GULF

RESEARCH

REPORTS

Volume 11
March 1999

ISSN; 0072-9027

Published by

The University of Southern Mississippi • Institute of Marine Sciences

GULF COAST RESEARCH LABORATORY

Ocean Springs, Mississippi

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999
Editorial
Mark S. Peterson

Gulf Coast Research Laboratory, mark.peterson^usm.edu

DOI; 10.18785/grr.ll01.01

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

Recommended Citation

Peterson; M. S. 1999. Editorial. Gulf Research Reports 11 (l): vii-vii.
Retrieved from http;//aquila.usm.edu/gcr/voll 1/issl/ 1

This Editorial 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 editor of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(Susm.edu.

EDITORIAL

In the early 1 960s, Dr, Gordon Gunter, then the Director
of Gulf Coast Research Laboratory, almost singlehandedly
developed the concept of G-ulf Research Reports (GRR) as
a mechanism . . devoted primarily to publication of the
data of the Marine Sciences, chiefly of the Gulf of Vlexico
and adjacent waters". The first issue appeared in April
1961 and since that time Gulf Research Reports has
produced 34 issues covering over 280 reports on the
resources and processes of the Gulf of Mexico and adjacent
waters. Many of the papers in those early issues focused
on local and regional issues, processes and problems.
Through the years, however, papers appeared front
authors outside the local and regional areas which focused
on organisms and/or processes relevant to the Gulf of
Mexico and adjacent waters. Papers have been published
from scientists in Denmark, Germany, Sweden, Canada,
Japan, Mexico, and the Caribbean Sea nations, giving a
more international flavor to the journal. The Director of
the Gulf Coast Research Laboratory (GCRL) served as
Editor of GRR until the 1997 issue.

The editorship of GRR was passed on to the late Dr.
Harold D. Howse from Dr. Gordon Gunter beginning with
the 1975 issue. At that time the journal was reformatted to
a larger page size and a nominal page charge was, for the
first time, asses.sed to help defray the cost of publication.
The first “Guide to Authors” appeared in that issue and
manuscripts had to be found acceptable by at least two
referees (Howse, editorial in GRR 5(1)). Dr. Howse was
Editor of GRR through 1 992 with volume 8(4). Dr. Thomas
D, McLlwain became Editor of GRR and guided the 1994
and 1995 issues to print. Interim GCRL Director, Dr. Robert
T. van Allcr, served as Editor of GRR for the 1996 issue.
From 1989 until 1996 Ms. Susan Griggs acted as Assistant
or Managing Editor of GRR and provided guidance with
her expert editorial and managerial skills.

I formally became Editor-in-Chief of GRR with the
1997 issue and currently serve in that capacity. Changes
in GRR procedures instituted in 1997 continue to be
modified and refined today. GRR now has an Editorial
Board that includes five GCRL scientists who, in
association with Maiuaging Editor Linda C. Skupien,
provide vital information and guidance for the production
of GRR. In 1998, the position of Editorial Associate was
added and has been filled by S. Dawnc Hard. The Editorial
Board is chaired by tjie Rditor-in-Chief, The role of the
Editorial Board is to make policy for GRR. All changes and
modifications to GRR are discussed, reviewed and voted
on by the Editorial Board. A group of Associate Editors
was appointed, including the scientists on the Editorial
Board, the Editor-in-Cliief and national and international
experts to bring disciplinary depth and international
perspective to GRR. All .Associate Editors have a tw'o-
year appointment. This major change in GRR policy has

been an important and fruitful one. At this time we removed
the page charges for published manuscripts and initiated
a nominal subscription fee. The 1997 issue included a
complete revision of the "Guide to Authors” and “Scope”
of GRR, a change in the volume numbering sequence of
GRR issues, and a minor redesign of the cover. The
Editorial Board modified the cover again in the 1998 issue
by including the new Inslilule of Marine Sciences logo in
lieu of the GCRL logo. Finally, in the 1998 issue (Volume
10), the abstracts from the annual meeting of the Gulf
Estuarine Research Society (GERS) were published in
GRR. GERS abstracts will continue to appear in GRR.
These changes were made to help our readership recognize
the changes within the Gulf Coast Research Laboratory,
the supporting structure of GRR, (see Preface of Dr. D.J.
Grimes in GRR Volume 10).

During )998, the Editorial Board in consultation with
Dr. Grimes began discussions about major changes in
GRR. The changes we envisioned will result in the ultimate
goal of making GRR a . widely recognized source of
scientific information that underpins the understanding,
planning, and management of Gulf of Mexico and
Caribbean natural resources and processes” (sec Preface
of Dr. D.J. Grimes in GRR Volume 10). Our goal was thus
to reformulate and repackage the original vision of GRR.
At the 1998 Editorial Board meeting in December, we
voted to again update the “Guide to Authors” and the
“Scope” to better reflect our mission and audience. We
also voted to remove a published submission deadline
such that more manuscripts might be submitted to the
journal with the ultimate vision of publishing two issues
annually. We voted to change the name of the journal from
Gulf Research Reports to Guff and Caribbean Research.
This change will become effective in Volume 12 published
in the year 2000. We feel this name change will more
accurately reflect the scope of the papers published in the
journal; and we hope our readership will enjoy our new
look and name, which vve feel will support and extend the
original vision of its founder Dr. Gordon Gunter. As this
issue was going to press, Dr. Gordon Gunter passed away
at the age of 89 on 19 December 1998. The Editorial Board
dedicates this issue to his memory and long standing in
the Marine Science Community, He will be long
remembered as (he founder of Gulf Research Reports.

Mark S. Peterson
Editor-in-Chief and Associate Professor
Gulf Coast Research Laboratory
Institute of Marine Sciences
The University of Southern Mississippi
703 East Beach Drive
Ocean Springs, MS 39564

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

Recent Trends in Water Clarity of Lake Pontchartrain

J.C. Francis

University of New Orleans

M.A. Poirrier

University of New Orleans

DOI: 10.18785/grr.ll01.02

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

Part of the Marine Biology Commons

Recommended Citation

FranciS;J. andM. Poirrier. 1999. Recent Trends in Water Clarity of Lake Pontchartrain. Gulf Research Reports 11 (l): 1-5.
Retrieved from http;//aquila.usm.edu/gcr/voll l/issl/2

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 editor of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(Dusm.edu.

Gulf Coast Research Reports Vol. II. 1-5, 1990

Manuscript received March 28. 1997'. accepted January 8. 1998

RECENT TRENDS IN WATER CLARITY OF LAKE PONTCHARTRAIN

J. C, Francis and M. A. Poirrier

Department of Biological Sciences. University of New Orleans, .Vcu' Orleans, Louisiana
70 NH. USA

ABSTRACT An analysis of Secchi disk transparency ob.servations from 3 sites on the Lake I’onlcharirain
Causeway indicates that water clarity has increased at the north shore and mid-lake sites, but has not changed
at the south shore site. Louisiana Department orEnviroiiiTienial Onaliiy data from I9K6 through 1995 were used
in the analysis. Lnrther analySi.s indicates that the increased transparency was not caused by changes m salinity
(jr wind speed. The best explanation lor the observed increase is the cessation of shell dredging m 1990.

Introduction

Lake Pontchartrain is an estuarine embaymeni located
in southeastern Louisiana, north of metropolitan New
Orleans. The lake has a mean salinity of about 4%. mean
depth of 3.7 in and surface area of 1,630 km-(Sikoraand
Kjerfve 1985). Several factors have contributed to the
environmental degradation of Lake Pontchartrain including
urban and agricultural runoff, shell dredging, saltwater
intrusion, operation of the Bonnet Carre Spillway and
industrial discharges (Houck el al. 1987), A major
environmental concern has been an assumed long-term
increase in turbidity based on Secchi disk transparency
observations (Stone cl al. 1 980).

Stone (1980) analyzed 4 sets of Secchi disk
transparency data and concluded that water clarity had
decreased almost 50% between 1953 and 1978. Francis et
al. (1994) also found that regression of the available
transparency data on time ( 1 953 through 1 990) suggested
a statistically significant decrease in transparency of
about 40%. The 1 953 to 1 990 data, however, were biased
in that they did not adequately represent the seasonal
effects of salinity and wind speed, fhere are strong
correlations between water clarity and salinity and wind
speed in Lake Pontchartrain, and both variables vary with
season. When the transparency data were adjusted for
the seasonal effects of salinity and wind speed or when
unbiased data sets were constructed, the data did not
support the hypothesis of a change in transparency from
1 953 to 1990.

Shell dredging was discontinued during the summer
of 1 990. It was known to have produced short-term. local
increases in turbidity, but may have had more w idespread
and lasting effects due to the production ofunconsolidated
bottom sediments that could be more easily resuspended
by wind (USACOE 1 987). I fshell dredging had long-term,
widespread effects on watej* clarity, then a comparison of
transparency data from the 1 986-90 and 199 1-95 periods
might reveal an increase in Iransparency that would be
indicative of recovery. Such evidence of recovery would

also suggest that a significant impact from sliell dredging
had occurred.

The present study wa.s conducted to determine
whether changes in water clarity as measured by Secchi
disk transparency had occurred since 1990, and thereby
provide a sequel to our earlier work (Francis etal. 1994),
and also to determine whether any observed changes
could be attributed to the cessation of shell dredging in
the lake.

Materials and Methods

Description of the Data Set

Secchi disk transparency, salinity, and turbidity data
for the 1986 to 1995 period were obtained from the Louisiana
Department of Environmental Quality (LADEQ). The data
were collected as part of an ongoing monitoring program
which includes monthly measurements at 3 stations on
the Lake Pontchartrain Causeway located approximately
4 miles (6.4 km) from the north shore, at mid-lake, and
approximately 4 miles from the south shore (Figure 1 ). A
few data points are missing in the 1986 through 1995 data
set because measurements were not taken in some months.
The missing data points were estimated by distance
weighted least squares.

Wind speed data for the 1986 to 1995 period were
recorded daily at the New Orleans International Airport.
The data set constructed for this study contains the
average wind speed for a 5-day period including the day
of transparency measurement and the 4 preceeding days.

Regional Effects of Wind

Wind probably has the same effect on transparency
in all regions of the lake. It is not possible, however, to
conduct a rigorous statistical test of that premise with the
available data. Multiple regression analysis was used
only to provide some support for the idea. Data were
selected from the LADEQ data sets for transparency and
salinity and from the wind speed data set recorded at the
New Orleans International Airport. The combined data set

1

Francis and Poirrihr

Figure 1. Map of I.ake Pontchartrain, Louisiana. The stippled area indicates areas where shell dredging was prohibited
(IJSACOE 1987). The three LADEQ monitoring sites on the Lake Pontchartrain Causeway are indicated by large dots.

has measurements of transparency, salinity and wind
speed for 119 months From 1986 through 1995. In 53
months salinity was sufficiently similar at the 3 sampling
sites to realize a coefficient of variation of 25 or less. These
data were chosen for analysis. The selection procedure
was intended to remove salinity as a significant variable
in the regression, The selection limit of25 was an arbitrary
choice. There was no autocorrelation in these data.

In regressions of transparency on salinity and wind
speed one won Id expect the partial regression coefficients
for salinity not to be significant because of the data
selection procedure, and those for wind speed to be
significant. If wind speed has the same effect on
transparency at the 3 sampling sites, then one would
expect 3 parallel regressions with different constants and
.slopes determined largely by wind. One would expect
further that the ratios of constant to slope would be the
same if the regressions are parallel.

When transparency was regressed on salinity and
wind speed, the partial regression coefficients for salinity
were not significant as expected, and those for wind speed
were significant at all 3 sites. Ratios of constant to slope
were 13.16, 13.51, and 1 1 .40 for the south shore, mid-lake
and north .shore sampling stations, respectively,

suggesting that a given wind speed produced
approximately the same percentage decrease in
transparency at the 3 sites, or that wind speed had
approximately the same effect in the different regions of
the lake.

T ransparency and Turbidity

Secchi disk transparency measurements were obtained
with a 20 cm disk with black and white quadrants,
■fransparency data were used in the present analysis to
facilitate comparison with historic data. Because Secchi
disk observations are somewhat subjective, the association
between transparency and turbidity data sets w as analyzed
to corroborate results. Pearson correlation coefficients
for transparency and turbidity were greater than 0.8
(p < 0,00 1 ) for the 3 sampling sites.

Statistical Methods

l’he4 time-scriesdatasets used in statistical analyses
(transparency, turbidity, salinity and wind speed) possess
low but statistically significant first order autocorrelation.
Autocorrelation was reduced to non-significance in each
data set by differencing with one period lag. Each dataset
thus fits a first order autoregressive model.

2

Water Clarity

Figure 2. Twelve-month moving averages of monthly Secchi disk transparency at the 3 sampling sites from 1986 through 1995.

Significance tests in analysis of variance and
regression analyses were performed with lagged data,
Residuals were analyzed to test for normality , homogeneity
of variance and independence.

Standardized partial regression coefficients may be
obtained with data transformed to standard normal form.
Standardized coefficients are useful for comparative
purposes because they are independent of scale.

Results

Twelve-month moving averages of monthly Secchi
disk transparency measurements from the south shore,
mid-lake and north shore sampling sites are presented in
Figure 2. Approximately the same transparency was
realized at all 3 sites through 1 990. After 1 990, transparency
increased at the north shore and mid-lake sampling sites,
but not at the south shore site. One-way analysis of
variance indicated that mean transparencies for the 3 sites
in the 1986-90 period were not significantly different,
p>0.5. In the 1991-95 period, however, mean
transparencies for the 3 sites were significantly different
from each other, p <0.05.

Lake-wide mean salinities were 3.98% and 3.17% in
the 1 986-90 and 1991-95 periods, respectively. The 95%
confidence intervals for these means overlap, indicating

that the higher transparencies measured in the 1991-95
period were not associated with a significant lake-wide
change in salinity. Twelve-month moving averages of
monthly salinity measurements from the south shore, mid-
lake and north shore sampling sites are presented in
Figure 3. Consistently lower salinities occurred at the
north shore throughout the 1986-95 period. The 95%
confidence interval for north shore mean salinity in the
1991-95 period docs not overlap the 95% confidence
intervals for mid-lake and south shore mean salinities. The
higher transparencies observed at the north shore in the
1991-95 period (Figure 2) were thus associated with
salinities lower (Figure 3) than were measured at other
regions of the lake.

Lake-wide mean wind speeds were 7.76 mph and
8.13 mph in the 1 986-90 and 1 99 1 -95 periods, respectively.
The 95% confidence intervals for these means overlap,
indicating that the higher transparencies measured at the
north shore in the 1991-95 period (Figure 2) were not
associated with a significant lake-wide change in wind
speed.

Multiple regression analysis was used to assess the
relative effects of salinity and wind speed on transparency
between the 1986-90 and 1991-95 periods for the south
shore and north shore sampling sites (Table 1), At the
south shore, the partial regression coefficient for salinity

3

Francis and Poirrier

Figure 3. Twelve-month moving averages of monthly salinity at the 3 sampling sites from 1986 through 1995.

was not statistically significant in both periods,
suggesting that the negative effect of wind speed was the
more prominent factor in determining transparencies.
Standardized regression coefficients for salinity at the
south shore had overlapping 30% confidence intervals as
did standardized coefficients for wind speed. At the north
shore, both partial regression coefficients were significant
in both periods (Table 1). Standardized regression
coefficients for salinity at the north shore had overlapping
30%confidence intervalsasdid standardized coefficients
for wind speed. These results indicate that the relative
effects of salinity and wind speed on transparency were
different at the 2 sampling sites. More importantly for the
purpose of this paper, the results also indicate that the
effects of salinity and wind speed were approximately the
same in both periods at a given sampling site.

Discussion

The similarity of standardized regression coefficients
in the 1 986-90 and 1 99 1 -95 periods at the south shore and
north shore sampling sites (Table 1) indicate that the
higher transparencies measured at the north shore in the
1 99 1 -95 period (Figure 2) cannot be explained by changes
in salinity or wind speed.

Salinity has a statistically significant positive effect
on transparency, and wind speed has a statistically
significant negative effect on transparency (Francis et al.

1994). Higher transparencies, therefore, are usually
associated with higher salinities and lower wind speeds.
An unusual feature of the reported results is that the
higher transparencies observed at the north shore in the
1 991-95 period were not associated with higher salinities
or lower wind speeds, but rather with lower salinities than
those measured at the mid-lake and south shore sampling
sites and with wind speeds that were the same ailhe 3 sites.

The higher transparencies (Figure 2) and higher
regression constant (Table 1) at the north shore during
the 1 99 1 -95 period may be explained by the posit ive effect
on transparency realized through cessation of shell
dredging. Sediment disruption produced by shell dredging
probably had a greater negative effect on transparency in
the lower-salinity waters of the north shore (Figure 3)
because of the tendency for lower-salinity waters to
retain particles in suspension longer (Francis ct al. 1994).
By reducing transparencies at the north shore in the 1 986-
90 period, shell dredging probably was responsible for
the lower regression constant for that period (Table 1).
Shell dredging was not present in the 1991-95 period
resulting in higher transparencies and a higher regression
constant.

Higher transparency peaks were apparent at the nortli
shore and mid-lake sampling sites by the fall of 1991
(Figure 2). This observation is consistent with expectation
because an immediate increase in transparency was not
anticipated. Unconsolidated sediments that are more

4

Wathr Clarity

TABLE 1

Regression analyses of transparency vs. salinity and wind speed for south shore and north shore sites in 1986 through
1990 and 1991 through 1995,

Site and Period

Coefficient

Standardized

Coefficient

P

South Shore 1 986-90

Constant

153.57

Salinity

5.55

0.25

0.213

Wind speed

-11.13

-0.54

0.021

South Shore 1 99! -95

Constant

163.84

7.42

0.18

0.389

Salinity

Wind speed

-13.93

0.61

<0.001

North Shore 1986-90

Constant

92.11

Salinity

14.12

0.55

0.006

Wind .speed

-6.01

-0.23

0.091

North Shore 1991-95

Constant

155,63

Salinity

34.02

0.57

0.003

Wind speed

13.94

-0.37

0.006

susceptible to resuspension by wind (USACOE 1987)
would persist for a period of time following dredging and
have a longer-term effect on turbidity. In addition, an
earlier expression of higher transparency may have been
mitigated by lower lake-wide salinities in 1990 and early
1991 (Figure 3) that would have lowered transparency.

Transparency remained essentially unchanged at the
south shore after shell dredging was stopped. Several
factors may have contributed to this outcome. Dredging
was proh ibited with in 3 m i les of the south shore extendi ng
from the Lake Pontchartrain Causeway east to Paris Road
in Orleans Parish, and near oil and gas facilities in Jefferson
Parish west of the causeway (Figure 1). Consequently,
dredging and its effects on transparency may have been
less intense near the south .shore sit(h The south shore is
subject to urban runoff from metropolitan New Orleans,
and it has a highly modified .shore line with no exchange
with natural streams and wetlands. Runoff introduces
nutrients that can promote algal growth with the result
that turbidity from phytoplankton growth may have
replaced turbidity from resuspended sediments.

Shell dredging began in 1933 and probably affected
transparency prior to the first transparency measurements
in 1953. The cessationofshelldredgingin 1990 reestablished
conditions favoring higher transparencies in some regions
of the lake. The change to higher transparencies cannot be
attributed to changes in salinity or wind speed.

Acknowledgment

We would like to acknowledge the generous financial
support of this work by Freeport-McMoRan, inc.

Lu er.mt're Cited

Francis. J.C.. M.A. Poirrier. D.E, Uarbe, V. WijesumJera and
M.M. M III ino. 1994. Historic trends in the Secchi disk
transparency of Lake Pontchartrain. GulfRcsearch Reports
9:1-16.

Houck. O. A., F. Wagner and J.D. Flslrott. 1987, To restore Lake
Pontchartrain. The Greater New Orleans Expressway
Commissiun. New' Orleans, LA. 269 p,

Sikora. W.B and B Kjerfve. 1985. Factors inlliiencing the
salinity regime ofLakc Pontchartrain. l.oui.siana. a shallow
coastal lagoon: Analysis of a long-term data set. Estuaries
8:170-180.

Slone, l.ll. (cd.) 1980. I'nvironmcnia! analysis of Lake
Poincharirnin. Louisiana, its surrounding wetlands, and
selected land uses, Vol 1 and 2. Louisiana Slate University
Center for Wetland Resources. Baton Rouge, LA, Prepared
for the U, S. Army Engineering District. New' Orleans.
Contract No. DACW-29-77-C-0253,

U, S. Army Corp.s of Engineers. 1987. Clam shell dredging in
Lakes Pontchartrain and Maurepas. Louisiana— draft
environmental impact siaicntcnl and appendixes. U. S.
Army Corps of Engineers. New' Orleans District, New
Orleans. LA,

5

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

Effects ofDiflubenzeron on the Ontogeny of Phototaxis hj Palaemonetes pugio

J.E.H. Wilson

Morgan State University

R.B. Forward Jr.

