIALS AND METHODS

From April 20, 1965 to September 6, 1966, plankton was col-
lected as near monthly as logistics and weather permitted from 6
sampling stations along two transects across Mississippi Sound —
from Marsh Point to Horn Island (Fig. 1) and from Ship Island to
Biloxi Beach (Fig. 1). The order of running the transects was sys-
tematically reversed from cruise to cruise to avoid sampling bias, but
on each cruise one transect was w'orked by day and the other by night.

Composite net plankton samples were taken by pumping water
from surface, mid-depth, and near bottom into a No. 20 plankton net

Figure 1. Map of Mississippi Sound showing sampling stations and
depths. GCRL represents the site of the Gulf Coast Research Laboratory.

302

Mulkana and Abbott

(76 ja mesh size) submerged in a water-filled container. The water
was pumped at the rate of 190-240 liters/min. for 30 min. with a pre-
calibrated pump. Leong (1967) has discussed plankton sampling by
pumping in oceanographic .situations. Our conditions were less severe
because we used 5 cm pump instead of 1,27 pump and pumped water
from a maximum depth of 8 meters rather than from 120 meters. As a
result there should be less churning effect and less fragmentation of
the planktonic organisms. A 19-liter aliquot of net filtrate was taken
at each station as a nannoplankton sample. Both this aliquot and the
net plankton sample were kepi chilled by ice until return to the labor-
atory.

The nannoplankton samples were flocculated with potassium alum
(KAl(SOi).,) to precipitate all suspended matter, plankton centri-
fuges having proved unsatisfactory for this purpose. The floes \vere
dissolved with just sufficient 3N hydrochloric acid and centrifuged at
1500 HPM for 15 minutes. Both net and nannoplankton concentrates
were freed from water and salt by filtration on 0.8 ju, Millipore type
A A filters with a rapid wash with isotonic ammonium formate (Par-
sons, Stephens and Strickland 1961). Humphrey and Wootton (1966)
have shown by pigment studies with Gijmnodinium, Nnnnochloris,
and sea water samples that delicate, small phytoplankton are retained
as well by this Millipore filter as by those of lesser pore size. Samples
were dried to constant weight at 105° C (Curl 1962) against a Milli-
pore filter blank, scraped from the filter, and homogenized by grind-
ing.

For measurement of protein content, portions of plankton homo-
genates, and albumin, used as standard, were hydrolyzed in a sealed
vial with 6N HCl for 12 hours at 120“ C (Welcher 1963). Hydroly-
sates were filtered, dried on a steam bath, and diluted to a standard
volume. Amino acid content of plankton fractions and albumin hydro-
lysates was determined colorimetrically a.s leucine equivalents using
a modification of the ninhydrin technique of Landua and Awapara
(1949), The albumin equivalent of leucine, determined experimental-
ly, was 6.97 /xmol leucine/mg albumin.

Plankton carbohydrates were determined by hydrolyzing por-
tions of homogenates, and oyster glycogen, used as standard, with 3N
HCl for 1^2 hours at 95° C. Hydrolysates were dried and glucose
content was determined colorimetrically by the method of Folin and
Wu (Department of the Army 1951 ) .

Nannoplankton samples were too .small for routine lipid analysis.
Lipid from net plankton wa.s extracted with chloroform-methanol-
water (2:2:1) in a screw-capped Waring blender jar (Bligh and Dyer
1959). The homogenates were filtered and allowed to .stand in separa-
tory funnels with Teflon stopcocks until phases separated. The chloro-
form layer was removed, washed with chloroform-methanol-water

Mississippi Plankton Crop

303

(3:48:47) (Folch, Lees and Stanley 1957), and dried with infrared
lamp. Total lipid in the aliquots was determined spectrophotometrically
by the method of Snyder and Stephens (1959) using triolein as stan-
dard.