J.D. Cosdow

DOI: 10.18785/grr.ll01.03

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Part of the Marine Biology Commons

Recommended Citation

Wilsori; J., R. Forward Jr. and J. Costlow. 1999. Effects ofDiflubenzeron on the Ontogeny of Phototaxis hy Palaemonetes pugio. Gulf
Research Reports 11 (l): 7-14.

Retrieved from http://aquila.usm.edu/gcr/voll l/issl/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 editor ofThe Aquila Digital Community. For more information, please contact Joshua.Cromwell^usm.edu.

Gulf Research Reports Voi, II, 7-14, 1999

Manuscript received July 30, 1997: accepted April 10, 1998

EFFECTS OF DIFLUBENZURON ON THE ONTOGENY OF
PHOTOTAXIS BY PALAEMONETES PUGIO

J.E.H. Wilson*, R.B. Forward, Jr.^ and J.D. Costlow^

‘Department of Biology, Morgan State University, Baltimore, Maryland 2 1 25 1 .USA
M 35 Duke Marine Lah Rd., Beaufort, North' Carolina 28516, USA
^201 Ann Street. Beaufort, North Carolina 28516, USA

ABSTRACTYhe pholoiaxis by larvae of the grass shrimp Palaemonetes pugio that hatched from embryos which
were exposed to a single pulse concentration of diflubenzuron (DFB: Dimilin^Sj) was quantified. Stage IV
embryos (6-day-old) were exposed to 0.5 pg/L of DFB for 4 days followed by transfer into clean seawater for
the rest of the incubation period. The phoioresponscs of lighl-adapled larvae from untreated embryos and
embryos treated with 0.5 pgfL DFB were monitored from 1 day through 8 day post hatch for phototactic
rcsponsc.s to 500 nm light. Larvae from untreated embryos exhibited strong positive photolaxis at high light
iiilensilics (3 x lO*- and 3 x 10 ' Wm’-) but became negatively photolactic at lower light intensities (3 x 10 ’ to
3 X 1 0'^ Wm'^). Thisphototaclic pattern continued during the monitoring period. On the other hand, larvae from
DFB-lreatcd embryos exhibited altered phototaxis for the firsts days. Alterations were especially evident on
Day 1. as larvae were only negatively photolactic. By Day 4. these larvae reverted to the normal pattern of
phoioresponscs shown by untreated larvae. These results indicated that the alterations in phoioresponscs of
larvae caused by embryonic exposure to DFB are only transitory and can be corrected within 4 days of hatching
if the larvae arc exposed to water lacking DFB.

Introduction and Costlow 1974). Also, ontogenetic changes in

photoresponses are observed in some crustaceans.
Diflubenzuron (DFB; Dimilin®) is an insect growth Generally, the younger stages are more positively
regulator that interferes with chitin formation and molting phototactic while negative phototaxis increases in the

in arthropods. It is approved for and is being used in the older larval stages, postlarvae, and adults (see review by

United States for control of a wide variety of insect pests, Pardi and Papi 1961, Dingle 1969). Because of the role of

including foliage feeders on soybeans, cotton-leaf phototaxisin vertical migration of crustacean larvae, any

perforator, and forest insects. In California DFB is used to alteration in this photoresponse as a result of exposure to

control mosquito larvae (Fischer and Len wood 1 992). The toxicants may affect the ecology and conceivably the

effects of DFB on non-target anthropods, especially larvae’s recruitment into the adult population,

aquatic organisms, is well documented (see review by Photo behavior has been shown to be very sensitive

Fischer and Lenwood 1 992). There is always the potential to changes in environmental factors such as temperature,

forDFBimpactingaquaticorganismsbecauseofoverspray salinity, and chemicals. Changes in photobehavior have

or spi I Is, especially where it is being applied close to water also been used in aquatic toxicology as a sensitive ind icator

or directly onto wetlands for mosquito control of anthropogenic stress (Rosenthal and Alderdice 1976,

Phototaxis and its ecological significance in Simonetetal. 1978, Langetal. 1981, Rand l985).Speciftcally

crustaceans is well documented in the literature (White for larval crustaceans, the following studies have employed

1 924, Thorson 1964, Forward 1974, Vernbergetal. 1974, changes in photobehavior as indicators of sublethal

Forward etal. l984,Sulkin 1984), Forexample, Ina review toxicity: Forward and Costlow (1976) for insect juvenile
by Thorson ( 1 964) of marine benthic invertebrates, of the hormone mimic on Rhithropanopeiis horrisii: Moyer and

14 1 species studied, 82%oftheearjy larval stages respond Barthalmus (1979) for the herbicide Weeder-64 on
positively to light. Phototaxis has also been reported to Palaemonetes pugio; Lang et al. (1980) for copper on
play an important role in diel vertical migration ofcrustacean Balanus improvisus. In all these studies, the larvae were

larvae (Forward 1 976, Forward and Cronin 1 980, Forward directly exposed to the toxicant followed by measurement
et al, 1 984, Forward 1 985). Vertical migration contributes of phototaxis. Only Wilson (1985) and Wilson etal. ( 1 985)

to the dispersal of crustacean larvae and helps in their have reported alterations in phototaxis by larval stages of

retention in the estuary (Sulkin 1 975, Cronin 1 979, Cronin crustaceans as a result of embryonic exposure to a toxicant,

and Forward 1 986). Both the level and sign of phototaxis were altered in Itght-

For larval stages of estuarine crustaceans, the adapted first stage larvae of P. /jr/g/o after 4 -day single
phototactic pattern, when tested in a narrow light field, is pulse exposure of the embryos to DFB. These alterations

generally negative phototaxis to low light intensities and in phototaxis were shown to be dependent on the DFB

positive phototaxis to moderate intensities (e.g. Forward concentration and the embryonic stage al exposure

7

Wilson f.t al.

(Wilson etal. 1985). The present study was conducted to
determine iTand when larval grass shrimp from DFB-
treated etnbryos which exhibit altered phototaxis regain
normal pattern of phototaxis during larval development.

M A l ERIALS AND METHODS

Ovigerous female grass shrimp P. pugio that were
induced to spawn in the laboratory (Duke University
Marine Laboratory, Beaufort. NC) were sorted according
to stage of embryonic development as described by
Wilson ( 1 985). Laboratory animals were used in this study
because they were relatively homogeneous and gave less
variable results than field animals. Only ovigerous females
carrying Stage IV embryos (6-day-old; body segmentation
stage, at 25 ± 1°C) were used in this study. Earlier studies
by Wilson (1985) and Wilson etaL(1985)haveshown that
Stage IV embryos are the most sensitive embryonic stage
and represent a midpoint in the embryonic development
of P, pugio. The shrimp were placed in largeculture dishes
(insidediameter = 20 cm) containing 0.5 pg/Lofwettable
powder (WP-25%) formation of DFB dissolved in 20%o
filtered (to 45 pm) seawater. Untreated 20%o filtered
seawater served as the control. This test concentration
was used because Wilson et al. ( 1 985) have shown that for
phototaxis, 0.5 pg/L is the lowest observed effect
concentration (LOEC) when various embryonic exposure
concentrations were used. The shrimp were maintained at
a density of 5 per liter of test solution for 4 days without
renewal (single dose exposure). After the 4-day exposure,
the shrimp were transferred into clean seawater (20%o),
which was changed every day until the eggs hatched. The
larvae were then used in phototaxis experiments. The
rationale for exposing embryos rather than larvae is that
this test protocol, delayed subleihal bioassay (DSB), has
been shown to be more sensitive than shrimp or crab larval
bioassays (see Wilson 1 985 for details). Ovigerous females
and larvae were reared in an environmental chamber set at
25"Cand 12L: 1 2D photoperiod, centered at 1 200 h. Animals
were fed freshly hatched Artemia sp. nauplii daily.

Experiments were performed to determine ontogeny
of phototaxis of larvae hatched from unexposed embryos
(control) and embryos exposed to 0,5 pg/L DFB. The
general protocol for all phototaxis experiments was that
described by Wilson etal. ( 1985) with few modifications.
Phototaxis was determ ined by measuring the direction of
swimming immediately following light stimulation. Ten to
15 larvae were placed in an acrylic trough measuring
14.9 x 8.3 X 3.5 cm containing approx unately 1 10 ml filtered
seaw'ater (20%o), The trough was divided into 5 equal
compartments by acrylic partitions which could be raised

or lowered simultaneously. The stimulus light was
presented horizontally from a slide projector fitted with a
300 watt incandescent bulb. The light was interference-
filtered to 500 nm(7 nm halfbandwidlh).Thisw'avelength
was selected because it has been shown to be the spectral
sensitivity maximum forP,/7wg/o (personal communication,
John K. Douglas, University of Arizona, Tucson, AZ
85721, unpublished) and A vM/gc?m( White 1924). Intensity
was regulated by neutral-density filters (Detric Optics,
Inc.) and measured with a radiometer (from EG&G
model 550).

Phototaxis measurements were performed in a
photographic darkroom between midnight and 0300 h.
This lime was chosen to coincide with the time of maximum
larval release by laboratory -maintained ovigerous females
(personal observations), thereby ensuring that larvae
were 24 ± 2 h old when first tested. By monitoring
phototaxis at the same time of day for all experiments,
complications due to biological rhythms in behavior (see
Forward and Cronin 1980) were avoided. Shrimp larvae
were light adapted for 4 -6 hto 12.53 Wm'Might intensity
(cool-white fluorescent lamps) prior to testing. Ten to 1 5
larvae were placed in the central compartment of the
acrylic trough and allowed to adapt in darkness for 30 s.
After this, the partitions were raised gently and the
stimulus light turned on simultaneously. Larvae were then
stimulated for 60 s then the partitions were lowered and
the stimulus light turned off. The number of larvae in each
compartment was recorded. Larvae were returned to rearing
conditions and tested on subsequent days. A new group
of larvae were then introduced into the trough and tested
as previously outlined. This procedure w'as repeated at
least 3 times before the neutral density filters were changed
to test a different intensity of the stimulus light. Six to 7
different light intensities were tested plus a“dark control’'
in which the movements of larvae in the test trough were
monitored without any stimulus light. Different larvae
were used for each stimulus light level. The larvae were fed
throughout the phototaxis experiments to reduce the
possibility of altered phototax is due to starvation (Cron in
and Forward 1980, Lang etal. 1980). The intensity versus
response curves for these larvae were again determined
on the second day (i.e., for2-day-oId larvae). Using the
same batch of larvae, this procedure was repealed every
day up to Day 4 and again on Day 8 . Examination of both
untreated and treated larvae on Day 4 indicated that they
had stalked eyes and thus had molted to the 2nd zoeal
stage.

Positivephototaxis wasdefmed as movement towards
the light source and negative phototaxis as movement
away from the light source. The animals in the 2

8

Ontogeny of phototaxis by grass shrimp larvae

compartments closest to the light source were regarded as
showing positive phototaxis; those in the 2 compartments
farthest from the light source as negatively phototactic.
The mean percentage positive and negative response and
their standard errors (S.E.) were calculated at each light
intensity. For statistical analysis, the percentages were
first arcsine transformed. Statistical tests determined the
difference between dark control (no light stimulus)
response levels due to movement in the test trough in
darkness and responses upon stimulation with light. Chi-
square tests and analysis of variance were performed on the
results as described by Sokal and Rohlf (1981), The level of
significance was set at P = 0.05 for all tests.

Results

Larvae from Unexposed Embryos

The intensity versus response curves for light-
adapted larvae from unexposed embryos during ontogeny
are shown (Figure 1 ). The pattern of phototaxis exhibited
by Day 1 larvae (Stage I) remains virtually the same
through Day 8 of development. As compared to the dark
control level of responsiveness, larvae were positively
phototactic (P < 0.05; ANOVA) at the stimulation
intensityof3 x I O'* (days 2,4, and 8) or at 3 x 10'^ Wm’^and
higher intensities (Days 1 and 3). Larvae were negatively
phototactic(P < 0.05; ANOVA) at lower light intensities
with the threshold being 3 x 10 ' Wm’* for Days 1 to 4 and
one log unit higher for Day 8.

There is some indication of increased activity by the
larvae with age as evidenced by the increase in the dark control
responses of larvae. The positive control (no light present)
increased from 26% on Day 1 to 40% on day 8 (Figure 1 ).

Larvae from Embryos Exposed to DFB

The ontogenetic changes observed in the
photoresponses of light-adapted larvae that hatched from
embryos (Stage IV) exposed to 0.5 |ig/L DBF are presented
in Figure 2, Positive phototaxiswasabsent(relativetothe
dark controls) at the stimulation intensities that normally
evoked significant positive responses in untreated larvae
(3 X 10'^ Wm'^ and higher; Figure 1). Compared with Day
1 untreated larvae (Figure l),the larvae from DFB-treated
embryos exhibited negative phototaxis (P < 0.05; ANOVA)
(F igure 2) over a much wider range of stimulus intensities
(3 X 10-' to 10'* Wm-^).

By Day 2, the first sign of a return to the normal
pattern of phototaxis was evident as seen by an increase
in positive phototaxis from the control level on Day 1 to
72% on the second day at 3 x I O ' Wm'^ stimulation intensity
(Figure 2). The positive responses at 3x1 O ' Wnr^on Days

2 and 3 by treated larvae are not significantly different
(P > 0.05; chi-square) from each other (Figure 2). At an
intensity of 3 x 10 - Wm *, Days 2 and 3 larvae remained
strongly negatively phototactic. However, by Day 4, the
larvae exhibited positive phototaxis at both 3 x 10 - and
3x10 ' Wm'^ (see Figure 2). Thus, the return to normal
photoresponse is complete by Day 4 for larvae from
embryos exposed toO.5 pg/L DFB. The response patterns
exhibited by 4- and 8-day-old larvae were almost identical.
The lowe.st light intensity evoking positive phototaxis
and the highest intensity that evokes negative phototaxis
for unexposed and exposed larvae are compared in Table I .
Although these threshold intensities were very different
for 1 -day-old treated and untreated larvae, they became
identical by Day 4.

Discussion

The phototactic pattern of Stage 1 larvae from the
grass shrimp P. pugio has been extensively documented
by Wilson ( 1 985) and Wilson et al. ( 1 985). The pattern of
phototaxis of light adapted Stage 1 larvae from untreated
embryos was positive phototaxis at high light intensities
(3x lO'^and 3x 1 O'* Wm -) and negative phototaxis at lower
light intensities (3 x 10'-' to 3 x 10'^ Wm'^; Figure 1; Wilson
et al. 1 985). This pattern of phototaxis persists for larvae
from untreated embryos irrespective of the age of the
embryos when incubation started in the laboratory (Wilson
1985, Wilson et al. 1985), For larvae that hatched from
DFB-treated embryos, both the magnitude and the sign of
the photoresponse were altered. Such larvae consistently
exhibited negative phototaxis at higher light intensities
that normally evoke positive phototaxis (3x10'^ and
3x 1 0 * Wnv^). These alterations in phototaxis varied upon
exposing embryos to concentration of DFB ranging from
0.3 to 1.0 pg/L (Wilson etal. 1985). However, at exposure
concentrations of ^ 2.5 pg/L, larvae exhibited severe
structural abnormalities, and the magnitude of both
positive and negative phototaxis was drastically reduced
(Wilson 1985).

Results of the present study indicate that for light-
adapted Stage I larvae from unexposed embryos, phototaxis
remains virtually unchanged during larval development.
Both the pattern of the stimulus light intensity versus
phototactic response curves and the magnitude of the
phototactic responses were similar for all the larval stages
tested (up to 8 days old). It should be pointed out that this
pattern of phototaxis by light-adapted larvae was also
observed up to Day 15 (Wilson unpublished data).
However, at the postlarval stage (unpublished data) both
positive and negative phototaxis are lost since the animals

9

PERCENT RESPONSE

Wilson et al.

Figure I. Palaemonetes pugio. Intensity versus response curves for different ages of light-adapted larvae hatched from
untreated embryos (i.e., incubated in seawater throughout embryonic development). Open circles, dashed lines
represent negative phototaxis. Closed circles, solid lines represent positive phototaxis. DC = dark control values for
larvae moving to the positive and negative chambers of the test trough in the absence of light. Data points are means + S.E.
The sample size (n) for each stimulus intensity was 3. Asterisks indicate means that arc significantly (P < 0.05) greater
or less than the appropriate dark control. Embryos were 6 days old when incubation started.

10

PERCENT RESPONSE

Ontogeny of phototaxis by grass shrimp larvae

Figure 2. Palaemoneles pugio. Intensity versus response curves for different ages of light-adapted larvae hatched from
embryos that were exposed to 0.5 ^g/L diflubenzuron starting when they were 6 days old. Open circles, dashed lines
represent negative phototaxis. Closed circles, solid lines represent positive phototaxis. DC = dark control values for
larvae moving to the positive and negative chambers of the test trough in the absence of light. Data points are means + S.E.
The sample size (n) for each light intensity was 3. Asterisks indicate means that are significantly (P < 0.05) greater or
less than the appropriate dark control.

11

Wilson et al.

TABLE 1

Comparison of lowest light intensity that evokes positive phototaxis and highest light intensity evoking negative
phototaxis in grass shrimp larvae from untreated control and diflubenzuron (DBF)-exposed embryos. NR is no
phototactic response.

Larval Age
(Days)

Positive Response
(Lowest Intensity) Wm'^

Negative Response
(Highest Intensity) Wm’^

untreated

DFB-exposed

untreated

DFB-exposed

1

NR

3x10'^

3x10'

2

3x10-'

3x1 0-‘

3

3x10'

3x10-

4

9

3x10-'

3x10'

8

9

3x10-'

3x10-'

were unresponsive to even the highest stimulation
intensity used (3 x 10"^ Wm’^atSOO nm light). Forward and
Costlow( 1 974) have reported a sim ilar pattern in phototaxis
during ontogeny for the mud crab, R. harrisii. Both the
action spectra and the intensity versus- response curves
for light- and dark -adapted animals were similar for all
zoeal stages On metamorphosis into the megalopa stage,
there was a dramatic change in behavior similar to that
reported here for the postlai vae of the grass shrimp.
These findings are different from those reported by Welsh
( 1 932) for the mussel crab and by Hunte and Myers (1984)
for estuarine amphipods, where changes from positive to
negative phototaxis were observed during larval
development. In some instances, (e.g. in Balanus) there
is a change from positive phototaxis in newly hatched
nauplii to negative in Stage II and back to positive in the
cyprid stage (Thorson 1964).

The lack of ontogenetic changes in pholotaxis of P.
pugio larvae from untreated embryos made it relatively
easy to determine when larvae from DFB-treated embryos
regained normal photobehavior. By comparing the pattern
of the intensity versus response curves for each age of the
larvae from untreated and DFB-treated embryos, it was
observed that a return to normal photobehavior started
w'ith Day 2 larvae and by the time they were 4 days old, the
response patterns were similar to that of the untreated
group. Thus, it is possible for larvae with altered
photobehavior resulting from embry'otoxicily of DBF to
regain their normal photoresponsiveness within 2 to 4 days
if reared in clean seawater during larval development.

Microscopic examination indicated that 4-day-old
treated and untreated larvae had molted to the 2nd zoeal
stage in the present experiment. Therefore, the change back
to normal pattern of phototaxis by light-adapted larvae from
DFB-exposed embr>'os was completed after the larvae molted
to the 2nd stage. Although there are reports of altered

phototaxis by crustacean larvae and adults resulting from
exposure to toxicant (Bigford 1977, Forward and Costlow
1976, Lang etal. 1980, Moyer and Barthalmus 1979, Wilson
et al. 1985), the present study is the first report of re-
establishment of normal phototaxis upon removal of the
toxicant during larval development.

In untreated Stage I larvae the eyes are sessile with
cuticular lens and apposition optics, i.e., the lenses form
small inverted images on the rhabdoms (Land 1984,
Fincham 1 984). For details on the structure and function of
grass shrimp eyes, see Parker ( 1 897), Douglass ( 1 986), and
Douglass and Forward (1989). Ontogenetic study of the
compound eyes of P. pugio from larval to postlarval stage
shows that the basic morphological and anatomical
organization of the eyes remain unchanged throughout
larval development (Douglass and Forward 1989). It is
therefore not surprising that the photoresponse of untreated
larvae remain the same during larval development in this
.study. 1'he altered photoresponse seen in larvae from DFB-
exposed embryos is conceivably the result of structural
modification of the visual system of the larvae. Grass shrimp
larvae hatched from embryos exposed to 0.5 pg/L DFB have
been shown to exhibit slight morphological abnormalities
(terata), which also affect swimming speed and vertical
distribution in a seawater column (Wilson etal. 1985, Wilson
etal. 1987).

Ultrastructural study of the exoskeleton of the mud
crab R. harrisii by Christiansen and Costlow (1982)
revealed that larvae exposed to DFB had disorganized and
swollen exocuticle. Since the thickness of the cuticle is the
same in Rhithropanopeus and Palaemonetes (Freeman
1993) and the effects of DFB on larval crustaceans is
similar, it can be presumed that larvae from DFB-treated
embryos may have swollen and malformed cuticular facets
in the eyes. Such swollen cuticular facets may alter the
entire optics of the larval eyes and could account for the

12

OntociENy ov phototaxis by grass shrimp larvae

observed reversal in phototaxis. In apposition eyes, the
cuticular facet acts as a lens which focuses light on the
rhabdom (Cronin 1 986). Conceivably, when the lens is not
properly formed, e.g., has granular disorganized
endocuticle (see Mulder and Gijswijt 1 973), or is swollen,
the amount of light passingthrough will be reduced. This
may explain why expo.sed larvae responded negatively at
light intensities to which they normally reactedpositively .
Normal phototaxis is restored upon moiling probably as
a result of formation of new cuticular facets with normal
thickness and endocuticle. It is also possible that the
distribution of the visual pigments in DKB-treated larvae
is altered as a result of biochemical changes. Irrespective
of what mechanism caused alteration in phototaxis, it is
clear from the present study that normal phototaxis was
restored after the larvae molt to the 2nd zocal stage.

Since larvae were tested in an unnatural light Held
(e.g. Forward 1 985), relating phototaxis to actual behavior
in nature isdifficult. Nevertheless, theresultsdo indicate
photobehavior W'as altered by exposure to DFB, and thus,
aspects of larval ecology that depend on photobehavior
would be altered. Photobehavior is involved in diel vertical
migration of the larvae, and hence their temporal vertical
distribution in an estuary (Allen and Barker 1 98.5) could be
altered. Since their vertical distribution affects horizontal
transport, recruitment to the adult population would be
affected. The ability to avoid predators could also be
reduced by alterations in photobehavior, since the negative
phototaxis participates in a predator avoidance shadow
response (Forward 1977). Also, Douglass et al. (1992)
demonstrated that P. pugio larvae have endogenous
phototaxis rhythm, which if altered would change the
photoresponse pattern throughout the tidal cycle in an
the estuary. Thus, the survival potential of the shrimp
population could be reduced by alteration in larval
photobehavior.

In summary, the pattern of phototaxis by grass shrimp
larvae from untreated embryos remains unchanged during
larval development. This pattern consists of a positive
phototaxis at high light intensity (> 3 x 10 ^ Wm’^) and
negativephototaxis at lower intensities (^ 3 x 10‘^ Wm *).
Although larvae from DFB-treated embryos had altered
phototaxis, photobehavior was gradually restored as the
larvae developed in clean water, and restoration was
complete upon molting to the 2nd zoeal stage. Hence,
altered phototaxis as a result of embryotoxiciiy to DFB is
only temporary in grass shrimp larvae.

Acknowledg.ments

This material is based on research supported in part
by AFGRAD Fellowship from the African American
Institute and National Science Foundation Grant No.
OCE-9596175 to J.E.H. Wilson, Duke University Marine
Laboratory graduate student research funds. The technical
assistance of M. Forward. M. Hartwill and A. Wilson is
gratefully acknowledged.

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14

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

Parasitization of Callinectes rathbunae and Callinectes sapidus by the Rhizocephalan Barnacle
Loxothylacus texanus in Alvarado Lagoon^ Veracruz^ Mexico

Fernando Alvarez

Universidad Nacional Autonoma de Mexico

Adolfo Gracia

Universidad Nacional Autonoma de Mexico

Rafael Robles

Universidad Nacional Autonoma de Mexico

Jorge Calderon

Universidad Nacional Autonoma de Mexico

DOI: 10.18785/grr.ll01.04

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

Part of the Marine Biology Commons

Recommended Citation

Alvarez, E, A. Gracia, R. Robles and J. Calderon. 1999. Parasitization of Callinectes rathbunae and Callinectes sapidus by the
Rhizocephalan Barnacle Lo:>£:of/iy /flcus te:>canus in Alvarado Lagoon, Veracruz, Mexico. Gulf Research Reports 11 (l); 15-21.
Retrieved from http:// aquila.usm.edu/gcr/voll l/issl/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 editor ofThe Aquila Digital Community. For more information, please contact Joshua.Cromwell^usm.edu.

Gulf Research Reports Vol, 11. 15-21. 1999

Manuscript received February 23, 1998; accepted June 21. 1998

PARASITIZATION OF CALLINECTES RATH BUN AE AND CALLINECTES
SAPIDVS BY THE RHIZOCEPHALAN BARNACLE LOXOTHYLACUS
TEXANUS IN ALVARADO LAGOON, VERACRUZ, MEXICO

Fernando Alvarez', Adolfo GraciaS Rafael Robles' and Jorge Calderon'

‘Colcccion Nacional de Crustdeeos, Instiiuto de Biologia, Vniversidad Nadonal A utonoma
de Mexico, Apart ado Postal 70-153, Mexico 045 10 D.F., Mexico

‘Inslituto de Ciencias del Mar y Limnologia, Vniversidad Nacional Autonoma de Mexico,

Apartado Postal 70-305, Mexico 045 10 D.F,, Mexico

A liSTRACT Calhnectesrathbunae anti Callinectessapidus in Alvarado Lagoon. Mexico, weresampled monihlv
tor one year to determine the extent of para.sitization by the rhizoeephalan cirripede l.oxothylacus lexanus.
Prevalence levels, host sex ratio, carapace width-weight variation, and disiribulionof the number ofparasiies
among hosts were analyzed, Loxothylacus texanus was present almost c.xdusi vely in C. rathbunae with a mean
prevalence of?. 58%, while less than 1% of all C. sapidus were parasitized. Callinectes rathbunae constitutes
a new host record for this parasite. A study of infection revealed significant variation in prevalence and host
size throughout the .study period. The sex ratio of parasitized crabs differed from that of the total sample with
males being para.silizcd more often, and the comparison of carapace width- weight relationships revealed lower
weights oi parasitized crabs.

Introduction

A number of studies on the rhizoeephalan barnacle
Loxothylacus texanus Bo.schma parasitizing the blue crab,
Callinectes supidiis. In the Gulf of Mexico have been
published in the last several decades describing: temporal
and geographic variation in prevalence (Adkins 1972*
Hochbergetal. 1992, Lazaro-Chavezetal. 1996), host size
distribution (Christmas 1969, Adkins 1972, Ragan and
Matherne 1974), morphological modifications of hosts
(Reinhard 1950, Alvarez and Calderdn 1996), and the
relationship between host size and parasite size (Wardle
andTirpak 1 991). The interest In the effect of this parasite
on the commercially important blue crab is renewed
whenever a nevv outbreak is detected (Wardle and Tirpak
1991) and few long-term prevalence records have been
kept(0’Brien and Overstreet 1991).

Until recently, no published information existed on
the extent of the blue crab-rhizocephalan interaction in
Mexican waters ofthe Gulf of Mexico, although parasitized
crabs have long been recognized by local fishermen.
Loxothylacus texanus is well established in the Gulf of
Mexico occurring in C. sapidus from southern Florida to
Campeche (Hochbergetal. 1992, Alvarez and Calderon
1996) and in C rathbunae from central Veracruz to
Terminos Lagoon, Campeche (AlvarezandCalderdn 1996).
Loxothylacus texanus has been reported outside the Gulf
of Mexico in Callinectes larvatus in the Canal Zone,
Panama (Boschma 1 950), and in C at 4 sites along

the Caribbean coast of Colombia (Young and Campos
1988, AIvarezandBIain 1993).

A one yearsurvey forL. texanus by monthly samplings
of C. rathbunae and C. sapidttswas conducted in Alvarado

Lagoon, southern Veracruz (Figure 1 ), to determ ine parasite
prevalence levels, host species selectivity, host carapace
width-weight variation, and distribution of number of
parasites per host.

M.\terials and Method.s

Monthly samples (12) of Callinectes spp. from
Alvarado Lagoon were examined from November 1 995 to
November 1996 (except October). Data were obtained
from the catch of local fishermen. Their catch was collected
and processed in the “Cooperativa Primero de Abril”, in
Alvarado, Veracruz. Crabs were identified, measured
(carapace width), weighed, and sexed. Male crabs were
classified as parasitized by L. texanits if they presented an
abnormally shaped abdomen and atrophied first pleopods.
Female crabs were considered parasitized ifthey presented
atrophied pleopods with mature abdominal shape. Crabs
of both genders were considered parasitized if they
exhibited the parasite externae, or bore scars in the
abdomen where externae had been attached. All crabs in
which morphological modifications were detected, but
which did not bear an externa were labeled as "‘feminized”.
When present, externae were counted and classified as
immature (small, mantle opening not developed) or mature
(full-sized, mantle opening fully developed). An average
of 1 77 crabs was examined monthly.

Statistical analysis of data included: Student's t-
test, analysis of variance (ANO V A), analysis of covariance
(ANCOV A), G-test of independence, and Ch i-square test.
All crab sizes are reported in millimeters (mm) and weights
in grams (g); mean values are followed by ± one standard
error.

15

Alvarez et al.

Results

A total of 2, 132 crabs was examined, which included
668 C. sapidusdX{6 1 ,464 C /•a//i6wwcr6r. Overall prevalences
were 0,75% (5 crabs parasitized) in C sapidm^nd 7,58%
(111 crabs parasitized) in C. rathbunae. The 5 parasitized
C sapidus were collected in January (1), March (3), and
November (1). Prevalence inC. rathbunae between
2% and 1 2% in ten of 1 2 collections; maximum prevalence
was recorded in December (23.68%) whereas no parasitized
crabs were collected in March (Figure 2).

One male and 4 female C sapidus were found to be
parasitized. Statistical analysis was not performed on this
species due to small sample size. Parasitized C. rathbunae
included 62 males and 49 females ( 1 .26 males per female),
while the unparasitized population was represented by
549 males and 804 females (1.46 females per male).
Comparison of these values shows that the parasitized
condition was not independent of sex (G-test, P < 0.005),
and that males were parasitized more often than females.

Mean size of parasitized crabs varied significantly
between host species (t-test). In C sapidus the overall

Figure 2. Prevalence of Loxoihylacus texanus in Callinecies rathbunae from Alvarado Lagoon (1995-1996). Sample size
indicated as number of parasitized crabs/total examined.

16

h, TEXANUS \>i CaLLINECTES SPP.

Figure 3. Mean size (CVV) of parasitized (solid circles) and unparasitized (open circles) Callinecfes rathbuitae in Alvarado
Lagoon (1995-1996); error bars represent ± one standard error.

mean was 1 1 1,60 ± 6.01 mm(n = 5,range92-l30 mm), while
in C. rathhunae it was 95.48 ± 0,80 mm (n = 111,
range 69-122 mm). Dueto the small number of parasitized
C. sapidus, no further analyses were performed. Mean
size for parasitized C. ralhbiinae w^s less than that of the
unparasitized population (99 i 3.61 mm in May to
78 'J: 9.04 mm in September); however, no significant
differences were encountered (AMOVA with months as
treatments) (Figure 3). Mean size of parasitized male
(94.25 ± 0.89 mm, n = 60, range 78^1 10 mm) and female
crabs ( 97.02 ± 1 .4 mm, n = 49, range 69- 1 22 mm) did not
differ statistically (t-test).

Carapace width- weight relationships forC. rathhunae
were significant for both parasitized (%, y = 1.62 X - 100.46,
n = 54,r = 0.68, P < 0.001;&,y = 0.84X - 25.95,n =41,
r = 0.46, P < 0.0 1 ; Figure 4) and unparasitized crabs (%,
y = 2.12X- 143.55, n= 116,r = 0.93,P<0.001;&,y= 1.87
X- 125.23,n= 142, r = 0.93, P < 0.0001 ; Figure 5). The
slopes of 4 regressions (ANCOV A with carapace width as
covariate, F^, = 26.09, P< 0.0001) were not

homogeneous even when the weight values of the 4
categories of crabs overlapped extensively in the 80- 1 1 0
mm of carapace width interval. Unparasitized males had
the highest slope, followed respectively by unparasitized
females, parasitized males, and parasitized females.

Of the n 1 parasitized C rathhunae, 19(17.1 2%) were
feminized ( 12 males and 7 females), and 92 (82.88%) bore
externae (50 males and 42 females). The number of parasite
externae per host varied from one to four: 64 .86% had one,
14.4 1 % had two, 2,7% had three, and 0.9% had four. The

observed pattern did not conform to a Poisson (random)
distribution (Table 1) and may reflect an aggregated
pattern since the observed frequencies ofmultiple externae
are much higher than expected and the coefficient of
dispersion is greater than one (CD = ! .45). Throughout
the year, the relative frequencies of internal (feminized
hosts), immature, and mature parasites did not seem to
follow a defined pattern (Figure 6).

Discussion

In Alvarado Lagoon, C rathhunae was the main host
for L. texanus, even though C, sapidus was locally
abundant. Callinecies rathhunae was parasitized by L.
texanus only south of Casitas, Veracruz (Alvarez and
Calderdn 1 996). To the north, throughout roughly half of
its distribution range, the C. rathhunae population was
not found to carry L, texanus. Examination of collections
of crabs from Tamiahua Lagoon, north of Casitas, has
shown that while L. texanus prevalence in C sapidus can
reach 5 1 .5%, no C. rathhunae are known to be parasitized
in the area (Lazaro-Chavez et al. 1996). In contrast, in
Alvarado Lagoon, only 5 C. sapidus were found parasitized
throughout the present study, while prevalence in C.
rathhunae reached 23.68%.

Most rhizocephalans exhibit a loose specificity,
commonly parasitizing 2 or more closely-related host
species, often of the same genus (Hoeg 1 995). Conditions
that may promote new host species acquisition when a
host species and a closely related potential host species

17

Alvarez lt al.

CARAPACE WIDTH (mm)

Figure 4. Carapace width-weight relationship of parasitized Callinectes rathbunae in Alvarado Lagoon (white
circles = females, black circle = males).

occur sympatrically have not been explored. In
Loxothylacus panopaei^ which parasitizes 4 species of
xanthid crabs along the east coast of North America, the
differential levels of parasitization in each host species
may be due to subtle di fferences in the spatial distribution
within the estuary as well as to that of infective parasite
larvae (Walker et al. 1992, Alvarez 1993). Within the Gulf
of Mexico, the apparent abandonment by L. texamis of C.

sapidus and its subsequent acquisition of C rathbunae
cannot be explained with the available data. However, the
observed pattern could also be the result of L. /exanus
parasitizing the less desirable C. sapidus where C,
rathbunae is not available.

Loxothylacus iexanus occurs outside the Gulf of
Mexico southward to Colombia (Young and Campos 1 988.
Alvarez and Blain 1 993). In Panama, C larvatus has been

CARAPACE WIDTH (mm)

Figure 5. Carapace width-weight relationship of unparasitized Callinectes rathbunae In Alvarado Lagoon (white
circles = females, black circles = males).

L. TEXANUS IN CaLLINECTES SPP.

Table 1

Distribution of externae of Loxothylacus texanus in 1,445 Callinectes rathbunae from Alvarado Lagoon. Feminized crabs
(n = 19) with no externae arc not included. Observed frequencies are compared (Chi-square test) to the expected
frequencies of a Poisson (random) distribution.

M umber of externae
per host

Observed frequencies

Expected frequencies

(O - Ef/E

0

1,353

1,332.61

0.312

1

72

107.94

11.966

2

16

4.37

30.951

3

3

0.118

70.38

4

1

0.0024

414.669

Total

1,445

1,445.04

= 528.288, p< 0.0001

reported as a host species for/.. ^e.Ya/7WA (Boschina 1950);
unfortunately no other data from the region are available,
and the parasitization of other species of Callinectes by
L. texamis cannot be ruled out.

As has been reported in other studies on blue crabs
parasitized by L /exanus in the GulfofMexico, in Alvarado
Lagoon there is significant variation in prevalence
throughout the annual cycle. This is probably due to the
varying intensity of host recruitment synchronized with
high temperatures and the parasite’s reproductive activity

(Hochbergetal. 1992, Lazaro-Chavezetal. 1996). Maximum
prevalences of L. /exanus in Alvarado Lagoon (3.09% in
C. sapidus and 23,68% in C raihbunae) are low and
intermediate, respectively, compared to those from other
reports from the GulfofMexico (Table 2). Mean prevalence
of L texanus in C. sapidus in the present study is extremely
low (0.5%), while in C. rathbunae it can be considered
high (6.28%). The size ranges of parasitized crabs of both
host species in Alvarado Lagoon are intermediate between
the smaller parasitized crabs from Louisiana and Texas and

>

o

ZSL

a:

q:

>-

2

-J

O

Cl

»-

>

O

UJ

<

LU

<

OL

<

3

3

3

LU

O

o

2L

Q

U-

<

—3

<

o

2

Figure 6. Frequency distribution of Loxothylacus texanus in Callinectes rathbunae by developmental stage: white bars
represent internal parasites (feminized hosts), gray bars represent immature parasites, and black bars represent mature
externae. In March 1996, no parasitized crabs were found in the sample. In October 1996, no sample was taken.

19

Alvarez et al.

Table 2

Mean and maximum Loxoihyiacus lexanus prevalence and host size range variation of Callinectes sapidus and Callinectes
rathbunae in the Gulf of Mexico; only externae bearing crabs are considered.

Authority

Locality

Host Species

Mean
prevalence
(%) ± 1 s.d.

Maximum
prevalence (%)

Host size
range (mm)

Adkins, 1972

Louisitma, USA

C. sapidus

4,83± 4.8

17.10

30-95

Wardle and Tirpak,

1991

Galveston, Texas,

USA

C. sapidus

8.22± 13.7

53.00

43-100

Hochberg et al.,

1992

west coast of Florida,
USA

C. sapidus

1.40± 1.3

5.10

35-170

Lazaro-Chavez et al.,
1996

Tamiahua Lagoon,
Mexico

C. sapidus

17.6±19.7

51.50

45-115

Present study

Alvarado Lagoon,
Mexico

C. sapidus

0.50 ± 1.06

3.09

95-130

Present study

Alvarado Lagoon,
Mexico

C. rathbunae

6.28 ± 6.51

23.68

69-122

the large parasitized individuals found in Florida (Table 2).
No pattern of variation associated with geographic
distribution is apparent, except that thesmallest parasitized
crabs occur in the northern Gulf of Mexico.

Although an abnormal abdominal shape combined
with atrophied pleopods in C sapidus and C. rathbunae
are unmistakable signs of parasitization by L. texanus,
reported prevalence values are mostly based on externae-
carrying crabs (Reinhard 1950, Alvarez and Calderon
1996). In Alvarado Lagoon 17. 12% of all parasitized crabs
showed signs of parasitization but did not bear externae,
and were classified as feminized, while in Tamiahua
Lagoon, almost half (48%) of all parasitized crabs were
feminized (L^zaro-Chavez et al. 1996). These 2 studies
show that the margin of error of prevalence estimates that
do not consider feminized crabs can be considerable.

The sex ratio of parasitized C rathbunae \n Alvarado
Lagoon suggests that malesare preferentially parasitized.
No explanation for this biased sex ratio is apparent, since
there is no evidence that infective female cyprid larvae
show any selective behavior, at least in L. panopaei
(Alvarez et al. 1995), In contrast, in Tamiahua Lagoon,
although males were more abundant, female C. sapidus
were parasitized more often (Lazaro-Chlivezet al. 1 996).

The number of C with multiple externae of

L. texamis occurred in a higher proportion than expected
under a random distribution. No mechanism other than
chance encounters between infective cyprid larvae and
susceptible hosts is currently known to determine the
number of parasite externae that emerge from a single host
(Walker etal. 1992).

Acknowledgments

We thank the Direccion General de Asuntos del
Personal Academico (DGAPA) of the Universidad
Nacional Autonoma de Mexico for providing funds for
this project through grant “IN 2 1 0595” to A. Gracia. We
also thank Mr. Eligio Gamboa for taking care of the
sampling logistics and the fishermen Mr. Abelardo Ruiz,
Mr. Pedro Ruiz and Mr. Ignacio Ruiz for their assistance
in the field.

Liter.vtdre Cited

Adkins, G. 1972. Notes on the occurrence and distribution of
the rhizocephalan parasite {Loxothylacus iexanus Boschma)
of blue crabs [Callinectes sapidus Rathbun) in Louisiana
estuaries. Louisiana Wildlife and Fisheries Commi.ssion,
Technical Bulletin 2:1-13.

Alvarez, V. 1 993. The interaction between a parasitic barnacle,
Loxothylacus panopaei (Cirripcdia: Rhizocephala). and
three of its crab host species (Brachy ura: Xanthidae) along
the east coast of North .America, Ph.D. Dissertation.
University of Maryland. College Park. MD. 180 p.
Alvarez, R. and L.M. Blain. 1993. Regisiro dc Loxothylacus
Boschma 1 928 (Crustacea: Cirripcdia: Sacculinidae) en el
suroeste del Caribe colombiano. Actualidades Bioldgicas
19:39.

Alvarez, F. and J, Caldcrbn. 1996, Distribution of Loxo//?y/acM.v
texanus (Cirripcdia: Rhizocephala) parasitizing crabs of
the genus Callinectes in the southwestern Gulf of Mexico,
Gulf Research Reports 9:205-2 1 0.