Caloric measurements on net plankton were made according to
Parr Manual No. 122 (1951) in a Parr Peroxide Bomb Calorimeter,
Model 1401. Dried net plankton samples were ground to pass a 60
mesh screen. Particle size is important, because combustion reactions
occur in a few seconds and large particles, if present, may not burn
completely. Benzoic acid was used as combustion aid.

Ash content of plankton samples w’as determined by hot nitric
acid digestion of the dried homogenate followed by ignition to con-
stant weight in a muffle furnace at 550*^ C.

Salinity and temperature were determined m situ at each station
by means of a Beckman Induction Salinometer. Figure 2 shows mean
surface water temperatures during the course of this study.

All data for biomass, nutrients, and caloric values have been com-
puted for a water column of 1 sq m area.

4

—I 1 1— T 1 ( 1 r 1 r — ^ — I 1 1 1 I 1 1 1 r

AMJ JASONDJ FMAMJJASO
1965 ' --1966

Figure 2. Mean surface temperature at six sampling stations from 20
April 1965 to 6 September 1966.

304

Mulkana and Abbott

RESULTS

Dry Weight: The net plankton dry weight values of 637 mg/sq m
and 452 mg/sq m (Fig. 3, Table 1) observed in April and May 1965,

o

X

E NANNOPLANKTON

NET PLANKTON

AMJ J ASONDJ FMAMJ JASO
-1965 1966

Figure 3. Seasonal variation in mean values of dry biomass of net plank-
ton and nannoplankton.

respectively, derived from a plankton increase because of vernal
warming in Mississippi Sound (Thomas and Simmons 1960), The
peak of 401 mg/sq m in August 1965, appeared to be due to increased
herbivore populations. The magnitude of spring and summer peaks in
1966 was considerably higher than peaks in 1965. Deevey (1960) indi-
cated that the zooplankton seasonal cycle in nearshore waters may be
extremely variable from one year to the next. On several occasions in
summer 1966 concentrations of a blue-green alga, Trichodesmium sp.,
were observed in surface waters. The algal abundance would contrib-
ute markedly to the upswing in standing biomass. Beginning in late
summer 1965, a gradual decrease in standing dry weight ensued, with
a minimum of 43 mg/sq m in January 1966. A similar trend was
noted again in 1966, when average values showed a sharp decline
during September, decreasing from 1560 mg/sq m to 681 mg/sq m
(Table 1).

Table 1

Summary of Mean Values for Dry Weight, Nutrients, Ash Content and Caloric Values in the Standing
Biomass of Net Plankton. Means have been Computed from Results at Six Stations

Date

Dry Weight
mg/sq m

Protein
mg/sq m
albumin

Carbohydrate
mg/sq m
glycogen

Lipid
mg/sq m
triolein

Ash

mg/sq m

Total nutrient
and ash as per-
cent dry weight

kcal/
sq m

1965

20 April

637

158

29

15

217

65

1.7

4 May

228

62

9

8

113

84

0.5

11 May

452

100

7

15

230

78

0.9

25 June

284

96

9

11

147

92

0.6

31 July

282

63

6

8

171

88

1.1

14 August

401

104

7

11

208

82

0.8

26 August

337

66

6

16

130

65

1.1

7 October

285

73

4

4

166

87

0.6

4 November

225

47

4

5

129

82

0.5

20 December

194

17

10

1

no

71

1966

23 January

43

15

1

1

22

90

8 April

508

92

24

9

175

59

1.3

28 May

898

172

15

11

463

74

2.2

17 June

444

76

7

16

219

72

1.0

1 July

739

217

37

12

389

88

1.9

15 July

581

197

34

16

276

90

1.1

19 July

740

271

20

19

402

96

1.8

13 August

566

185

32

14

292

92

1.2

26 August

1560

470

32

48

778

85

3.3

6 September

681

235

16

34

310

87

1.7

Grand Mean

504

136

15

14

247

81

1.3

Mississippi Plankton Crop 305

306

Mulkana and Abbott

The nannoplankton dry weight exceeded the net plankton dry
weight manyfold, and showed a mean ratio to net plankton of 140:1,
with the maximum in winter and the minimum in summer (Tables 1,
2 and 3). Yentsch and Ryther (1959), on the basis of chlorophyll a
values, and Johannes (1964), on the basis of dry biomass, pointed out
that nannoplankton comprised the major fraction of the total plank-
ton.

The nannoplankton crop showed no definite seasonal trend, bio-
mass peaks at various stations appearing in spring, fall, midsummer,
and midwinter (Fig. 3). Mean dry weight values ranged from 7.50
g/sq m to 21.8 g/sq m in spring and from 7.9 g/sq m to 25.4 g/sq m

Table 2.

Summary of Mean Values for Dry Weight and Nutrients in
Nannoplankton. Means have been Computed from
Results at Six Stations

Date

Dry

Weight
g/sq m

Protein
g/sq ID
albumin

Carbohydrates
g/sq m
glycogen

1965

20 April

21.8

1.4

0.5

4 May

7.5

0.8

0.1

25 June

10.9

1.3

0.5

31 July

25.4

1.-8

0.6

14 August

8.0

0.9

0.3

26 August

7.9

0.5

0.3

7 October

37.0

1.6

1.1

4 November

46.7

1.3

1.4

20 December

67.6

1.1

0.5

1966

23 January

42.8

1.0

2.3

28 May

95.2

2.7

0.5

17 June

123.5

6.3

6.0

1 July

22.8

2.3

1.4

15 July

28.5

1.7

1.0

29 July

61,2

3.5

1.3

13 August

17.2

3.2

0.7

26 August

12.4

2.5

0.4

6 September

14.8

2.9

0.6

Grand Mean

36.2

2.0

1.9

Mississippi Plankton Crop

307

Table 3.

Seasonal Variation in NannopIankton-to-Net Plankton Ratios for Dry
Weight, Proteins, and Carbohydrates

Date

Dry Weight
nanno/net

Proteins

nanno/net

Carbohydrate

nanno/net

1965

20 April

34

9

17

4 May

33

13

17

25 June

38

13

62

31 July

90

29

100

14 August

20

9

48

26 August

24

7

57

7 October

130

22

278

4 November

208

27

310

20 December

348

64

55

1966

23 January

996

65

2086

28 May

106

16

34

17 June

278

82

903

1 July

31

11

39

15 July

49

9

87

29 July

83

13

65

13 August

30

17

22

26 August

8

5

14

6 September

22

12

37

in summer 1965 (Table 3). An average nannoplankton rise was no-
ticed starting in fall and reaching a maximum, 67.6 g/sq m, in Decem-
ber 1965. The nannoplankton-to-net plankton dry weight ratio was
996:1 in January 1966 and 8:1 in late August 1966, the highest and
lowest for the entire study period (Table 3). Such an extreme differ-
ence in ratio probably resulted from low production of net plankton
and little or no spawning and breeding activity of herbivores at a
time when production at the nannoplankton level was high. Roth frac-
tions maintained higher average standing dry weights in summer
than in winter, with a differential, larger increase in net plankton.

Protein : Protein comprised the largest component of net plank-
ton nutrients. The dry weigh t-to-protein ratio was 3.7:1, equivalent
to 27% protein. Parsons, Stephens and Strickland (1961), Raymont

308

Mulkana and Abbott

(1963), and Blazka (1966) have shown considerably higher relative
protein levels in net plankton, apparently because their samples were
not contaminated by the large amounts of detritus, clay, and silt usu-
ally present in collections from Mississippi Sound. The high ash con-
tent in our net plankton samples should account for the low relative
protein level.

Mean net plankton protein for all samples was 136 mg/sq m. The
level varied from 62 mg/sq m to 158 mg/sq m between April and
August 1965 (Fig. 4). With the seasonal temperature decline, protein

1965 —1966

Figure 4. Seasonal variation in mean values of protein of net plankton
and nannoplankton.

values decreased to 15 mg/sq m in January 1966. As temperatures
rose again with the onset of spring, net plankton protein values again
increased, reaching 470 mg/sq m in August 1966. The summer maxi-
mum for 1966 was considerably higher than the summer values for
1966 and followed the same trend as net plankton dry weight.