20

L. TEXANVS IN CaLLINECTES SPP.

Alvarez, F., A.H. Hines and M.L. Rcaka-Kudla. 1995. The
effects of parasitism by the barnacle Loxothylacus panopaei
(Cirripedia: Rhizoccphaia) on growth and survival of the
host crab Rhilhropanopem harrisii (Brachyura: Xanthidae).
Journal of Experimental Marine Biology and Ecology
192:221-232.

Boschma, 11. 1950. Notes on the Sacculinidae, chiefly in the
collection of the United States National Museum.
Zoologische Vcrhandelingen 7.1-55.

Christmas, J.Y. 1969. Parasilicbarnacles in Mississippi estuaries
with special reference to To-To//i>'/ac«5 Boschma in

the blue crab {Callinectes sapidufs). Proceedings of the
22nd Annual Conference of the Southeastern Association
of Game and Fish Commissioners, p. 272-275.

Hochberg, R.J., T.M. Bert, P. Steele and S.D. Brown, 1992.
Parasitization of Loxothylacus texanus on CaJlinectes
aspects of population biology and effects on host
morphology. Bui letin of Marine Science 50; 1 17-132.

Hoeg, J.T. 1 995. The biology and lifecycle of the Rhizocephala
(Cirripedia). Journal ofthe Marine Biological Association
of the United Kingdom 75:517-550.

Lazaro-Ch^vez, E-,F. Alvarez and C. Rosas. 1996. Records of
Loxothylacus /exflwns(CiiTipedia;Rhizocephala) parasitizing
the blue crab Callinectes sapidus in Tamiahua Lagoon,
Mexico. Journal of Crustacean Biology 16:105-1 10.

O’Brien, J. and R. Overstreet. 1991. Parasite-host interactions
between the rhizocephalan barnacle, Loxothylacus texanus,
and the blue crab, Callinectes sapidus. American Zoologist
31:91.

Ragan, J.G. and B.A. Mathcrne. 1974. Studies oi Loxothylacus
texanus. In: R.L. Amborskl, M,A. Hood and R.R. Miller,
eds.. Proceedings, 1 974 Gulf CoastRegional Symposium
on Diseases of Aquatic Animals, Louisiana State University
Sea Grant Publication 74-05, p. 185-203.

Reinhard, E.G. 1 950. An analysis of the effects of a sacculinid
parasite on the external morphology of Callinectes sapidus.
Biological Bulletin 98:277-288.

Walker, G., A.S. Clare, D. RIttschof and D. Mensching. 1992.
Aspects ofthe life cycle of Loxothylacus panopaei{Q\ss\ct),
a sacculinid parasite of the mud crab, Rhithropanopeus
harrisii (Gould): a laboratory study. Journal of
Experimental Marine B iology and Ecology 1 57: 1 8 1 - 1 93 .

Wardle, W. J. and A.J. Tirpak. 1991. Occurrence and distribution
of an outbreak of infection of Loxothylacus texanus
(Rhizocephala) inblue crabs in Galveston Bay, Texas, with
special reference to size and coloration of the parasite’s
external reproductive structures. Journal of Crustacean
Biology 1 1:553-560.

Y oung, P. S. and N.H. Campos. 1 988. Cirripedia (Crustacea) de
la zona intermareal e infralitoral de la regibn de SantaMarta,
Colombia. Anales del Instituto dc Invcstigaciones Marinas
de Puntade Betin 18:153-164.

21

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

A Survey of the Reef-Related Medusa (Cnidaria) Community in the Western Caribbean
Sea

E. Suarez-Morales

El Colegio de la Frontera Sur, Mexico

L. Segura-Puertas

Universidad Nacional Autonoma de Mexico

R. Gasca

El Colegio de la Frontera Sur, Mexico

DOI: 10.18785/grr.ll01.05

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

Part of the Marine Biology Commons

Recommended Citation

Suarez-Morales; E., L. Segura-Puertas and R. Gasca. 1999. A Survey of the Reef-Related Medusa (Cnidaria) Community in the
Western Caribbean Sea. GulfResearch Reports 11 (l): 23-31.

Retrieved from http://aquila.usm.edu/gcr/voll l/issl/5

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 editor of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

Gulf Research Reports Vol. 11, 23’31, 1999

Manuscript received March 18, 1998; accepted May 12, 1998

A SURVEY OF THE REEF-RELATED MEDUSA (CNIDARIA)
COMMUNITY IN THE WESTERN CARIBBEAN SEA

E. Sufirez-Morales’, L. Segura-Puertas^ and R. Gasca*

'El Colegio de la Frontera Sur-Unidad Chetumal, P.O. Box 424, Chetumal, Quintana Roo
77000, Mexico

Wniversidad Nacional Autonoma de Mexico, Institute de Ciencias del Mar y Limnologia,
Estacion Puerto Morelos, P.O. Box 1 152, Cancun, Quintana Roo 77501, Mexico

The species composition, distribution, and abundance of medusae collected during a 4-day plankton
survey in a reef system of the Mexican Caribbean were stud icd, Highest mean medusae abundance was observed
over the fore-reef zone and in daytime samples. Lowest abundances occurred in the reef lagoon and at dusk.
Seventeen species were identified, with Liriope tetraphylla.Aglaura hemistoma.,Cubaia aphrodite, andSarsia
prolifera being the most abundant. They belong to a group of medusae dominant along the world’s second largest
barrier reef Cluster analysis revealed primary (fore-reef) and secondary (reef lagoon, channel) oceanic grou ps.
showing the strong oceanic influence along and across the reef system. Day-to-day variation in the reef medusan
community seemed relatively unimportant. The community structure of the reef medusa fauna appeared to be
quite uniform despite the expected migratory behavior of these predators, tidal exchange across the reef,
introduction of oceanic species, and time of day. The species composition was most closely related to that of
the Campeche Bank and oceanic Caribbean waters. Dominance of oceanic medusae within the reef lagoon was
attributed to the narrowness of the continental shelf and the mesoscale hydrological features of the zone.

Introduction

The medusa fauna of coastal, neritic and oceanic
waters of the Northwestern Tropical Atlantic has been
investigated by several surveys (Phillips 1972, Burke
1975, Segura-Puertas 1991, 1992, Segura-Puertas and
Ord6nez-L6pez 1994, Suarez-Moralesetal. I997,Suarez-
Moralesetal. 1998). However, relatively little emphasis
has been placed on coastal environments, where medusae
can play a relevant role as predators in the zooplankton
food webs (Raymont 1983). Studies dealing with these
cnidarians have been developed in estuarine and littoral
systems of the Mexican Caribbean (Collado ct al. 1988,
Zamponi el al. 1 990, Zamponi and Su&rez-Morales 1991,
Su^rez-Morales et al. 1998). Along this coast runs the
world’s second largest barrierreef system (Jordan 1 993).
Coral reef zooplankton has been surveyed mainly for the
most abundant groups such as copepods (Renon 1977,
1993, McKinnon 1991), but not for the less numerous
zooplankters of a higher trophic level, such as medusae.
There are no previous works dealing with the medusa
fauna dwelling in this Mexican reef system. The closest
regional antecedent for reef-related medusae is the
qualitative survey of Larson (1982) from samples collected
in the Carrie Bow Cay reef area off Belize.

This study describes changes in the numerical
abundance, composition and diversity of the reef-related
medusa fauna of the Mahahual reef system, Mexican
Caribbean Sea. The survey comprised a 4-day period, (30
December 1990-2 January 1991), and describes the small-
scale space and time variation of the medusan community.

Previous works on the plankton of this reef area refer to
zooplankton groups (Castellanos and Sudrez-Morales 1997)
andto ichthyoplankton (Vasquez-Yeomansetal. 1998).

Study Area

TheMahahual reef area liesbetween 1 8°43' and 1 8®46'N
and 87® 42' and 87°42'27” W, on the southern portion of the
Mexican coast ofthe Caribbean Sea (Figure 1 ). The entire
coast receives the influence of Caribbean waters before
flowing into the Gulf of Mexico through the Yucatan
Channel. The shelf is narrow along this coast and depth
increases rapidly offshore (Merino and Otero 1991). A
large barrier reef runs along the Mexican Caribbean, from
Isla Contoy in the north down through the Belizean coast
(Jordan 1993). Mahahual is a small fishing village located
on the southern portion of the Yucatan eastern coast. In
this area the reefbarrier forms a shallow (1 .5 m) and narrow
(30-180 m) reef lagoon. Benthic vegetation within the
lagoon is dominated by beds of Thalassta iestudinum.
Coral cover is minimal along the shallow portions ofthe
lagoon, but increases towards the fore-reef. Surface water
temperature is highest in July-August (32®C), and lowest
in December- January (2 1 ®C). Mean annual salinity along
this coast varies within the 32-36%o range. Oceanographic
conditions over this zone are influenced by the Yucatan
Current, which flows northward and by a coastal counter
current which flows southward. Interaction of both currents
produces inshoreward, semi-circular trajectories of drifting
objects (Merino 1986). This flow, coupled with tidal
currents and turbulence, seems to be the most relevant
hydrological phenomenon affecting the reef zooplankton
(Su^rez-Morales and Rivera-Arriaga 1998).

23

Suarez -Morales et al.

87 ® 42 ’ 30 ”

87 ® 42 ’ 00 "

Figure 1. Surveyed area with zooplankton sampling stations, Mahahual reef zone, Mexican coast of the Caribbean Sea.

Materials and Methods

A 4-day zooplankton sampling program was carried
out from 30 December 1 990 to 2 January 1991, during the
full moon. Stations were located to investigate the three
main reef-related zones: fore-reef (FR), Stations 1 and 2;
channel (CH), Station 3; and reef lagoon (RL), Station 4
(Figure 1). Daytime sampling was made hourly between
0700 and 1200; evening (dusk) samples were collected
between 1730 and 1930. No night collections were made
on Day 4. Zooplankton was collected by surface hauls (0-
50 m) using a square-mouthed (0.45 m per side) standard
plankton net (0.3 m m mesh). This gear allowed collection
ofsmall and medium-sized medusae. A digital flowmeter
was attached to the net mouth to estimate the volume of
water filtered. The mean amount of water filtered during

each trawl was 160 m\ At least one replicate tow was
performed at each sampling station. Zooplankton samples
were fixed and preserved in buffered 4% formaldehyde
solution (Smith and Richardson 1979). Medusae were
sorted from the entire sample and then identified and
counted to obtain the species density (org./lOO m*).
Zooplankton density dataware not significantly different
among collections (V^squez-Yeomans et al. 1997).
Shannon-Wiener’s Diversity Index (bits/individual, which
represents the degree of uncertainty about the identity of a
given species) and the Index of Importance Value (IIV, a
dominance measurement) were estimated for each collection.
TheBray-Curtis Similarity Index(Ludwig and Reynolds 1 988)
was used in the construction of a dendrogram clustering the
stations. These calculations were performed with the aid of
the ANACOM software computer program (De la Cruz 1 994).

24

Reef medusae of the Western Caribbean Sea

Results

Conditions throughout the surveyed period were quite
uniform. Mean surface temperature during the surveyed
period ranged from 26° to 2 8°C. Salinity averaged 36%o, and
ranged from 34 to 38%o.

Total medusa densities showed temporal variation
through the survey period. Highest total mean densities were
recorded during the morning of the first day, the highest two
beingatStation2(578org./100 m^), and at Station 1 (469 org./
100 m^), both representing the fore-reef zone. Values at the
other localities ranged from 7 to 280 org./ 1 00 m^ Highest mean
medusae density occurred in Day 1 over the fore-reef (Station
2,421 org./100 m’).

Overall data for the three reef zones considered herein
showed that medusae were most abundant over the fore-reef
(mean density 1 85 org./ 1 00 m^), followed by the channel(l 8
org./lOO m^)and by thereeflagoon(16.7org./100 m^).Upto
87% of the total medusae numbers occurred over the fore-
reef, and only 4% in the reef lagoon. Total density was 1 .4
times higher in the morning (91 org./ 100 m’) than at dusk
(67 org./lOO m^), with 64%ofthe individuals being collected
during daytime samplings. Over the fore-reef, density values
at daytime (190 org ./1 00 m^)andatdusk(l76org./100 m^)were
similar. At the reef lagoon, values were 28 org./ 1 00 m’ (AM)
and6org./100 m^ (PM);at the channelzone values were 18.4

and 15.2,respectively(Figiu'e2). Overall mean density varied
day to day. Values recorded were as follows: Day 1, 135
org./100 m’;Day2,54.35org./l00 m^; Day 3, 45.1 org./100 m^;
Day 4, 97.6 org./lOO Up to 40% of the total medusan
numbers were collected during Day 1 , 1 3% in Day 2, 1 9% in
Day 3, and 28% in Day 4 (only AM).

A total of 17 medusan species were identified (Table 1 ).
The most abundant, Liriope tetraphylla (Chamisso and
Eysenhardt 1821), accounted for 4 1 % of the medusae, with a
mean density of 33 .3 org./ 100 m\ Also abundant were /Ig/aura
/2em/5/o/naP<5ronandLesueur 181 0(22%; 17.8 org./lOO m’),
CubaiaaphrcKiitey\.dyQv\%9^{\ 1.6%;9.4 org,/ 100 m^^Sarsia
prolifera Forbes 1 848 (8.2%; 6.6 orgV 100 nP), and Oheliasp.
(7.11 %; 5.7 org./ 1 00 m^). These fivecomprised about 90% of
the total overall medusan catch. The relative abundance,
estimated density, and frequency of all the medusan species
recorded in the area are presented in Table I .

Liriope tetraphylla showed an overall mean density
in daytime samples of 48 org. /1 00 m^ with lower values in
dusk samples (40 org./ 1 00 m^). The same tendency in day
vs dusk samples was observed for Obelia sp., 12 org./
100 m^vs4 org./lOO m^\Clytiafolleata(}AcCv^<iy \%59),
5.7 org./lOO m^ vs 1 .5 org./lOO m^; and S, prolifera. 7.8
org./lOO m^ vs 1.8 org./lOO m\ Values for .4. hemistoma
were equal in day ( 1 7.25 org./l 00 m’) and night samples
(18.5 org./ 100 m^).

Figure 2. Mean day/night densities (org./lOOm^) of medusae in the three reef-related environments.

25

SuArez-Morales et al.

Liriope tetraphylla was most abundant at the fore- in the reef lagoon (L. ietraphylla, C. aphrodite, H. disticha,

reef. More than 90% of the total numbers of this species Zanclea costata, and S, protifera). Overall diversity

occurred in this environment. Only 8,3% occurred in the (Shannon- Wiener) was highest at the fore-reef (1 .66 bits/

channel, and the remaining 1.4% reached the lagoon. ind.). In this environment, day samples were more diverse
Aglaura hemistoma was collected only at the fore-reef. (1 .84 bits/ind.) than those collected at dusk ( 1 .38 bits/ind.).

Cubaia aphrodite was most abundant at the fore-reef The reeflagoon (0.4 bits/ind.) and the channel zones (0.6 bits/

(57%), and was more abundant at the channel zone (27%) ind.) showed low'er overall diversity values,

than at the reef lagoon (15%). occurred Clustering with the Bray-Curtis Index produced a

mostly over the fore-reef (80.6%), and was scarce at the dendrogram (Figures) which two large groups of stations
channel zone ( 1 5%) and the reef lagoon (4.3%). were defined. One group included all the fore-reef stations,

Several species occurred in either day or dusk samples, and in the other group the remaining stations (reef lagoon

and in a specific environment. Occuring only in fore-reef and channel) were clustered and mixed,

samples at dusk were Podocoryne minima (Trincil903),

Amphinemadinema(PeTor\ and Lesueur 1 809) and Halitiara Discussion

formosa Fewkes 1882. Amphinema rugosum (Mayer 1 900)

and Cunina octonaria (McCrady 1852) were recorded only Only 44% of the species recorded at Mahahual have

in fore-reef day time samples. HaIocor<fyIedisticha (Goldfuss been previously reported from the reef area off Belize (Larson

1 820) was observed only in the reef lagoon at dusk. 1982), while 50% are known from neritic and oceanic waters

The species richness was highest at the fore-reef, where of the Gulf of Mexico (Phillips 1 972, Burke 1 975), and 72%

16 out of the 17 medusa species were recorded. Only three from the Campeche Bank andthe Mexican Caribbean (Phillips

species(l. tetraphylla, S.prolifera, andC. aphrodite) v/ere 1972,ZamponietaI. 1990,ZamponiandSu^ez-Morales 1991,
recorded in the channel zone, and only five were observed Segura-Puertas 1992, Segura-Puertas and Ord6nez-L6pez

Figure 3. Dendrogram from clustering by Bray-Curtis Index showing distribution of the clusters in the three reef-related
environments during the surveyed period.

26

Medusan species density (org./100m^) by environment, sampling day, and time of day at Mahahual

Reef medusae of the Western Caribbean Sea

27

Rhopalonema velatum

Suarez-Morales et al.

1 994, Su^ez-Moralesetal. 1995, Suarez-Morales etal. 1998).
Only one species collected at Mahahual (S. prolifera) has
not been recorded previously in the region. It has been
reported from the northeastern Atlantic (Ranson 1925,
Sanderson 1930, Russell 193 8), and even from the Black Sea
(Thiel 1935). This is the first record of this species in the
northwestern Atlantic.

The number of species collected in this survey ( 1 7) is
relatively low when compared with the medusa richness
recorded in adjacent zones. Sixty-two species have been
recorded from the Campeche Bank and the Mexican
Caribbean (Phillips 1972, Segura-Puertas 1992, Segura-
Puertas and Ord6flez-L6pez 1994, Suarez-Morales et al.
1998). More than 20 species were found in a large
embayment on the central portion of the Mex ican Caribbean
coast (Suarez-Morales et al. 1 997).

The reef-related medusa fauna recorded off Belize by
Larson (1982) can be compared with that recorded over
Mahahual reef. Both belong to the same barrier reef
system. Larson recorded 71 species in reef-related areas
of Carrie Bow Cay, of which 80% were recorded in the fore-
reef and 64% in the reef lagoon. The corresponding
values for Mahahual were 88% at the fore-reef, 27% at the
channel zone, and only 20% at the reef lagoon. It is

difficult to explain the differences in species richness with
respect to Larson’s (1982) results in a reef environment.
To obtain most of the samples, he used a net with a 0.56
mm mesh opening, filtered an average of 250 m’, and made
surface tows; hiscollections were made between 1730 and
1830. Up to this point, Larson’s methods are similar to
those we used at Mahahual. The main difference was
probably related to material analyzed by Larson resulting
from qualitative collections performed while diving, using
light traps at night, and sampling with a beach seine and
with dip nets. Medusa densities are commonly low in reef
environments (Sammarco and Greenshaw 1 984, Morales
and Murillo 1996). The overall mean density recorded at
Mahahual is similar (83 org./lOO to that recorded by
Larson ( 1 982) in plankton trawls from Carrie Bow (92.5 org./
100 m’). However, there is no estimate on the species
richness from plankton net collections. Larson (1982)
recognized only 13 species as dominant; of this group, 8
are shared with the Mahahual community. In both cases,
L. tetraphylla^A. hemistoma^Solmundellabitentaculata,
and C. aphrodite were among the most abundant medusae.
However, abundance of the dominant species in both
systems showed several differences (Table 2). This
suggests that although the number of species is almost

TABLE2

The medusae collected in this survey at Mahahual and previously from the Campeche Bank, the Mexican Caribbean, and
Belize. Key for citations in this table: 1. Larson (1982), 2. Phillips (1972), 3. Segura-Puertas (1992), 4. Segura-Puertas
and Orddiiez-L6pez (1994), 5. Zamponi et al. (1990), 6. Zamponi and SuArez-Morales (1991), 7. Suirez-Morales et al.
(1995), and Suarez-Morales et al. (1997). *Not previously recorded in the Caribbean Sea or Gulf of Mexico.

Mahahual
(this survey)

Campeche Bank
(3,4)

Mexican
Caribbean
(2, 5, 6, 7)

BeUze

(1)

Amphinema dimma

X

X

Amphinema rugosum

X

X

X

Zanclea costata

X

X

Obelia sp.

X

X

Clytia mccradyi

X

X

Clytia folleata

X

X

Solmundelb bitentaculata

X

X

X

Liriope tetrophylla

X

X

X

X

Aglaura hemistoma

X

X

X

X

Rhopalonema vela turn

X

X

X

Carybdea marsupialis

X

X

X

Podocoryne minima

X

X

Sarsia prolifera

X

Haliiiara formosa

X

X

Cubaia aphrodite

X

X

Hahcmlyle disticha

X

X ,

Curana octonaria

X

X

X

28

Reef medusae of the Western Caribbean Sea

TABLES

Density of the five dominant medusa species at two Caribbean reef environments.

Mahahual (this survey)

Carrie Bow Cay (Larson 1 982)

Relative density
(%)

Density

(org./100nf)

Relative density
(%)

Density

(org/lOOm?)