The dry weight-to~protein ratio in nannoplankton was 18:1,
equivalent to 5.5% protein. Similar low levels of protein in smaller
plankton organisms have been reported by Blazka (1966), Raymont
(1963), and Parsons, Stephens and Strickland (1961). This supports
the concept that nannoplankton are mainly composed of photosyn-

Mississippi Plankton Crop

309

thesising elements and, hence, are mainly carbohydrate, not protein
(Fig. 5) . Nannoplankton protein ranged from 0.5 g/sq m to 1.8 g/sq m
during the April-December 1965 period, with maximum and mini-
mum occurring in the summer (Table 2). Data for 1966 indicated
greater variability in available protein. Values fluctuated between 1.0
g/sq m and 6.3 g/sq m, with marked variations during the summer.
Although the protein level in nannoplankton was extremely low, the
nannoplankton-to-net plankton mean protein ratio on a water column
basis was 24:1 (Table 3).

^ LI PIDS

ililii C AP BO H Y DRAT ES

NANNOPLANKTON

20 10 0 10 20

Figure 5. Average nutrient composition and dry weight of Mississippi
Sound plankton.

310

Mulkana and Abbott

Carbohydrate : Carbohydrate composed 3% of the dry net plank-
ton sample, Riving a carbohydrate-to-protein ratio of 1 :9. Various
biochemical studies on plankton agree that carbohydrate level in net
plankton is relatively low and protein makes up most of the nutrient
component (Raymont and Conover 1961, Parsons, Stephens and
Strickland 1961, Blazka 1966). Carbohydrate values for April 1965
and April 1966 were 29 mg/sq m and 24 mg/sq m, respectively (Table
1). From late spring to winter 1965, values remained at or below
10 mg/sq m with a minimum of 1 mg/sq m in January 1966 (Fig. 6).

I

nannoplankton

6i -

5 -

4 ■

3 ■

2 ■

1 ■

r

A

1965 ^ 1966

Figure 6. Seasonal variation in mean values of carbohydrate of net
plankton and nannoplankton.

Nannoplankton mean carbohydrate content was 1.9 g/sq m,
of the dry weight, with a mean ratio of 1 :1 :18 for carbohydrate : pro-
tein :dry weight (Table 2). Carbohydrate and protein formed only
about 10% of the dry weight, the low nutrient content in nannoplank-
ton apparently resulting from large amounts of detritus and clay ad-
mixed with nannoplankton samples. Ash weight determinations on
15% of the samples gave values as high as 85% ash. Although nanno-
plankton showed a lower percentage nutrient content, on a unit area
basis average carbohydrate level was a total of 235 times that of net
plankton. Protein and carbohydrate maintained almost the same levels
in the nannoplankton and showed similar seasonal fluctuations.

Mississippi Plankton Crop

311

Livid: The average net plankton lipid was of the dry

weight. The maximum, 16 mg/sq m, and minimum, 1 mg/sq m, values
were observed in August and December, respectively, in 1965 (Fig.
7). In 1966, minimum, 1 mg/sq m, and maximum, 48 mg/sq m, oc-
curred respectively, in January and August. From August 1965 on-
ward, lipid values declined sharply to a minimum in winter and grad-
ually increased with the onset of vernal conditions in 1966. The lipid
content relative to the dry weight, during August lipid peaks, was
4.7% in 1965 and only 3% in 1966 when a considerably higher stand-
ing crop was noted (Table 1). Wimpenny (1938) reported similar
results from the North Sea, and indicated that lipid content was lower
when higher standing biomass was present. Fisher (1962) pointed out
that size, spawning, maturity, and season affect lipid content of plank-
ton, but he attributed the difference in lipid content to a differential
in Species composition from one year to the next. Lipid content in
spring samples, which consisted predominantly of phytoplankton, was
lower than in summer samples which included considerable zooplank-
ton (Table 1). Blazka (1966) indicated that lipid content per unit bio-
mass increases from algae to zooplankton in fresh water habitats.
Possibly this is true in marine environments as well.

Nannoplankton samples were too small for routine determination
of lipid content, but based upon a single cruise, when 18 gallons of net
filtrate from each of the six stations were processed, lipid content was
found to be 1.2% of nannoplankton dry weight.

1965 — 1966

Figure 7. Seasonal variation in mean value of lipids of net plankton.

312

Mulkana and Abbott

Figure 4 summarizes dry weight and nutrient composition data
for net plankton and nannoplankton.

Caloric values and ash content : With one exception, net plankton
caloric values^ averaged for six stations on each date, fell within the
range 3. 6-5. 4 kcal/ash-free g (Fig. 8). This is a remarkable consis-

(M

1965 1966

Figure 8. Seasonal variation in mean caloric values and ash weights of
net plankton.

tency^ considering that net plankton dry weights in the same sample
series varied from 225 mg/sq m to 1560 mg/sq m. The mean value for
the entire series w^as 4.85 kcal/ash-free g. Richman (1958) and Golley
(1961) reported similar values for Daphnia, but caloric values ob-
tained for Copepoda and Cladocera by Sitaramiah (1967) were as
high as 7.02 kcal/ash-free g. Paine (1964) indicated that higher
values are usually found for eggs and resting stages, although well-
fed zooplankton also show a high caloric content periodically. The ex-

Mississippi Plankton Crop

313

treme value found in the mixed net plankton samples in the present
analyses was 9.9 kcal/ash-free g. This extremely high result derives
from an anomalous datum from one station on one date, and may re-
flect a technical error.

The plankton samples yielded high ash contents throughout the
period of study. Paine (1964) suggested that in organisms high in
ash content, such as sponges and opisthobranchs, at least some low
caloric estimates may result from difficulties in determining accurate-
ly the amount of noncombustible material present. Corrections for en-
dothermic combustion reactions due to high ash content may be made
as suggested by Paine (1966), but plankton samples may at times con-
tain large amounts of low energy-yielding organic detritus, thus caus-
ing low estimates. The w^et-ashing procedure employed in this investi-
gation destroys carbonates and leads to low ash estimates, thus giving
rise to slightly low results w'hen caloric values per unit weight are
computed on an ash-free basis.

Caloric content on a unit area basis followed a general trend simi-
lar to that shown by other parameters. Thus, the highest caloric value
for 1965, 1.7 kcal/sq m on April 20, corresponded to the highest value
for dry w^eight in 1965. In 1966, highest values for all parameters ex-
cept carbohydrate corresponded to highest standing dry weight. Al-
though caloric data for winter and early .spring are not available, a
trend towards decline in values w^as apparent after the peak in sum-
mer 1965. This may be associated with seasonal decrease in tempera-
ture. Vernal rise in caloric content w'as again noted in 1966, along
with the rise in other parameters. The caloric value obtained in Au-
gust 1966, 3.3 kcal/sq m, was the highest for the year. Other para-
meters in 1966 showed similar trends.

DISCUSSION

Nannoplankton, by definition, is that portion of the total plank-
ton not obtained in tow-net collections. Since it is difficult to collect in
quantity and even more difficult to study by classical, systematics-
oriented methods, nannoplankton, historically, has been ignored to a
large extent in favor of net plankton.

Nevertheless, various workers, approaching the assay of the total
plankton from several standpoints, have all agreed that, at least in
temperate and tropical w^aters, nannoplankton dominates the primary
producer trophic level. Thus^ Pomeroy and Johannes (1966) found
that 94-99% of total plankton respiration resulted from organisms
too small to be retained by a fine net; Banse (1964) estimated that
nannoplankton made up more than half of the total plankton; and
Yentsch and Ryther (1959) reported that, in Vineyard Sound, net
plankton represented, in various samples, 2-47% of total plankton

314

Mulkana and Abbott

biomass. The latter authors also tabulated other earlier results, in-
cluding- a report by Riley from Tortugas that net plankton amounted
to only one percent of total plankton.