Clytia mccradyi + C. folleata
(as Phialucium in Laison 1 982)

4.1

35.5

32.0

28.0

Solmundella bitentaculata

1.3

1.04

1.5

1.2

Liriope tetraphylla

41.0

33.3

57.0

48.6

Aglaura hemistoma

22.0

18.0

3.3

2.9

Cubaia aphrodite

11.6

9.4

0.4

0.3

four times higher at Belize, the distribution of the species
richness, the overall density, and the abundance of the
dominant species are similar in the two surveys. This
probably relates to the local abundance of C. aphrodite
and of A. hemistoma at Mahahual; both were relatively
scarce at Carrie Bow. A^laura hemistoma is probably
even more abundant in Mahahual during summer as
recorded off the Caribbean (Su^rez-Morales et al. 1 998).
Differences between the medusa fauna of Carrie Bow and
Mahahual could be related to the physiographic features
of each particular reef section.

In both areas, most of the remaining medusan species
occurred in low numbers, which is a common feature of the
medusa communities (Gili and Pages 1 987). The arrival of
these oceanic species effects a local enrichment of species,
but does not produce a major increase in the overall
number of individuals. This pattern agrees with parallel
results of Gili et al. (1988) from studies of cnidarian
zooplankton in the western Mediterranean.

The nearshore hydrographic structure along the
Caribbean coast of Mexico is related to the flow of a
coastal countercurrent moving southward (Merino 1986)
from the northernmost edge of the Caribbean coast. Its
influence would explain, at least partially, the high affinity
of the local medusa fauna with that of the Campeche Bank
and the southern Gulf of Mexico (Phillips 1972, Segura-
Puertas and Ord6flez-L6pez 1994), and the relatively low
affinity with the adjacent Belizean reef, which lies to the
south (Larson 1982).

Segura-Puertas and Ord6flez-L6pez ( 1 994) reported
6 species (A^ hemistoma, L, tetraphylla, Nausithoe
punctata^ Rhopalonema y datum, Eutima gracilis and
Z. costata) as being the most common in the Campeche
Bank and the oceanic Mexican Caribbean Sea. Our results

and those of Larson (1982) show that L. tetraphylla and
A . hemistoma are also successful over reef environments
(Table 3). Aglaura hemistoma has been reported as highly
abundant in other tropical and subtropical environments
(Gili and Pages 1987, Gili et al. 1 988). The seasonal (March-
May) occurrence of the aggregating scyphozoan Linuche
unguiculata seems to be a distinctive and dominant
feature of the western Caribbean neritic and near ocean ic
environments (Larson 1982, Su^rez-Moralesetal.|998).
Reflecting its well-known seasonal behavior, this species
was absent from our samples.

Communities of planktonic cnidarians are frequently
dominated by a few of the most common species (Pugli
and Boxshall 1 984, Gili and Pag6s 1987). Our results are
similar. The most dominant medusae were distributed
throughout the surveyed area; L. tetraphylla, C.
aphrodite, and hemistoma were dominant in the three
environments sampled. Uniformity in the distribution of
planktonic cnidarian species is related to their high
adaptability (Gili et al. 1988). This would explain, at least
partially, the wide distribution of these medusae in the
Mahahual reef area, and probably along the western
Caribbean coasts. However, other groups, such as
copepods, show a sharp difference in composition and a
higher density within the reef lagoon than outside (Alvarez-
Cadenaetal. 1998).

In the Mexican Caribbean, a number of oceanic
medusae reach neritic and even estuarine waters (Suarez-
Moralesetal. 1997,Su^rez-Moraleselal. 1998). This has
been shown also for other zooplankton groups (Sudrez-
Morales and Gasca 1996). According to the results of
Merino ( 1 986) with drifting bottles, planktonic organisms
transported northward by the western edge of the Yucatan
Current tend to drift inshore. This would explain the

29

Suarez-Morales et al.

presence of oceanic medusae over the innermost portions
of the narrow shelf. A relevant factor in the mesoscale
distribution of zooplankton along the western Caribbean
is the strong effect of tidal currents (Greer and Kjerfve
1982, Kjerfve 1982, Kjerfve etal. 1982) which bring an
inflow of oceanic water to the lagoon through the channels
and over the reef crests. This has been reported also for
Mahahual (Castellanos and Sudrez-Morales 1 997). There
is a strong import of oceanic fauna into the Mahahual reef
area, as reflected in the dominance of oceanic forms and
the high species richness over the fore-reef. This effect
has been described also in the general reef zooplankton
community at Carrie Bow Cay (Ferraris 1 982).

The two assemblages defined by the Bray-Curtis
Index showed a clear separation of the sampling stations
in the surveyed area. The first, which comprised all the
fore-reef stations, represents the primary influence of the
oceanic fauna over the reef front. At this point, separation
between the fore- reef and the reef lagoon seems to be
sharp. However, the second group, which included the
channel and reef lagoon stations, showed a secondary
oceanic influence. This was represented mainly by the
occurrence of the most common oceanic species in the
area, a much lower species richness, and the occurrence
of coastal species. Therefore, the main difference between
the fore-reef and the reef lagoon medusa communities is
the species richness, all areas being dominated by a few
oceanic species. Migration and exchange of water into
and out of the reef lagoon are seen to be relatively
unimportant in determining the across-reef medusa
community structure. This pattern is useful to describe
the small-scale distribution of the medusae across the reef
from day to day. Due to the expected uniformity of the
zooplankton community along this reef system (Suarez-
Morales and Rivera- Arriaga 1 998), this pattern is probably
valid along the entire reef system.

Apparently, the effect of tlie coastal countercurrent
prevents the formation of a distinct seaward gradient of
medusae, which is common in some other shelf-related areas
studied (Pages and Gili 1 992). The occurrence of euryhaline
medusae mixed with the oceanic ones has been reported also
by Aral and Mason (1 982) in the Strait of Georgia, and by Gili
et al. ( 1 988) in tlie Mediterranean.

From the known ecological affinities of the medusae
recorded at Mahahual reef, three general groups can be
recognized: 1) oceanic species (L, tetraphylla, S.
bitemaculata, A. hemistoma, R. velatuniy N. punctata,
Cyiaeis tetrastyla, and CarybJea marsupialis), which
represented 60% of the medusa population; 2) neritic/
coastal species (A. dinema, A. rugosuniy Obelia sp., C.
folleata and Clytia mccradyi), which accounted for 30%

of the medusae, and 3) coastal species (P. minima, Zanclea
costata). The groups showed an overlapping distribution
through the surveyed area.

Acknow ledgments

We received financial support from CONACYT (Projs. :
D1 12-904520 and 1 189-N9203) for the collection trip. We
gratefully acknowledge the support of L. Vasquez-
Yeomans, E. Sosa, andl. Castellanos for their participation
in this project.

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F.A.O, Dociimento Tdenico de Pcsca 175:1-107.

Suarez-Mo rales, E. and R. Gasca. 1996. Planktonic copepods
of Bahia de !a Ascension, Caribbean coast of Mexico, a
seasonal survey, Crustaceana 69: 1 62- 1 74.

Snarcz-Morales, E, and E. Rivera- Arrkga. 1998. Zoopbneton
c hidrodinamica en zonas litoralcsyartecifalesde Quintana
Roo, M6tico. Hidrobiotdgica, in press.

Suarez-Morales, E., L. Segura-Puertas and R. Gasca. 1995.
Medusas (L'nidaria: Hydrozoa) de ta Bahia de Chetumal,
Mexico (1 990- 1991). Caribbean Journal of Science 31:243-
251.

Suarez- Morales, E., L. Segura-Puertas and R, Gasca. 1998.
Medusan (Cnidana) assemblages off the Caribbean coast
of Mexico. Journal of Coastal Research 14, in press.

Sudrez* Morales, E., M.O. Zamponi and R. Gasca. 1997.
Hydromedusac (Cnidaria: Hydrozoa) of Bahia de la
Ascension, Caribbean coast of Mex ico: a seasonal survey.
Proceedings of the 6th International Conference on
Coetenteralc Biology 1995, National Naiuurhistorisch
Museum, Leiden, The Netherlands, 16-21 July 1995,
p. 465-472.

Thiel, M.E. 1935, Zur Kenntnis der Hydromedusenfaana des
Schwarzen Meeres. ZooiogiscKcrAnzeiger 3:161-174,

Visquez-Yeomans,L., U, Orddflez-Lopczand E'. Sosa-Cordero.
1998. Fish larvae adjacent to a coral reef in the western
Caribbean Sesoff MahahuaL Mexico. Bulletin of Marine
Science 62:245-261.

Zamponi, M.O. and E, Sudrez- Morales. 1991. Algunas
medusas del Mar Caribe Mexicano con la deseripcidn
de Tetraotop^rpa siankaan&nsis gen. et sp, nov.
(Narcomedusae: Aeginidac). Spheniscus 9:41.-46,

Zamponi, M.O., E. Suarcz-Moralcs and R. Gasca. 1990.
HidromedusasfCoelenterata: Hydrozoa) yescifomedusas
(Coelenterata; Scyphozoa) de la Bahia de la Ascensidn,
Reservade Sian Ka'an, In; D, Navarro and J.G. Robinson,
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Sian Ka*an, Quintana Roo. Mexico. CIQRO/PSTC. Unlv,
of Florida, Mexico, p. 99-107.

31

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

An Annotated Checklist and Key to Hermit Crabs of Tampa Bay Florida^ and
Surrounding Waters

Karen M. Strasser

University of Southwestern Louisiana

W Wayne Price

University of Tampa

DOI: 10.18785/grr.ll01.06

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

Part of the Marine Biology Commons

Recommended Citation

Strasser, K. M. and W. Price. 1999. An Annotated Checklist and Key to Hermit Crabs of Tampa Bay Florida, and Surrounding Waters.
Gulf Research Reports 11 (l): 33-50.

Retrieved from http://aquila.usm.edu/gcr/voll l/issl/6

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 editor of The Aquila Digital Community. For more information, please contact Joshua.Cromwell(®usm.edu.

Gulf Research Reports Vol. 1 1, 33-50, 1999

Manuscript received April 22, 1998; accepted August 20, 1998

AN ANNOTATED CHECKLIST AND KEY TO HERMIT CRABS OF
TAMPA BAY, FLORIDA, AND SURROUNDING WATERS

Karen M. Strasser^ and W. Wayne Price*

'Department of Biology, University of Southwestern Louisiana, Post Office Box 42451,
Lafayette, Louisiana 70504-2451, USA, Phone: 318-482-5403, email: kms@usl.edu
^Department of Biology, University of Tampa, Tampa, Florida 33606, USA, Phone: 8 13-253-
3323, email: wprice@alphaMtampa.edu

ABSTRACT Fourteen species of hermit crabs, belonging to 5 genera, were identified from 35 stations in Tampa
Bay and adjacent continental shelf waters. Ninety-two samples were taken from the intertidal zone to 1 5 m with
a variety of gear including dip nets, trawls, dredges, and hand collections using SCUBA. Pagurus maclaughlinae,
Pagurus longicarpus^ and Pagurus pollicaris w'ere distributed throughout the bay. These species were often
sy mpatric, and were commonly found inseagrass beds, sandy substrates, and sand/mud substrates, respectively.
Clibanarius vittatus, Pagurus gymnodactylus, and Pagurus stimpsoni inhabited the highcrsalinity waters of the
bay entrance. Pagurisies sp., Paguristes hummi, Pagurus impressus and Pelrochirus diogenes were collected
from the lower bay to offshore on hard substrates and sand. Paguristes puncticeps, Paguristes sericeus and
Pagurus carolinensis were collected only offshore on hard substrates. The latter species is reported from the
Gulf of Mexico for the first time. Isocheles wurdemanni appears to be restricted to high energy beaches. An
illustrated key as well as information on distribution, reproductive biology, taxonomic problems, symbionts,
and coloration are presented.

Introduction

Tampa Bay, the largest open-water estuary in Florida
(Tampa Bay National Estuary Program 1 996), supports a
rich diversity of invertebrates which often occur in high
densities (Simon 1 974). However, the hermit crab fauna of
this embayment and adjacent waters is poorly known.
Although prior to the present study 1 5 species of hermit
crabs were documented from the shallow waters ( 1 5 m or
less) of the west coast of Florida, only 5 have been
recorded from the Tampa Bay area (Table 1). The first
species reported was Pagurus pollicaris Say, 1817, by
I ves ( 1 89 1 ) near the entrance of the Manatee River, which
flows into Tampa Bay. Over 50 years later, Paguristes
hummi Wass, 1955, was collected in tidal pools at the
mouth of Tampa Bay. Provenzano (1959), in a major
taxonomic paper on the shallow-water hermit crabs of
Florida, cited only 1 species from the Tampa Bay area,
Pagurus longicarpus Say, 1817. In the most recently
published survey of macro in vertebrates of Tampa Bay,
Dragovitch and Kelley ( 1 964) found Pelrochirus diogenes
(Linnaeus, 1 758) as well as Pagurus longicarpus and P.
pollicaris. During the next 20 years, several systematic
accounts were published on hermit crabs from Florida
waters (Mclaughlin and Provenzano 1974a, 1974b,
McLaughlin 1975, Garcia-G6mez 1982, Lemaitre 1982,
Lemaitre et al. 1982), but they included no records from
Tampa Bay. McLaughlin and Gore (1988) reported P.
maclaughlinae Garcia-Gomez, 1 982 from Tampa Bay, in a
study on the larval development of this species.

The present study was undertaken to assess the
species composition and distribution of hermit crabs

inhabiting the Tampa Bay area, and provide an illustrated
key as an aid to their identification. In addition, information
on reproductive biology, coloration, and taxonomic
considerations is included.

Materials and Methods

More than 90 samples (over 850 specimens) of hermit
crabs were taken at 3 5 locations in the Tampa Bay, Florida,
area to a depth of 1 5 m (Figure I ). Most collections were
made by the authors from 1991-1 997; however, additional
material was examined from the University of Tampa
Invertebrate Collection and the Florida Marine Research
Institute, St. Petersburg, Florida. Specimens were collected
with a variety of gear types and techniques; these are
included in Appendix 1 with the station number (Figure 1),
bottom type, temperature, salinity, depth, and species
found at each station. Morphological terminology used
for identification in the key is given in Figure 2, Unless
otherwise noted, illustrations were prepared with the aid
of a dissecting microscope and drawing tube.

Synonymies (restricted to primary taxonomic
publications), material examined, distribution, and notes
on ecological and reproductive biology are provided for
each species in the systematic account. For species in
which detailed coloration notes are available in the
literature, only key color characters have been provided.
For the other species listed below, descriptions of
coloration for living specimens are reported for the first
time, or additional detail is given to supplement existing
notes. The material examined is presented in the following

33

Strasser and Price

manner: station number: date collected (number of
specimens). Ovigerous females are designated with an (o).
Collection dates followed by an asterisk indicate specimens
borrowed from the Florida Marine Research Institute, St.
Petersburg, Florida. Collections dates before 1991 that are
not followed by an asterisk are from the University of
Tampa Invertebrate Collection. Specimens collected during

the present study are deposited in the University of Tampa
Invertebrate Collection except forrepresentative specimens
of each species which are deposited in thehiational Museum
ofNatural History, Smithsonian Institution, Washington,
DC, (catalog number of specimens referred to as
Paguristes sp. is USNM 265379).

Table 1

Hermit crab species reported from the west coast of Florida (Florida/Alabama border south to Cape Sable) to a depth of
15 m. Species records contained in this table were compiled from published literature as indicated. Lemaitre et al. (1982)
concluded after a study of the species of the Provenzanoi Croup, the distribution of Pagurus annulipes did not include the
west coast of Florida. The authors did not examine Wass* material, and assigned his material to Pagurus

maclaughlinacf P. stimpsonij P. gymnodactylusy and/or P. criniticornls.

Location

Reference

Family Hiogenidae:

Clibanarius vittatus

Pensacola

Cooley 1978

St, Joseph Bay

Brooks and Mariscal 1985a

Sopchoppy

Hazlctl 1981

Alligator Harbor

Wass 1955

Tampa Bay

Present study

Little Gasparilla Pass

Ives 1891

Isocheles wurdemanni

Perdido Key

Rakocinski et al. 1996

St, George Island

Caine 1978

Alligator Harbor

Wass 1955; Provenzano 1959

Tampa Bay

Present Study

Paguristes hummi

Perdido Key

Rakocinski etal. 1996

Pensacola

Cooley 1978

Dog Island

Sandford 1995

Alligator Harbor

Wass 1955; Wells 1969

Clearwater Beach

Provenzano 1959

Tampa Bay

Wass 1955; Present study

Sanibel Island

Gunter and Hall 1965

Marco Island

Provenzano 1959

West Coast of Everglades

Rouse 1970

Paguristes puncticeps

Northwest Coast of Florida

Provenzano 1959

off Tampa Bay

Present study

Paguristes sericeus

off Horseshoe Cove

Provenzano 1959

off St. Petersburg Beach

Provenzano 1959

off Tampa Bay

Present study

Paguristes tortugae

Marco Island

Provenzano 1959; McLaughlin and Provenzano 1974a

Everglades

Rouse 1970

Paguristes sp.

Tampa Bay

Present study

Petrochirus diogenes

Pensacola

Cooley 1978

Alligator Harbor

Wass 1955

Tampa Bay

Dragovich and Kelley 1964; Present study

Everglades

Rouse 1970

Family Paguridae:

Pagurus annulipes ?♦

Alligator Harbor

Wass 1955

Pagurus hrevidactylus

St. Andrews State Park

McLaughlin 1975

Pagurus carolinensis

off Tampa Bay

Present study

34

Hermit Crabs of Tampa Bay, Florida

TABLE 1 (Continued)

Location

Family Paguridae (continued):

Pagurus gymnodactylus Perdido Key

Pensacola
Cedar Key
Anclole Anchorage
Tampa Bay
Marco Island

Pagurus impressus Pensacola

Dog Island
Alligator Harbor
Sea Horse Key
Clearwater Beach
Tampa Bay
Sanibei Island
Everglades

Pagurus longicarpus Perdido Key

Pensacola
St. Joseph Bay
Dog Island
Alligator Harbor
Panacea
Wakulla Beach
Cedar Key
Crystal River
Clearwater Beach
Tampa Bay
Sanibei Island
Rookery Bay
Everglades
Cape Sable

Pagurus maclaughlinae Crystal River

Anclote Anchorage
Tampa Bay
Estero Bay
Rookery Bay
Everglades

Pensacola
St. Joseph Bay
Dog Island
Alligator Harbor
Panacea
Cedar Key
Tampa Bay
Lemon Bay
Little Gasparilla Pass
Charlotte Harbor
Sanibei Island
Rookery Bay
Everglades

Anclote Anchorage
Tampa Bay

Iridopagurus caribbensis off Panama City

Reference

Rakocinski etal. 1996
Lemaitre 1982
Lemaitre 1982
Lemaitre 1982
Present study
Lemaitre 1982

Cooley 1978
Sandfordl995
Wass 1955; Wells 1969
Proven zano 1959
Provenzano 1959

Benedict 1892 (see Williams 1984); Present study
Provenzano 1959
Rouse 1970

Rakocinski etal. 1996
Cooley 1978

Brooks and Mariscal 1985a

Sandford 1995

Wass 1955; Wilber 1989

Wilber and Herrnkind 1982

Wilber and Herrnkind 1982, 1984; Wilber 1989

Provenzano 1959

Lyons etal. 1971

Provenzano 1959

Provenzano 1959; Dragovich & Kelley 1964; Present study
Provenzano 1959; Gunter and Hall 1965
Sheridan 1992
Rouse 1970

Tabb and Manning 1961

Garcia-G6mez 1982
Lemaitre etal. 1982

McLaughlin and Gore 1988; Present study
Garcia-G6mez 1982
Sheridan 1992
Garcia-Gdmez 1982

Cooley 1978

Brooks and Mariscal 1985a, 1985b
Sandford 1995
Wass 1955; Wells 1969
Brooks 1989
Provenzano 1959

Ives 1891; Dragovich and Kelley 1964; Present study
Provenzano 1959
Provenzano 1959
Provenzano 1959

Provenzano 1959; Gunter and Hall 1965
Sheridan 1992
Rouse 1970

Lemaitre etal. 1982
Present study

Williams 1984

Pagurus poUicaris

Pagurus stimpsoni

35

Strasser and Price

Figure 1. Location of collection sites in the Tampa Bay area.