The study described here relates to observations on the total
plankton crop of Mississippi Sound. Turbulent conditions are the rule
in this body as in other Gulf Coast estuaries and lagoons, because of
shallow waters, extensive wind stirring, river discharge from the
mainland, and tidal mixing from the Gulf of Mexico through various
passes between barrier islands. As a result, clay particles and detritus
stay in su.spension and become admixed with plankton samples, there-
by giving rise to high ash values. Further, limited data Indicate that,
probably because of fine clays, ash is differentially increased in the
nannoplankton fraction.

Despite this, nannoplankton, on a unit-area, total-water-column
basis shows a total organic content greater by orders of magnitude
than net plankton organic content. Thus, for an 18-month period in
1965 and 1966, mean nutritional composition of net plankton was 136
mg/sq m protein, as albumin, 15 mg/sq m carbohydrate, as glycogen,
and 14 mg/sq m lipid, as triolein. During the same period, nanno-
plankton samples from the same stations and times showed means of
2.0 g/sq m protein and 1.9 g/sq m carbohydrate. These values were
associated with mean dry weights of 604 mg/sq m for net plankton
and 36.2 g/sq m for nannoplankton, including ash contents of 49%
for net plankton and, based on very limited samples, 65% for nanno-
plankton.

The net plankton nutritional values amount, on a dry weight
basis, to 27% protein, 3% carbohydrate, and 3% lipid. These levels
are considerably lower than published re.sults based on clay-free sam-
ples or individual species (Parsons, Stephens and Strickland 1961,
Linford 1965, Raymont and Linford 1966, Blazka 1966). For nanno-
plankton, comparable data are not at hand. However, it has Vjeen
pointed out by Gunter (1938, 1941, 1967) that most abundant species
of fishes on the northern Gulf Coast feed at the ba.se of the food chain.
These are the menhaden, Brevoortia patroniis and B, gzmteri, the an-
chovy, Anchoa mitckilli^ and the mullet, Mugil cephalus. Quite prob-
ably a considerable portion of their food is nannoplankton but there
are no good data, although Peck (1894) said that menhaden filtered
out dinoflagellates and minute plankton. There seems to be little doubt
that oysters and other pelecypods feed to a considerable extent on
nannoplankton (Nelson 1925, 1947) and filter feeding smaller crusta-
ceans may also be in part based on nannoplankton.

No geographically uniform seasonal trend was found for nanno-
plankton standing crop, although net plankton did exhibit such a
trend, a result in accord with the findings of Yentsch and Ryther
(1959). Unfortunately, no method was available for estimating the

Mississippi Plankton Crop

315

relative contribution of salt marsh detritus to the two plankton frac-
tions. This detritus might either emphasize or mask a trend, since
Odum and de la Cruz (1967) suggested that outpourings from Georgia
salt marshes provided the major organic load in Georgia estuaries,
and that the majority of this w'as “nanno detritus.'" Study on the year-
ly cycle of nannoplankton carbohydrate in Louisiana estuaries by Mul-
kana (1969), however, shows a normal seasonal trend associated with
rise and decline of temperature.

Net plankton caloric values, on an ash-free weight basis, were
consistent with various reported values in the literature. On the aver-
age, the caloric value of 1 g, ash-free, of net plankton was equivalent
to 1.3 g glucose. On an area basi.s, caloric values were relatively
steady. There was one extreme value of 3.3 kcal/sq m in August 1966,
but the remainder fell in the range 0. 5-2.2 kcal/sq m.

Nannoplankton samples were insufficient to permit caloric deter-
minations, but should show larger values than net plankton per unit
area consistent with the much larger standing crop of nannoplankton.

SUMMARY

1. On a unit area basis, mean nannoplankton dry weight was 72
times that of net plankton from April 1965 to September 1966 in
Missis.sippi Sound,

2. The mean net plankton protein, carbohydrate and lipid were found
to be 136 mg/sq m, 15 mg/sq m and 14 mg/sq m, respectively. The
mean nannoplankton protein and carbohydrate were, respectively,
2.0 g/sq m and 1.9 g/sq m.

3. Mean levels of net plankton protein, carbohydrate and lipid were
27, 3, and 2.8 7 / > respectively, of net plankton dry weight. In nan-
noplankton, protein and carbohydrate were 5.5 and 5%, respec-
tively, relative to dry weight.

4. Although nutrient levels were higher in net plankton on unit
weight basis, total nutrients available from nannoplankton were
substantially higher. The mean nannoplankton-to-net plankton
protein and carbohydrate ratios per unit water column were 24:1
and 235:1, respectively.

5. Some seasonal trends were apparent as changes in the standing
biomass of net plankton, associated with temperature rhythm.
Nannoplankton exhibited no definite seasonal trends. Present re-
sults suggest that factors other than temperature, such as grazing
activity of herbivores and detritus influx, strongly influence nan-
noplankton dry weight.

6. Caloric values of net plankton showed seasonal trends similar to

316

Mulkana and Abbott

those shown by dry weight. Average net plankton caloric value
during the study period was 1.3 kcal/sq m.

LITERATURE CITED

B'Mise, K. 1964. On the vertical distribution of zooplankton in the sea. hi: Prog-
ress in Oceanography. Ed. by M, Sears. Perganion Press, New York, pp. 53-
12.5,

Blazka, P. 1966. The ratio of crude protein, glycogen and fat in the individual
steps of the production chain, hi: Hydrobiological Studies. Ed. by J. Hrbacek.
Academia Pub. House, Czechoslovac Acad. Sci., pp. 395-408.

Bligh, E. G., and W. J, Dyer. 1959. A rapid method of total lipid extraction and
purification. Can. J. Biochem. Physiol. 37, pp, 911-17,

Christmas, J. Y,, G. Gunter and P. Musgrave. 1966. Studies on annual abundance
of postlarval penaeid shrimp in the estuarine waters of Mississippi, as re-
lated to subsequent commercial catches. Gulf Res. Rep. 2:177-212.

Curl, H. Jr, 1962. Standing crops of carbon, nitrogen, and phosphorus and trans-
fer between trophic levels in continental shelf waters south of New York.
Rapp. P.— v. Reun Cons. perm. int. Explor, Mer 153, pp. 183-89.

Department of the Army. 1951. Methods for Medical Laboratory Technicians.
Manual No, TM8-227. Superintendent of Documents, U. S. Printing Office,
Washington 25, D. C.

Deevey, G. B. 1960. The zooplankton of the surface waters of the Delaware Bay
region. Bull. Bingham occanogr. Coll. 17, pp. 6-63.

Fisher, L. R, 1962, The total lipid material in some species of marine zooplankton.
Rapp. P.-v. Reun Cons, perm, int. Explor. Mer 153, pp. 129-35.

Folch, J., M. Lees and G. H. Sloane Stanley. 1957. A simple method for the isola-
tion and purification of total lipids from animal tissues. J, Biol. Chem. 226,
pp. 497-509.

Golley, F. B. 1961, Energy values of ecological materials. Ecology 42, pp. 581-83.

Gunter, G. 1938. The relative numbers of species of marine fish on the Louisiana
coast. The American Naturalist, 72:77-83.

. 1941. The relative number of shallow fishes of the northern Gulf of

Mexico, with some records of rare fishes from the Texas coast. The American
Midland Naturalist, 26(1) : 194-200.

1962. Shrimp landings and production of the State of Texas for the

period 1956-1959, with a comparison with other Gulf .states. Publ. Inst. Ma-
rine Sci., Univ. Texas. 8:216-26.

1963. The fertile fisheries crescent. J, Miss. Acad. Sci. 9:286-290.

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