Key to the Hermit Crabs of the Tampa Bay Area

1. Third maxillipeds approximated at base (Figure 3a)
[Family Diogenidae] 2

Third maxillipeds widely separated at base (Figure 3b)
[Family Paguridae] 8

2. No paired appendages present on first 2 abdominal

segments of either sex; dactyl of fourth pereopod

subterminal (Figure 3e) 3

Paired appendages present on first 2 abdominal

segments of male (Figure 3c), and first only of female
(Figure 3d); dactyl of fourth pereopod terminal (Figure 3f)
5

3. Chelipeds dissimilar and unequal, right slightly larger

than left, right with calcareous tip (Figure 4a)

Petrochirus diogenes

Chelipeds similar and subequal, both with corneous
tips (Figures 4b, c) 4

4. Finger tips spooned (Figure 4b); antennal flagellum

long and not setose Clibanarius vittatus

Finger tips accuminate (Figure 4c); antennal flagellum

short and very setose (Figure 4d)

Isocheles wurdemanni

5. Rostrum broadly rounded or pointed, not extending
beyond lateral projections of cephalic shield (Figure 4e)

Paguristes hummi

Rostrum slender and clearly extending beyond level of

lateral projections (Figures 4f, g, h) 6

6. Ocular acicles ending in more than one terminal spine
(Figure4f) Paguristes sp.

Ocular acicles ending in simple spine (Figures 4g, h)
7

7. Anterior and lateral margins of cephalic shield meeting

at broadly obtuse angle (Figure 4g)

Paguristes puncticeps

Anterior and lateral margins of cephalic shield

meeting at near right angle (Figure 4h)

Paguristes sericeus

8. Ocular acicles ending in more than one spine or with

submarginal spines (Figure 4i)

Pagurus carolinensis

Ocular acicles ending in a single terminal spine or with
subterminal spine (Figure 4J) 9

36

Hermit Crabs oe Tampa Bay, Florida

9. Antennal flagellum with paired setae, 3-8 articles in
length, at least every second article proximally, decreasing

in length distally (Figure 4k)

Pagurus gymnodaciylus

Antennal flagellum with setae 1 article in length or less
(Figure 41), or irregularly short and long setae over entire
length 10

10. One or both chelipeds broad, right chela dorsoventrally

flattened (Figures 4m, n) 11

Roth chelipeds narrow, right chela not dorsoventrally
flattened (Figures 4o, p, q) 12

1 1 . Dactyl of right cheliped with sharply produced angle
on outer margin; lacking depression on dorsal surface of

proprodus of either cheliped (Figure 4m)

Pagurus pollicaris

Dactyl of right cheliped without sharply produced
angle on outer margin; with depression on dorsal surface

of proprodus of both chelipeds (Figure 4n)

Pagurus impressus

12 Dactyls of 2nd and 3rd pereopods each withoutrow of
corneous spines on ventral margin (Figure 4r); eyestalks

short, length approximately 3 times the width

Pagurus longicarpus

Dactyls of 2nd and 3rd pereopods armed with row of
strong corneous spines on ventral margin (Figure 4s);

eyestalks long, length at least 4 times the width

13

1 3. Left chela with longitudinal ridge on dorsal surface of
propodus, unarmed or with weak spines or turbercles

(Figure 4p) Pagurus stimpsoni

Left chela without ridge on dorsal surface of propodus,
midline armed with a single or double row of strong spines
(Figure 4q) Pagurus maclaughlinae

Systematic Account

Family Diogenidae Ortmann, 1 892
Clibanarius vittatus (Bose, 1802)

Pagurus vittatus . — Bose 1802:78, Plate 12, Figure 1.

Clibanarius vittatus. — Stimpson 1 8 62 : 8 3 . — Hay and
Shore 19 18:4 10, Plate 30, Figure 9. — ^Provenzano 1959:371,
Figure 5D.—Holthuis 1959: 1 41, Figures 26,27. — Williams
1965:120, Figure 97. — Forest and de Saint Laurent
1 967 : 1 04 Coelho and Ramos 1 972: 1 70 .—Felder 1973:32,
Plate 3, Figure 20. — Williams 1984:194, Figure 135. —
AbeleandKim 1986:29,339d,e,

Material. Station 14:3 Aug 1993(1). — Station 20:25
June 1993(1).— Station23:May 1973(2).

Figure2. Schematic drawing of a hermit crab in dorsal view
(after McLaughlin 1980)

Known range. Potomac River, Gunston, Virginia, to
Florianopolis, Santa Catarina, Brazil (Forest and de Saint
Laurent 1967).

Remarks. Only 4 specimens of C. vittatus were
collected at the mouth of Tampa Bay in seagrass, sand/
mud and rock jetty habitats. This species is commonly
found in shallow subtidal and intertidal zones of harbor
beaches, mud flats (Pearse et al. 1942), rock jetties, bay
shores (Whitten et al. 1950), salt marshes near the ocean
(Heard 1982), and seagrass- sand/mud areas (Lowery and
Nelson 1988). Although C. vittatus is euryhaline (10-
35%o) (Heard 1 982), it is more commonly found at higher
salinities, which may be necessary for egg development
(Lowery and Nelson 1988). Although higher salinity
habitats were sampled at different seasons in the present
study, few animals were found. Thus, it appears that C
vittatus is uncommon in the Tampa Bay area.

Ovigerous females of C. vittatus ware reported from
North Carolina in June (Kircher 1 967), South Carolina in
July and August (Lang and Young 1977), east coast of
Florida from A pril-September (Lowery and Nelson 1988),
southern Florida in October (Provenzano 1959),
northwestern Florida in June (Cooley 1978) and Texas
from May-August (Fotheringham 1975). No ovigerous
females were collected during this study.

Coloration. Light longitudinal stripes on the second
and third pereopods. See Provenzano (1959) for additional
detail.

37

Strasser and Price

Figure 3. a) Third maxillipeds of Diogenidae, b) third maxillipeds ofPaguridae (a and b redrawn from Provenzano 1961),
c) PaguristeSy ventral surface of male, gonopores on coxa of Fifth pereopods, d) Pagurisies, ventral surface of female, Mxp
3 = coxa of third maxilliped, gonopores on coxa of third pereopod, e) Clibanarius vittatuSy distal end of fourth pereopod,
dactyl subterminal (scale -2.5 mm), f) Pagurisies 5er/c<ms, distal end of fourth pereopod, dactyl terminal (scale -2.5 mm).

Isocheies wurdemanni Stimpson, 1862 Petrochirus diogenes (Linnaeus, 1758)

Cancer diogenes — Linnaeus 1758:63 1 .

Cancer bahamensis — Herbst 1796:30.

Petrochirus granvlatvs — Stimpson 1 859:234.
Petrochirus bahamensis — ^Benedict 1 90 1 : 1 40. — Hay
and Shore 19 18:4 10, Plate 30, Figure 6. — Schmitt 1935:206,
Figure 66. — Provenzano 1959:378, Figure 8.— Provenzano
1961:153.

Petrochirus Holthuis 1 959: 1 5 1 . — Williams

1965:122, Figure98. — Provenzano 1968: 147, Figures. 1-
12. — Felder 1973:30, Plate 3, Figure 14. — Williams
1984:198, Figure 138.— AbeleandKim 1986:3 l,353e,f.

Materia). Station 10: 28 May 1966*(1). — Station 14:
23 Jan. 1993(1).— Station 23: 9 Feb. 1965* (1).— Station 26:
8May 1983 (3), 24 Oct. 1992(1).— Station 27: May 1978(1),
30 Aug. 1980(1).— Station 30: 2 Oct. 1993(1),

Known range. Off Cape Lookout, North Carolina,
through Gulf of Mexico and West Indies south to off Ilha
de S3o Sebastiao, Brazil, 23‘’42.5' S, 45“ 14.5' W (Forestand
de Saint Laurent, 1967).

Remarks. Petrochirus diogenes is rare in shallow
waters of the Tampa Bay area. Most specimens were
collected on sand near hard substrates at the mouth of
Tampa Bay or in offshore waters. This species has been
reported on mud, mud/shell and sand bottoms in

Isocheies wurdemanni — Stimpson 1862:85. —
Provenzano 1959:375, Figure7. — Felder, 1973:32, Plate 3,
Figure 2 1 . — Abele and Kim 1986:29, 353d.

Material. Station 28: 1 June 1991 (3).

Known range. Texas, Louisiana, west coast ofFlorida
and Venezuela (Provenzano 1959).

Remarks. Whereas this species was only collected in
shallow offshore waters along the high energy beaches of
Anna Maria Island, it is probably found in similar habitats
along the entire west coast of Florida. This is consistent
with observations made by Caine (1978) who studied
activities of I. wurdemanni along the Gulf of Mexico beaches
of St. George Island, Florida. In his study, the majority of
specimens were collected within 3 m of the splash zone or
on the beach side of sand bars, 20-50 m offshore. Peak
abundances were reported in the fall and spring with
densities reaching 286 m‘^ along the offshore sand bars.

Ovigerous females of /. wurdemanni were reported
from St. George Island, Florida, in the months of May,
June, September, Octoberand November (Caine 1978). No
ovigerous females were collected in the present study.

Coloration. Body color white, see Stimpson (1 859),
Wass ( 1 955), and Provenzano ( 1 959) for additional detail.

38

Hermit Crabs of Tampa Bay, Florida

Figure 4. Hermit crabs of the Tampa Bay area, a) Chelipeds of Peirochirus diogenes^ b) chelipeds of Clibanarius vlttatus,
c) chelipeds of hocheles wurdemanni^ d) antennal flagellum of Isockeles wurdemanni, e) cephalic shield and ocular acicles
of Pagurisies hummiy 0 cephalicshieid and ocular ocitlesof PagurisUs&p., g) cephalic shield and ocular acicles of Paguristes
puncticepsy h) cephalic shield and ocular acicles of Paguristes sericeusy i) ocular acicles of Pagurus caroiinensiSy j) ocular
acicles of Pagurus maclaughlinaey k) antennal peduncle of Pagurus gymnodactylusy I) antennal peduncle of Pagurus
maclaughiinaey m) right cheliped of Pagurus pollicariSy n) right cheliped of Pagurus impressusy o) right cheliped of Pagurus
longicarpuSy p) left cheliped of Pagurus stimpsoniy q) left cheliped of Pagurus maciaughUnaey r) dactyl and propodus of second
pereopod of Pagurus iangicarpuSy s) dactyl and propodus of second pereopod of Pagurus maclaughlinae. Scales equal 2 mm
for k and 1 and 1 mm for all other illustrations.

39

Strasser and Price

continental shelf waters on the Tortugas shrimping
grounds (Provenzano 1959), off Mississippi (Franks et al.
1972), on brown shrimp grounds in the western Gulf of
Mexico (Hildebrand 1954), and has been found as deep as
1 28 m (Wenner and Read 1 982). It may be fairly common
in deeper continental shelf waters off Tampa Bay.

Ovigerous females were reported in June from Texas,
in August from west Florida (Provenzano 1968), and in
March from the Virgin Islands (Provenzano 1961). No
ovigerous females were found during this study.

Coloration. Body color generally reddish with color
fading at joints. Antennal flagellum with red and white
bands; cornea blue and black. See Provenzano ( 1959) for
additional detail.

Paguristes hummi Wass, 1955

Paguristes hummi — Wass 1955: 148, Figures 1-4. —
Provenzano 1 959:38 1 , Figure 9. — Felder 1973:3 1 , Plate 3,
Figure 16. — Williams 1984:200, Figure 139. — Abeleand
Kim 1986:30, 343a. — Campos and Sanchez 1995:576,
Figure 7.

Material. Station 13: 26 Sept. 1992(1). — Station 14:13
June 1993(1).— Station 16: 120ct. 1983(1).— Station 18:
20ct. 1993(3). — Station 24: 3 Jan. 1966*(1). — Station25:
31 May 1 966* (1).— Station 27: 1 Sept. 1991 (1).— Station
28: 1 June 1991 (4), lOct. 1991 (1).— Station 30: 2 Oct. 1993
( 1 ), 29 April 1 994 (4).— Station 3 1 : 26 July 1 995 (4, 1 o).

Known range. Newport River, North Carolina, to
Sapelo Island, Georgia; Marco Island, southwestern
Florida, to off Isles Dernieres, Louisiana (Williams 1 984);
Caribbean coast of Colombia (Campos and Sanchez 1 995).

Remarks. Paguristes hummi found both offshore
and in lower Tampa Bay, usually associated with hard
substrates. Wass (1955) reported P. hummi inhabiting a
variety of gastropod shells in the intertidal zone only on the
south side of Mullet Key at the mouth of Tampa Bay where
it was abundant at times. This species was found in shelly
areas of Beaufort, North Carolina, but was more abundant
offshore on rocky outcrops (Kellogg 1971). In the Alligator
Harbor-Dog Island area of northwest Florida, P. hummiha^
been found to inhabit sponges (Wass 1955, Wells 1969,
Sandford 1995), which have been identified as the hermit
crab sponge Spongosorites suberitoides (Sandford and
Kelley-Borges 1 997). All specimens collected in the present
study were found in gastropod shells.

Ovigerous females of P. hummi were reported from
northwestern Florida in January and July (Cooley 1978),
and from southwestern Florida in February (Provenzano
1959), October, and November (Rouse 1970). The only
ovigerous female collected in this study was taken in July.

Coloration. Body color generally white. Striking blue
color mark, ringed by black and yellow, present on the

inner surface of the merus of both chelipeds. See Wass
( 1955) and Provenzano ( 1 959) for additional detail.

Paguristes puncticeps Benedict, 1901

Paguristes puncticeps— Benedict 1 90 1 : 144, Plate 4,
Figure4,Plate5, Figure 2. — Provenzano 1959:384, Figure
10a. — Abeleand Kim 1986:30, 347e. — Campos and S^chez
1995:572, Figure2.

Material. Station 25: 31 Dec. 1966*(1). — Station 26:
8May 1983 (l),40ct. 1992(1), 19 April 1997(1, lo).—
Station 27: 1 May 1978(1), Oct. 1981 (1), 1 Sept. 1991 (2).^
Station 30: 2 Ocl. 1993 (4).

Known range. Northwestern Florida; south Florida to
Jamaica, probably throughout the West Indies (Provenzano
1 959); Caribbean coast of Columbia (Campos and Sanchez
1995); off Tampa Bay, Florida (present study).

Remarks. This report of P. puncticeps is the first
from a locality that occurs between northwestern Florida
and Miami and is indicative of a probable continuous
distribution of the species along the west coast of Florida
and throughout the Caribbean Sea. This species was only
found offshore of Tampa Bay in association with hard
substrates in depths of 10-15 m. Paguristes puncticeps
has been collected as deep as 19 m from the fortugas
shrimp grounds (Provenzano 1959). One ovigerous female
was collected in April during the present study, and one
was reported from Cuba in January (Provenzano 1959).

Paguristes sericeus and P. puncticeps are
morphologically similar species and were collected
together in continental shelf waters off Tampa Bay. Some
confusion exists in the literature concerning the length of
the antennal peduncles in relation to the antennal acicles
for these 2 species. All illustrations except Figures 93a
and 142a of Williams ( 1965, 1 984), respectively, show the
relationship of these characters to be similar in both
species: the antennal peduncle is slightly longer than the
antennal acicle (Milne Edwards and Bouvier 1 893, Benedict
1901, Provenzano 1959). The relationship of these
characters is not mentioned in descriptions of either
species (Milne Edwards 1 880, Milne Edwards and Bouvier
1 893, Benedict 1901, Provenzano 1 959), with the exception
of Williams ( 1 965, 1 984) who states correctly, “Antennal
peduncles slightly exceeding acicles.” However, an error
exists in Figure 93a (Williams 1 965, reproduced as Figure
I42a in Williams 1984), In these figures the antennal
peduncle of P. sericeus is shown to be considerably
shorter than the antennal acicle. Abele and Kim (1986)
used this inaccurate illustration along with a probable
misinterpretation of the word “acicle” in the, passage
above as a basis for separating P. sericeus and P.

40

Hermit Crabs of Tampa Bay, Florida

puncticeps. They appear to have interpreted Williams’
use of “acicle” to mean ocular acicle, whereas he was
instead referring to the antennal acicle in that section.
Using this interpretation and Williams’ illustration, P.
puncticeps appears to have a much longer antennal
peduncle in relation to the ocular acicle than does P.
sericeus. However, since the relationships among the
lengths of the antennal peduncle, antennal acicle and
ocular acicle are similar for both species, these characters
cannot be used to distinguish them.

As indicated in couplet 7 of the key and Figures 4g,h
of the present study, the shape of the antero^ lateral
margins of the cephalic shield appears to be the most
reliable character which separates P. sericeus from P.
puncticeps. Provenzano (1959) discussed the contrast
between the sloping angles of the shield in P. puncticeps,
and the near right angles found in P. sericeus {=P.
rectifrons sensu Provenzano). The presence of white
spots on the ocular peduncles of fresh P. puncticeps is
also mentioned by Provenzano as a differentiating
characteristic. However, this color pattern is not always
present in live material and should be used with caution.

Coloration. Body color red with white spots. At
times, juveniles bright red and adults rust red. Ocular
peduncles reddish orange, usually with white spots;
cornea bright blue. Antennular and antennal flagella
reddish. Proximal and distal ends of each segment lighter
in color than middle on all walking legs; setae fringing
dorsal and ventral areas occasionally green from
accumulation of algae. See Provenzano ( 1 959) for additional
coloration notes.

Paguristes sericeus Milne Edwards, 1880

Paguristes sericeus — Mi Ine Edward s 1 8 80 :44 . — Milne
Edwards and Bouvier 1893:46, Plate 3, Figures 14-22. —
Provenzano 1961:155. — Williams 1965:1 17, Figure 93. —
Provenzano and Rice 1966: 54, Figures 1-10. — Felder
1973:32, Plate 3, Figure 19. — PequegnatandRay 1974:242,
Figure 44, — Williams 1984:203, Figure 142. — Abeleand
Kim 1986:30, 347c, d.

Paguristes tenuirostris — Benedict 1 90 1 : 1 43 , Plate 4,
Figure 1.

Paguristes rectifrons — Benedict 1901 : 145, Plate 4,
Figure 7.

Material. Station 26: 8 May 1983 (3), 30 Apr. 1995
(2).— Station 27; 1 Sept. 1991 (1).

Known range. Off Cape Lookout, North Carolina;
West Flower Garden Bank, northwest Gulf of Mexico to
the Virgin Islands (Williams 1 984).

Remarks. This species was collected only offshore
of Tampa Bay on sand near limestone outcroppings at a
depth of 15 m. Paguristes sericeus has been found on
sand and coral rubble (Provenzano 1 96 1 ) at depths of9 to
145m(Williamsi984).

Ovigerous females were reported from off St.
Petersburg Beach, Florida, in July (Provenzano 1 959), on
the Dry Tortugas shrimping grounds in March and May
(Provenzano 1959, Rice and Provenzano 1965), and in the
V irgin Islands in March and April (Provenzano 1 96 1 ). No
ovigerous females were collected during the present study.

For taxonomic considerations see remarks under P.
puncticeps.

Coloration. Similar to P. puncticeps, except overall
color generally more orange-red, and eyestalks without
white spotting. See Provenzano ( 1 959, 1961), Provenzano
and Rice (1966), and Williams (1984) for additional
coloration notes.

Paguristes sp.

Material. Station 13:2 Mar. 1 99 1 ( 1 ), 26 Sept. 1 992
(6).— Station 14: Apr. 1979(1), 18 June 1992(1, 3o), 3 Aug.
1993(1).— Station 26:24 0ct. 1992(3), 19 Apr. 1997(1).—
Station27: 1 Sept. 1991 (7).— Station 29: 120ci. 1991 (2).—
Station 30: 2 Oct. 1993(3).— Station3 1:26 July 1995(3, lo).
— Station 33: 29 Sept. 1996(3, lo),

Remarks. These specimens appear to be of an
undescribed species most similar to Paguristes tortugae
Schmitt, 1 933 . The most obvious differences occur in the
color patterns. Paguristes tortugae has reddish-purple,
transverse bands on the pereopods whereas our
specimens arc unbanded with a brownish-green body
color (see coloration section). Future work with these
species should yield additional characters for their
distinction.

Paguristes sp. is relatively common in lower Tampa
Bay, especially near Bishop Harbor (station 13) where it
was often found in large groups on or near basket sponges.
It was rarely taken offshore, but was found near hard
substrates in all collections. Ovigerous females were
found in the summer and fall.

Coloration. Cephalic shield green or brownish-green
with yellowish-orange and white spots; posterior part of
thorax pinkish with irregular red spots and occasionally
blue patches laterally; area postero-medial to cephalic
shield yellowish-orange with green and white patches;
posterior border of carapace red. Proximal one-fourth of
ocular peduncles brown or greenish-brown, distal part
white, circumscribed with one proximal orangish-yellow
and one distal dark brown band; cornea black. Proximal

41

Strasser and Price

half of ocular acicles brown, distal half white. Antennular
peduncles marked with 3 brown or brownish-green and
white bands; flagella brown. Antennal peduncles brown
with white spines, distal segments circumscribed with 2
brown and 2 white bands; flagella colorless, every other
article white distally, middle part of each article solid
brown or with brown streaks laterally. Third maxillipeds
with brown and white bands. Chelipeds with dactyls and
fixed fingers yellowish, proximal part of propodi and
remaining segments greenish-brown; proximal one-half
of dactyls and three-fourths of propodi with reddish,
white-tipped tubercles or spines; spines on dorsomesial
margins of propodi and carpus reddish proximally, followed
by yellow rings and brown tips; merus with yellowish
reticulations and white dots mesially and laterally, and
reddish-orange patches along dorsal margin, Pereopods
generally greenish-brown with white or bluish-white spots
and reticulations; dactyls with brown spines, other articles
with reddish, white-tipped spines; carpi with dorsal one-
half reddish proximally. Abdomen yellowish with red
patches and white spots; transverse blue streaks laterally.

Family Paguridae Latreille, 1803
Pagurus caroUnensis McLaughlin, 1975

Pagurus near bonairensis — Pearse and Williams
1951:143.

Pagurus brevidaciyluS' — Provenzano 1959:413,
Figure 20 .—Williams 1965:132, Figure 107.

Pagurus caroUnensis — McLaughlin 1975:365,
Figures4-6. — Lemaitreetal. 1982:677, — Williams 1984:212,
Figure 150. — ^AbeleandKim 1986:33, 375f,g.

Material. Station 26: 24 Oct. 1992 (1), 4 Mar. 1997
(1).— Station 27: Oct. 1991 (1).— Station 30: 2 Oct. 1993
(3).— 3 1:26 July 1995 (2).

Known range. Off Newport River (Kellogg 1971) and
Cape Lookout, North Carolina, to southeastern Florida
(Williams 1 984); off Tampa Bay, Florida (present study).

Remarks. This is the first record of P. caroUnensis
in the Gulf of Mexico, Only 6 specimens were collected
offshore in association with hard substrates at depths of
5-15 m. This species has been reported to prefer hard
bottom in areas of good water circulation (Provenzano
1959) at depths of 2 to 53 m (Lemaitre et al. 1982).

Ovigerous females were reported in June, July, and
August from North Carolina, November, July-October in
Georgia and March-August in Florida( Williams 1 984). No
ovigerous females were collected in the present study.

Pagurus caroUnensis^ reported from the Gulf of
Mexico for the first time in the present study, is
morphologically very similar to P. brevidactylus. A\\hQ\iL^

this latter species was not found in the Tampa Bay area,
it occurs in northwest Florida. Future studies may
document an overlap in the ranges of these 2 species in
the Gulf of Mexico similar to their overlap in southeast
Florida (Lemaitre et al. 1982). The spination of the left
chelae may be used to separate these 2 species. Pagurus
hrevidaciylus (Stimpson, 1 859) has a longitudinal row of
strong or moderately strong spines near the dorsolateral
margin of the propodus, while P. caroUnensis may have
small or no spines in this area. In addition, P. brevidactylus
has shorter setae on the articles of the antennal flagella
and longer, more slender ocular peduncles than P.
caroUnensis. Coloration may be used to separate live
specimens of these species. Pagurus brevidactylus has
dark green to brownish black continuous stripes on the
pereopods, and striped chelipeds. Pagurus caroUnensis
has rust red to maroon stripes on the pereopods that do
not extend to the distal and proximal margins of each
segment, and the chelipeds are not striped (Lemaitre et al.
1982).

Coloration. See remarks above. Additional coloration
notes are found in Provenzano [1959 (=P. brevidactylus)].

Pagurus gymnodactylus Lemaitre, 1982

Pagurus annuUpes — Felder 1973:26, Plate 3, Figure
4 [notP. annw/jpes (Stimpson)].— Williams 1974:41.

Pagurus gymnodactylus — Lemaitre 1 982:657, Figures
1,2, 4c, d, 5a, b. — Lemaitre et al. 1 982: 687. — Abele and Kim
1986:33,377h,iJ.

Material, Station 14:3 Aug. 1 993 (4). — Station 18:2
Oct. 1 993 (8, 1 o).— Station 32 : 26 July 1 995 ( 1 ).

Known range. Gulf of Mexico from Mexico to west
coast of Florida (Lemaitre et al. 1982).

Remarks. Pagurus gymnodactylus was collected on
sand and hard substrates in shallow subtidal depths at the
mouth of Tampa Bay. This species has been found from
the subtidal zone to 1 9 m (Lemaitre et al. 1 982).

No information is available on the reproduction of
this species. However, in the present study, one ovigerous
female was found in October.

Coloration. While some specimens appeared to be
almost completely white, those with color displayed the
following characteristics: carapace mottled yellow-brown,
occasionally with green and red splotches, red flecks
laterally. Abdomen transparent blue. Ocular acicles,
eyestalks, and antennular flagella transparent with red
and white flecks; eyestalks sometimes with central,
horizontal, blue-green band. Antennal flagella transparent,
marked with white every 2-5 articles; peduncle transparent
with red and white flecks. First and second maxillipeds

42

Hermit Crabs of Tampa Bay, Florida

mottled red and white at bases. Third maxillipeds with blue
to red transverse bands. Merus, carpus and propodus of
right cheliped mottled brown, distal part of propodus and
dactyl white. Dactyls, propodi, carpi, and meri of second
and third pereopods with mottled brown transverse bands.

Pagurus impressus (Benedict, 1892)

Eupagurus impressus — Benedict 1 892:5.

Pagurus impressus — Provenzano 1959:399, Figure
15 —Williams 1965: 129, Figure 104.— Felder 1973:27, Plate
3, Figure 9. — Williams 1984:2 15, Figure 153. — ^Abeleand
Kim 1986:33, 377a,b,c.

Material. Station 13: 26 Sept 1992(4). — Station 14:
Apr. 1982 (3), May 1983(1 ), 23 Jan. 1 993 (25+o), 3 Aug.

1993 (25+), 1 1 Sept. 1993(2).— Station 15: May 1983 (2).—
Station 16: 120ct. 1983(1).— Station 19: 19Feb. 1982(2),
25 June 1993(1),— Station 28: 1 Oct. 1 990 (2).— Station 29:
120ct. 1991(25+).— Station 30:2Oct 1993 (6), 29 April

1994 (2).— Station 3 1 :26 July 1995 (9).

Known range. NorthCarolinatoCape Canaveral, Florida;
Florida Bay north to Pensacola, Florida; Port Aransas, Texas
(Williams 1984); Padre Island, Texas (Felder 1973).

Remarks. This species is very common at the mouth
of Tampa Bay and in shallow offshore waters. It was often
found in congregations on sand near hard substrates.
Pagurus impressus has been reported to inhabit areas of
sand, seagrass beds or pilings, and has been found in
hermit crab sponges (Wass 1955, Wells 1969, Sandford
1995). In the Dog Island area, P. //wpre-wus has been shown
to move into the intertidal zone close to the shoreline in
January, with many individuals inhabiting the hermit crab
sponged. (Sandford and Kelley-Borges 1997).

Ovigerous females were collected from the Carolinas
and Georgia in January and February, and in Florida in
February and April (Williams 1984), In the present study,
ovigerous females were collected in January only.

Coloration. Eycstalks dark brown with white specks
on dorsalsurface, red at base, and longitudinal blue stripe
on ventral surface. Cornea black with translucent yellow
covering. Antennal and antennular flagella yellow,
sometimes red at base. Cephalic shield mottled yellow and
brown. Thorax generally reddish-brown with white spots;
laterally, darker red with white spots. Third maxillipeds
brown with white spots, white at joints. First and second
maxillipeds reddish with white spots. Propodi and dactyls
of chelipeds almost solid brownish-orange to rust-red on
dorsal surface, sometimes with small white spots, ventral
surface darker brown with white spots; carpi and meri
mottled dark brown with white transverse bands. Dactyls
of second and third pereopods mottled brownish orange,

with thin longitudinal stripe on lateral and mesial faces;
propodi, carpus and meri mottled brown, with white
transverse bands near joints. Joint between carpus and
merus of all walking legs reddish in color. See Provenzano
( 1 959) for additional coloration notes.

Pagurus longicarpus Say, 1817

Pagurus longicarpus — Say 1817:163. — Hay and
Shore 1918:411. — Provenzano 1959:394, Figure 13. —
Williams 1965: 125, Figure 101. — Felder 1973:27, Plate 3,
Figure 7. — Williams 1984:216, Figure 154. — Abeleand
Kim 1986:33, 38 lc,d,e.

Material. Station 1 : May 1 986 ( 1 8), 5 Feb. 1 99 1 (24),

1 3 Jan., 1992 (2), 23 June 1992 ( 10), 1 Sept. 1992 ( 12),2 1 Jan.
1993 (5), 29 Nov. 1993 (6).— Station 2: 5 May 1977(7),—
Station3: 1 Feb. 1992 (4), 5 May 1992(5), 18 June 1992(10),
19 Sept. 1992(6), 13 Jan. 1993, 11 May 1993 (6, lo).—
Station4: 1 1 Nov 1991 (2), 4 Jan. 1993(4).— Station 5: 26
Sept. 1976(1), 28 Sept. 1976(2),Sept. 1991 (3), 16 Jan. 1993
(3), 1 1 May 1993 (9).— Station 6: 8 June 1978(12).— Station
9: 1 8 Sept. 1992 (2), 6 Jan. 1 993 ( 1 ), May 1993 (3).— Station
12:7May 1983(1).— Station 14:Oct. 1979(6), 3 Aug. 1993

(6) , 1 1 Sept. 1993 (4).— Station 17: 31 Dec. 1964* (1).—
Station 18: 1 1 Dec. 1965* (8).— Station 19:2Nov. 1991

(7) . — Station 20: 8 Jan. 1965* (4). — Station 21 : 9 Feb.

1 965* ( 1 ).— Station 22: 25 June 1 993 .

Known range. Minas Basin and Chignecto Bay, Nova
Scotia (Bousfield and Liem 1960) to Hutchinson Island,
Florida (Camp et al. 1977); southwestern Florida to the
coast of Texas (Whitten et al. 1950, Provenzano 1959,
Rouse 1970).

Remarks. Pagurus longicarpus is commonly found
on sand, sand/mud, grass, and hard substrate habitats
throughout the intertidal and shallow subtidal waters of
the entire Tampa Bay area. This species has been reported
from harbor beaches and channels on a variety of
substrates (Williams 1984), from the intertidal to 200 m
(Wenner and Boesch 1979). Its ubiquity in bays and
estuaries prevents its use in distinguishing shallow water
habitats (Alice 1923).

Ovigerous females of P. longicarpus were collected
from April-Septcmber in Massachusetts (Carlon and
Ebersole 1995), February-September in North Carolina,
March-July in Georgia (Williams 1 984), September- April in
Florida (Wass 1955, Dragovich and Kelley 1964, Lyons et
al. 1 97 1 ), and winter in Texas (Fotheringham 1975). In the
present study, ovigerous females were collected in May.

Coloration. Abdomen and thorax brown, sometimes
with white spots on cephalic shield. Ocular acicles white,
eyestalks white with brown near black corneas, Antennular

43

Strasser and Price

peduncles brown and white; flagella white. Antennal
peduncles and acicles brown; flagella brown with white
article every 2-4 articles. Maxillipeds brown proximally.
Right cheliped white or off-white, with 3 longitudinal
brown, rust or yellowish-brown stripes; stripes joined at
merus, then separated distally on mesial, dorsal, and
lateral margins. Second and third pereopods with
longitudinal stripe on lateral and mesial faces. See
Provenzano (1959) for additional coloration notes.

Pagurus maclaughlinae Garcia-G6mez, 1982

lEupagurus annulipes — Ives 1891:193. [not E.
annulipes Stimpson].

Pagurus annulipes — Schmitt 1935:205 (in part). —
Provenzano 1959:407, Figure 18 [not P. annulipes
(Stimpson)]. — Williams 1965: 130 (in part), Figure 105. —
Forest and de Saint Laurent 1967: 127 (in part).

Pagurus bonairensis~—¥Q\AQX \ 9iy2(> (in part), Plate
3, Figure 5. [notP. bonairensis Schmitt].

Pagurus maclaughlinae — Garcia-Gomez 1 982:647,
Figures 1,2. — Lemaitreetal. 1982:691. — AbeleandKim
1 986:33, 377d,e,f,

Material, Station 1: 13 Jan. 1992 (1), 21 Jan, 1993
(25+).— Station 3: 28 Jan. 1992 (25 +o), 1 Feb. 1992(1), 28
Feb. 1992(25+), 5 May 1992(25+), 18 June 1992(1), 13 Jan.
1993(25+), 1 1 May 1993 (25+).— Station 5: 18 Sept. 1992
(25+), 16 Jan. 1 993 (2 5 +o), 11 May 1993(25+).— Station 9:
6 Jan. 1993(3).— Station I l:20ct. 1992(25+o), 16 Jan. 1993
(25+0), 12 May 1993 (25+o), 17 July 1993 (2 5 +o),— Station
13:26 Sept. 1992(5).— Station 14:23 Jan. 1993 (25+),3 Aug.
1993 (25+0), n Sept. 1993 (25+).— Station 15: 1 June
1991(5).— Station 20: 25 June 1993 (15). —Station 28: 1
June 1991 (3). — Station34: 15 Apr. 1995(4, lo). — Station
35:28 Apr. 1996(3,10).

Known range. Wassaw Sound, Georgia, to Puerto
Rico; northern Gulf of Mexico to Florida Keys (Garcia-
G6mez 1 982, Lemaitre el al. 1 982)

Remarks. Pagurus maclaughlinae is one of the most
common species found in the shallow subtidal waters of
Tampa Bay. Although this species is typically found in
seagrass beds, specimens have also been collected on
hard substrates and high energy beaches. At Station 14,
individuals were found dinging to the gorgonlan
Leptogorgia virgulaia. Pagurus maclaughlinae has been
reported at depths of 1 -5 m (Lemaitre et al. 1 982).

Ovigerous females were collected each month of the
year in Indian River Lagoon, on the Atlantic Coast of
Florida, with peaks (> 50%) occurring in August-October
andFebruary-June(Tunbergetal. 1994). In Tampa Bay,
P. maclaughlinae appears to reproduce throughout the

year since ovigerous females were found during each
season.

Coloration. Antennal flagellum with blue and white
transverse bands. Pereopods with brown and white
transverse bands. Chelipeds light brown with white
tubercles, distal ends of dactyl and fixed finger white. See
Garcia-G6mez(1982)foradditionaldctail.

Pagurus poUicaris Say, 1817

Pagurus poUicaris — Say 1817:1 62. — Hay and Shore
1918:41 l,Plate30, Figure 1 . — Provenzano 1959:401, Figure
16.— Williams 1965: 128, Figure 103.— Felder 1973:27, Plate
3, Figure 8. — Williams 1984:220, Figure 157. — Abeleand
Kim 1986:33, 375h,i.

Material. Station 1: 13 Jan. 1992(1), 1 Sept. 1992(2).—
Stations: 1 1 May 1993(1).— Station 4: 3 July 1992 (3), 4
Jan. 1993 (4).— Station 5: 26 Sept. 1976(1 ),28 Sept. 1976
(l),Sept. 1991(1), 11 May 1993(1).— Station7: lODec.
1982(1).— Station8:4Jan. 1974(1).— Station 9: ISSept.

1 992 (3), 6 Jan. 1 993 ( 1 ), 11 May 1 993 (2) . —Station 12:7
May 1 983 ( 1 ) .—Station 1 3 : 7 May 1 983 ( 1 ) , Apr. 1 99 1 ( 1 ),
26 Sept. 1992(2).— Station 14: 14 Apr. 1970(1), April 1979
(4), Oct. 1979 (1), 3 Aug. 1993 (3), 1 1 Sept. 1993 (3).—
Station 15; Apr. 1979(3).— Station 19: 1 Feb. 1992(3).—
Station 30: Oct. 1 993 (2). — Station 33 : 29 Sept. 1 996 ( 1 ). —
Station 34: 15Apr. 1995(2).— Station 35: 28 Apr. 1996(2).

Known range. Grand Manan, New Brunswick, to
northeastern Florida; Key West, Florida, to Texas
(Provenzano 1959, Williams 1984).

Remarks. Pagurus poUicaris was collected
throughout Tampa Bay , was usually found alone on sand
in the shallow subtidal zone, and was occasionally near
hard substrates. This species is known to inhabit shallow
estuaries, deep harbor channels, and littoral waters
(Williams 1984), although it has been collected to a depth
of 1 1 2 m ( Wenner and Boesch 1 979).

Ovigerous females were collected from early spring to
June in Massachusetts (Nyblade 1970, Carlon and F.bersole
1 995), January and February in North Carolina, and in the
winter in Texas (Fotheringham 1 975). Ovigerous females
were taken from northwestern Florida in February (Cooley
1 978), near Crystal River in December (Lyons et al. 1971),
in Tampa Bay in November and December (Dragovich and
Kelley 1964), and in southwestern Florida in March
(Provenzano 1 959). No ovigerous females were collected
during this study.

Coloration. Eyestalks white with dark brown
surrounding cornea on dorsal part, light yellow near
cornea; cornea light blue-grey with black ring. Antennular
peduncles tan to green; flagella mostly drab green with

44

Hermit Crabs of Tampa Bay, Florida

red and white bands. Antennal peduncles with thin,
reddish, longitudinal stripe; flagella with 2-4 tan or green
articles to every white article. Right chela white to light
brown from merus to area of propodus at insertion of
dactyl; dark brown L-shaped patch beginning at proximal
end of propodus and ending at insertion of dactyl; adjacent
mesial margins of dactyl and propodus darker brown. Left
chela with similar coloring, L-shaped patch less defined.
Second and third pereopods light brown, darker on dorsal
and lateral surfaces. See Provenzano ( 1 959) for additional
coloration notes.

Pagurus stimpsoni (Milne Edwards and Bouvier, 1 893)

Eupagurus stimpsoni — Milne Edwards and Bouvier
1 893 : 1 44, Plate 10, Figures 1 3- 1 8.— Alcock 1905: 1 82.

Pagurus annul ipes — Schmitt 1935:206 (in part), [not
P, annulipes (Stimpson)],

Pagurus bonairensis — Schmitt 1936:376. — Felder
1973:26 (in part), [not Plate 3, Figure 5].

Pagurus bender soni — Wass 1963:144, Figure 5.

Pagurus stimpsoni — Lemaitreetal. 1982:687, Figure

2 .

Material. Station 14: 18 June 1992(2o),23 Jan. 1993
(1).— Station 18: 2 Oct. 1993(3).— Station 30: 2 Oct. 1993
( 1). — Station 32: 28 Oct. 1 996 ( 1 o). — Station 33 : 29 Sept.
1996 (lo).

Known range. North Carolina to Florida; Gulf of
Mexico; Carribean coast of South America (Lemaitre et al.
1982).

Remarks. Only 9 specimens of P. stimpsoni were
collected at the mouth of T ampa Bay or in offshore waters.
Specimens were found on hard substrates with P.
maclaughlinae at Station 14, and P. carolinensis at
Station 30. This species may have an unusually wide
depth range. While most reports are from the shallow
subtidal to depths of 30 m (Lemaitre et al. 1982), Wass
(1963) reported it in the Straits of Florida at depths of
228mand347-512m.

Ovigerous females of P. stimpsoni were collected
during the present study in June, September and October.
Wass (1963) reported a gravid female from the Straits of
Florida in August.

Coloration. Antennal flagellum with brown and white
transverse bands. Pereopods with white and brown
transverse bands.Chelipeds mottled brown and white;
distal ends of dactyl and fixed finger white.

Discussion

Distribution within the Tampa Bay Area

Pagurus maclaughlinae y P. longicarpus and P,
pollicarisvfttt distributed throughout the shallow waters
of Tampa Bay and were often collected together. They
were the only species taken in the upper part of the bay,
including Old Tampa Bay and Hillsborough Bay (for
subdivisions of Tampa Bay see Lewis and Whitman, Jr.

1 985); however, no subtidal hard substrates were examined
in these areas. Savercool and Lewis (1994) documented
several hard-bottom communities in Old Tampa Bay and
collections on these limestone outcroppings and oyster
reefs may reveal additional hermit crab species. Pagurus
maclaughlinae was found in a variety of subtidal habitats,
but was the dominant species collected in seagrass beds.
Pagurus longicarpus and P. pollicaris were most
commonly taken in intertidal or shallow, subtidal waters
on sand and sand/mud substrates. Because no seasonal
quantitative sampling was conducted in subtidal areas, it
was impossible to determine whether these 2 species
underwent seasonal migrations. Along the Texas coast,
both species are subtidal, but migrate to the upper subtidal
zone briefly during the winter, presumably to breed
(Fotheringham 1975).

Clibanarius v Hiatus ^ Pagurus gymnodactylus and
P. stimpsoni inhabited shallow waters of the bay entrance
near hard substates, sand and seagrass beds. Four species,
Paguristes hummi, Paguristes sp., Petrochirus diogenes
and Pagurus impressus were collected from lower bay
waters to offshore of Tampa Bay, mainly on hard substrate
and sand habitats. Paguristes puncticeps, P. sericeus
and Pagurus carolinensis were taken only offshore on
hard substrates in depths of 5-15 m. Although several
species were collected occasionally on high energy
beaches, Isocheles wurdemanni appears to be the only
species restricted to this habitat.

Herm it crab species richness was greatest on the hard
substrate habitats of the bay entrance and shallow offshore
waters where 12 of the 1 4 species found in the study were
taken. The number of species decreased to only 3 in the
lower salinity waters of upper Tampa Bay and less
drastically in the deeper offshore waters.

Zoogeography

Of the 15 species of hermit crabs reported previously
from the shallow waters of the west coast of Florida
(Table 1), 13 were found inthe Tampa Bay area during this
study. Only Iridopagurus caribbensis (Milne Edwards
and Bouvier, 1893), Paguristes tortugae and Pagurus
brevidactylus were not represented in the survey.

45

Strasser and Price

Iridopagurus caribbensis appears to be a rare species
ranging from off South Carolina to the Caribbean Sea in
depths of 10 to 180 m (Williams 1984). There is only one
report of this species from the west coast of Florida (Table
1). Paguristes tortugaehas been found from the Carolinas
through theCaribbean to northern Brazil (Williams 1984).
In the Gulf of Mexico, this species has been documented
only along the coast of southwest Florida (Table 1).
Pagurus breyidactylus ranges from Bermuda and northeast
Florida through the Caribbean to northern South America
(Lemaitre et al. 1 982). Its only documented occurrence in
the Gulf of Mexico is from northwest Florida, but the
distribution of this species may extend to the Texas coast
(McLaughlin 1 975). It is highly probable that the species
diversity of the hermit crab fauna of the Tampa Bay area
is greater than the 14 species reported in this study. Only
additional sampling, especially on the continental shelf,
will help to determine the extent of the faunal richness of
this area.

Tampa Bay is considered by some authors (Hedgpeth
1953, Rehder 1954, Earle 1969, Humm 1969) to be the
boundary between the warm-temperate Carolinean
province and the tropical Antillean province for marine
organisms along the Gulf coast of Florida, The hermit crab
fauna of the Tampa Bay area reflects the transition between
these 2 provinces. Thirty-nine per cent of the species
have widespread distributions including the U.S. east
coast, Gulf of Mexico and Caribbean Sea {Clibanarius
vittatuSy Petrochirus diogenes, Paguristes sericeus,
Pagurus maclaughlinae, P. stimpsoni). Five (39%)
species have a temperate distribution and have been
found along the U.S. east coast and the Gulf of Mexico
{Paguristes hummiy Pagurus carolinensiSy P. impressus,
P. longicarpuSf P. pollicaris). A les.scr tropical influence
is indicated by the presence of only 2 species (15%),
Isocheles wurdemanni and Paguristes punciiceps, with
distributions in the Caribbean and Gulf of Mexico only.
One species, Pagurus gyrnnodactylus, appears to be
endemic to the Gulf of Mexico. Although the Tampa Bay
fauna contains elements from both provinces, as expected,
there is no evidence to support the assertion that this area
serves as a biotic boundary for shallow-water hermit
crabs. McCoy and Bell (1985) came to the same conclusion
about Tampa Bay.

Symbionts

The porcellanid crab Porcellanasayana {Loach 1 820)
was associated with 4 hermit crab species collected in the
Tampa Bay area. This species was found in shells with
Petrochirus diogems (Station 30), Pagurus impressus
(Stations 30, 31), Paguristes punciiceps {StaLionslGyll)
andP. ser/cews (Stations 26, 27). While only one or 2 crabs

were typically found per hermit crab, 3 specimens of
Porceiiana sayami were collected with Petrochirus
diogenes. Porceiiana sayana appears to show little host
specificity and has been reported with Petrochirus
diogenes (Telford and Daxboek 1978, Williams 1984),
Pagurus pollicaris (Williams 1984), Paguristes grayi,
Dardanus yenosusy the queen conch Strombus gigas
(Telford and Daxboek 1978), and the decorator crab
Stenocionops furcata (Hildebrand 1954). fhe large
reported depth range of Porceiiana sayana^ shallow to
92 m (Gore 1974) and 713 m? (Schmitt 1935), has led to
speculation that more than one species may be represented
in these reports (personal communication D. L. Felder).

A male-female pair of bopyrid isopods tentatively
identified as Parathelgcs sp. (personal communication
R.W. Heard, Gulf Coast Research Laboratory, Ocean
Springs, MS 39564) was found attached to the abdomen
of a specimen of Paguristes sp. (Station 26).

Acknowledgments

We are indebted to David K. Camp, formerly at the
Florida Marine Research Institute; Paula Mikkelson,
formerly at Harbor Branch Oceanographic Institution;
and Julio Garcia-G6mez, formerly at the Rosenstiel School
of Marine and Atmospheric Science, for providing
specimens from their collections. Fred Rhoderick, Jesse
Cruz and students from several marine zoology classes
from the University of Tampa helped in the collection of
specimens. Fred Punzo and Stan Rice made helpful
suggestions at various stages of the research and
preparation of the manuscript. We would also like to thank
Rafael Lemaitre, Sara LeCroy, Jerry McLelland, David
Camp, Floyd Sandford and an anonymous reviewer for
their constructive comments on the manuscript.

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Appendix 1. Station data and occurrence of species.

L Southwest side of Courtney Campbell Causeway;
sand /mud; 1 8. 5-26%o salinity; < 1.5 m; triangular dredge.
Species present: Pagurus longicarpus^ P. maclaughlinae,
P. poUicaris.

2. Northwest side of Courtney Campbell Causeway;
sand/mud, Spartina marsh; <1.5 m; dip net. Species
present: Pagurus longicarpus.

3. Southeast side of Courtney Campbell Causeway;
sand/seagrass beds; <1.5m; dip net; Species present:
Pagurus longicarpus^ P. maclaughlinae, P. poUicaris.

4. Northwest side of Gandy Bridge; sand/mud, seagrass
beds; 22%o salinity; <1.5 m; dip net; Species present:
Pagurus longicarpus, P. poUicaris.

5. Picnic Island; sand/seagrass beds; 22-32 °C;22-27%o
salinity; <1.5 m; dip net; Species present: Pagurus
longicarpus, P. maclaughlinae, P. poUicaris.

6. McKay Bay; mud/sand; dip net. Species present;
Pagurus longicarpus.

7. Hooker Point; dip net. Species present: Pagurus
poUicaris.

8. Spoil Island, Hillsborough Bay; dip net. Species
present: Pagurus poUicaris.

9. Ballast Point, sand/seagrass bed; 2 1-33. 5 °C;20-26%o
salinity; <1 m; dip net, hand collection. Species present:
Pagurus longicarpus, P. maclaughlinae, P. poUicaris,

10. Coffeepot Bayou; 1.5 m; hook and line. Species
present: Petrochirus diogenes.

1 1 . Cockroach Bay; mud, oyster reefs, seagrass beds; 20-
29®C; 1 8-30%o salinity; <1.5 m; dip net. Species present:
Pagurus maclaughlinae.

12. Piney Point; sand; <1.5 m. Species present:
longicarpus, P. poUicaris.

13. Bishop Harbor, limestone outcroppings, sponges,
sand; 27-32%o salinity; 3.5 m; hand collection, SCUBA.
Species present; Pagurisfes hummi, Pagur isles sp.,
Pagurus impressus, P. maclaughlinae, P. poUicaris,

14. Northeast Skyway Bridge jetty; sand, concrete
blocks; 28-32%o salinity; <3.5 m; hand collection, SCUBA.
Species present: CUbanarius vittatus, Petrochirus
diogenes, Paguristes hummi, Paguristes sp., Pagurus
gymnodactylusy P. impressus, P. longicarpus, P.
maclaughlinae, P. poUicaris, P. siimpsoni.

15. Blackthorn Memorial Park; seagrass beds; 32%o
salinity; <1.5 m; dip net. Species present: Pagurus
impressus, P. maclaughlinae, P. poUicaris.

49

Strasser and Price

16. BocaCiega Bay. Species present: Paguristeshummi,
Pagurus impressus.

17. Near Shell Key off Pass-a-Grille Beach. Species
present; Pagurus longicarpus.

18. West Tierra Verde south of Pass-a-Grille Channel;
sand, seagrass beds; 0.6 m; hand and tater rake/scooper/
dipnet. Species present: Paguristes hummi, Pagurus
gymnodactylus, P, longicarpus, P. stimpsoni.

19. Fort Desoto Beach; sand; <3 m; hand collection,
snorkeling. Species present; Pagurus impressus, P.
longicarpus, P. pollicaris.

20. Mullet Key Bayou; mud, seagrass beds; <1 .5 m; dip
net. Species present: Clibanarius vittatus, Pagurus
longicarpus, P, maclaughlinae.

21. Mullet Key bayside. Species present; Pagurus
longicarpus.

22. Fort Desoto Pier; sand, algal mats; <0,5 m; hand
collection. Species present: Pagurus longicarpus.

23. Egmont Key, bayside; seagrass beds; 1.2 m; frame
trawl with rollers. Species present: Clibanarius vittatus,
Petrochirus diogenes.

24. 4 miles westof Egmont Key; sand, crushed shell; 6 m;
dredge. Species present: Paguristes hummi.

25. 8 miles west of Egmont Key; sponge, coral, shell;
13.5-15 m; trawl. Species present: Paguristes hummi, P.
puncliceps.

26. Larry’s Ledge; sand, limestone outcroppings, corals,
sponges; 32%o salinity; 15 m; hand collection, SCUBA.
Species present: Petrochirus diogenes, Paguristes
puncticeps, P. sericeus, Paguristes sp., Pagurus
carolinensis.

27. Jack’s Hole; sand, limestone outcroppings, corals,
sponges; 1 5 m; hand collection, SCUBA. Species present:
Petrochirus diogenes, Paguristes hummi, P. puncticeps,
P. sericeus, Paguristes sp. , Pagurus carolinensis.

28. North Anna Maria Island front beach; sand; 3-4 m.
Species present; Jsocheles wurdemanni, Paguristes
hummi, Pagurus impressus, P. maclaughlinae.

29. Molasses Barge off Anna Maria Island; sand, barge
remains; 7 m; hand collection, SCUBA. Species present:
Paguristes sp., Pagurus impressus.

30. St. Petersburg Artificial Reef; concrete, boat remains,
sand; 10 m; hand collection, SCUBA. Species present:
Petrochirus diogenes, Paguristes hummi, P. puncticeps,
Paguristes sp., Pagurus carolinensis, P. impressus, P.
pollicaris, P. stimpsoni.

31. I Mile Artificial Reef off Anna Maria Island; sand,
35%o salinity; concrete pilings; 5-9 m; hand collection,
SCUBA. Species present: Paguristes hummi, Paguristes
sp., Pagurus carolinensis, P. impressus.

32. Egmont Key, front beach; sand; 35%o salinity;
1.5 m; hand collection. Species present: Pagurus
gymnodaciylus, P. stimpsoni.

33. Egmont Key, front beach; concrete, fort remains;
24°C; 34%o salinity; 3 m; hand collection, SCUBA. Species
present: Pagurus pollicaris.

34. Lower Tampa Bay, off Lewis Island; shell; 3-4 m; otter
trawl, Species present: Pagurus maclaughlinae, P.
pollicaris.

35. Lower Tampa Bay, off Point Pinellas, seagrass beds;
2 m; otter trawl. Species present: Pagurus maclaughlinae,
P. pollicaris.

50

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

The Planktonic Copepods of Coastal Saline Ponds of the Cayman Islands with Special
Reference to the Occurrence ofMesocydops ogunnus Onabamiro^ an Apparently
Introduced Afro -Asian Cyclopoid

Eduardo Suarez-Morales

El Colegio de la Frontera Sur, Mexico

Jerry A. McLelland

Gulf Coast Research Laboratory , Jerry.McLelland(^usm.edu

Janet Reid

National Museum of Natural Historyj WashingtoUj D.C.

DOI: 10.18785/grr.ll01.07

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

Part of the Marine Biology Commons

Recommended Citation

Suarez-Morales, E., J. A. McLelland and J. Reid. 1999. The Planktonic Copepods of Coastal Saline Ponds of the Cayman Islands with
Special Reference to the Occurrence oi Mesocy clops ogunnus Onabamiro, an Apparently Introduced Afro-Asian Cyclopoid. Gulf
Research Reports 11 (l): 51-55.

Retrieved from http://aquila.usm.edu/gcr/voll l/issl/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
Research by an authorized editor of The Aquila Digital Community. For more information, please contact Joshua.Cromwell^usm.edu.

Gulf Research Reports Vol. 11, 51-55, 1999

Manuscript received July 24, 1998; accepted November 4, 1998

THE PLANKTONIC COPEPODS OF COASTAL SALINE PONDS OF
THE CAYMAN ISLANDS WITH SPECIAL REFERENCE TO THE
OCCURRENCE OF MESOCYCLOPS OGUNNUS ONABAMIRO, AN
APPARENTLY INTRODUCED AFRO-ASIAN CYCLOPOID

Edua rdo Suarez- Morales', Jerry McLelland^ and Janet Reid'

Cotegio de la Fronlera Sur (ECOS UR). A P 424. Chetumal, Quintana Roo 77000, Mexico
^Gulf Coast Research iMboratory, Institute of Marine Sciences. The University of Southern
Mississippi, 703 East Beach Drive, Ocean Springs, Mississippi 39564, USA
^National Museum of Natural History, Smithsonian Institution, Department of Invertebrate
Zoology, Washington, DC 20560-01 63, USA

ABSTRACT Taxonomic analysis of the copepod specimens collected from 29 Cayman Island ponds revealed
the presence often species including the nearly ubiquitous cyclopoidApocyclops panamensis. Th is species was
widespread throughout the islands, being collected at 27 of the sampling sites. Another common calanoid,
Mastigodiaptomus nesus, occurred at nine sites on Grand Cayman and one on Cayman flrac. A cyclopoid of Afro-
Asian origin, Mesocyclops ogunnus. was collected at two nearly fresh water sites on Grand Cayman and was
considered to be a recent introduction. Because of its known adaptability to fluctuating environmental
conditions, it is likely that M. ogunnus will successfully compete with and probably displace some of the native
species and may become a dominant zooplankter on Grand Cayman.

Introduction Material and Methods

The coastal saline ponds of the Cayman Islands
represent a variety of habitats and. like those of most small
Caribbean islands, are subject to hypersaline conditions
during the dry seasons and flooding during the summer
rainy season. Some ponds are also connected via sinks
and seeps to brackish, anoxic, anchialine cave systems,
and as such arc somewhat affected by tidal flow. Coastal
ponds provide a feeding habitat for a variety of resident
and migratory waterfowl that forage on poeciliid fish and
a variety of small benthic invertebrates including insect
larvae, snails and crustaceans. In conjunction with a
biological assessment conducted in 1996-97 by the Cayman
Island National Trust, plankton samples were collected
from 29 coastal and inland sites on Grand Cayman and the
two sister isles, Little Cayman and Cayman Brae during
August 1996 and January and June 1997. The habitats
sampled included shallow roadside borrow pits and ponds,
tidally influenced mangrove swamps, Typha swamps,
sedge swamps, seasonal pools on grasslands, and the
mouth of an anchialine cave. Salinities at most of the
sampled locations varied from hypersaline in the fall and
winter to nearly fresh in the summer when inundated
during the extensive rainy period. A brief description of
localities where copepods were collected is presented in
Table 1 along with associated data on salinity (%o),
temperature (®C), pH, and dissolved oxygen (D.O., mg/1).
The general location of the collecting sites is shown on
Figure 1.

Fifty non-quantitative plankton samples were taken
using a plankton net with a mesh size of 0.07 mm at 29
coastal and inland pond localities in the Cayman Islands
(Figure 1). All collections were taken from slightly below
the surface of the water (0-0,5 m) by hand-towing the net
a distance of about 1 0-1 5 m. Copepods were examined live
soon after collection, and representative specimens were
sorted from the sample, fixed with 1 0% formalin, and later
preserved in 70% ethanol. Hydrographic data were
collected within the upper 0,25 m at each site using a YSI
multi-parameter system (model 85) and a pH pocket meter.
Geographic coordinates were recorded with a portable
GPS unit. Preserved specimens were examined by the
senior author and identified to species with the aid of
taxonomic descriptions published by Sewell (1940), Van
de Velde (1 984), Bowman ( 1986), Campos-Hemdndez and
Suarez-Morales (1994), and Su^rez-Morales et al. (1996).

Results and Discussion

Taxonomic analysis of the copepod specimens
collected from Cayman Island ponds revealed the presence
of 10 species. These included the nearly ubiquitous
cyclopoid panamensis (Marsh 1913), which
was widespread throughoutthe islands at27 of the sampling
sites, and the common calanoid, Mastigodiaptomus nesus
Bowman, 1 986, which occurred at 9 sites on Grand Cayman
and one on Cayman Brae. More isolated were the

51

Suarez-Morales et al.

TABLE 1

Cayman Island Pond station data and copepod occurrence records. GC = Grand Cayman, LC = Little Cayman, CB = Cayman
Brae, NT = Not Taken, Key to species: k? = Apocyclops panamensisy AC -Acartia ionsoy MA = Macrocy clops albidusy
yW = Mastigodiapiomus nesusy ML = Mesocy clops longisetusy MO = Mesocyclops ogunnuSy M3 = Metis jousseaumeiy
XT = Thermocyclops tenuis yT^ = Tropocyclops exiensuSyTl^ = Tropocyclops prasinus cf, aztequei.

Temp.

Salinity

D.O.

Copepod

Site

Habitat

Date

%o

mg/1

pH

species

Betty Bay Pond, GC

Slightly brackish, borrow pit,

1/16/97

29.8

2.6

MM

MN, MO

19MT50"N/8ril'30"W

mangrovc/woodland fringe, Chara mats

6/1 1/97

34.4

5.8

■■

AP

Collier’s Pond, GC

Permanent, shallow brackish, mangrove

1/16/97

25.8

2.7

5.4

9.8

AP,MN

19“20'03"N/8P0510"'W

fringe, Ruppia beds

6/11/97

29.9

2.6

1.1

8.9

AP

Governor’s Pond, GC

Small inland Typhof Urochloa mutica

1/27/97

3.0

9.4

AP, MN, TP

19‘’16’39"N/8n8'30"W

fringe, seasonal, temporary

6/12/97

■■

6.9

8.6

MN

Least Grebe Pond, GC

Small inland XypWsedge fringe,

8/28/96

34.6

0.8

8.9

MN,MO

19°16'4R”N/8ri8’17"W

seasonal, temporary

1/27/97

24.3

0.2

1.53

9.4

AP, MN, TP

6/12/97

30,4

1.0

1.05

8.4

AP.MN

Malporta.s Pond, GC

Shallow, brackish, mangrove fringe

1/16/97

26.7

■■

9.6

AP, MN

19°20’35”N/8J°12’17”W

6/11/97

33.2

10.7

AP

Meagre Bay, GC

Shallow, brackish, mangrove fringe

26.1

10.6

10.5

AP,MN

19°lT38"N/8n3'44”W

28,8

15.9

4.1

10.5

AP

Palmetto Pond, GC

Shallow, brackish-hypersaline, mixed

1/17/97

26.9

14.5

4,4

9.5

AP,MN

19'’23'i6'’N/81^2l'58'’W

mangrove Fringe

6/13/97

27.9

19.7

5.1

9.4

AP

Pease Bay, GC

Shallow, brackish, mangrove fringe.

1/16/97

1.6

10.5

10.1

AP,MN

19"17l5"N/8ri4'26"W

rock outcroppings, Ruppia beds

6/12/97

■■

19.5

1.9

10.1

AP

Point Pond, GC

Shallow, brackish, temporary, mixed

1/26/97

32.0

5.8

12.7

11.2

AP,MN

19"20'58''N/8n3’2r’W

woodland fringe, Ruppia beds

Sea Pond, GC
19°23'14*W81°22’32"W

Tidally influenced mangrove swamp

1/15/97

29.4

25.9

8.4

9.1

AT

Vulgunncr’s Pond, GC

Shallow, hypersaline lagoon, small tidal

1/14/97

33.9

22.9

12.1

9.5

AP, TE

19'’23’I0"N/81"22’59"W

creek inlet, Ruppia beds

6/10/97

30.9

26.8

7.4

9.8

AP, TT, MJ

Bittern Pond, LC

Marshland, Meagre tern {Acrostichum)

6/3/97

28.9

2.1

6.5

9.1

AP

19“39'36“N/80“05'46"'W

fringe, Ruppia beds

Booby Pond, LC

Seasonal, brackish-hypersaline. mixed

1/18/97

19.0

24.3

5.0

9.8

AP

19‘’39’58"N/80‘’04'15’’W

woodland/mangrove fringe, rock
outcroppings, sinkholes and underground
seep influence

6/3/97

27.0

3.3

4.4

8.2

AP

Bulldozer Pond, LC

Marshland, seasonal, shallow, ironshore

1/20/97

23.0

21.9

4.0

9.3

AP

19"39'38"N/80°06'02"W

rock pools

6/4/97

5.0

3.8

9.9

AP

Coot Pond, LC

Temporary, seasonal,meadow pond.

6/5/97

31.0

0.1

0.1

7.9

ML,TT

19M1’53"N/79“58’18"W

sedge fringe

Easterly Pond Complex, 1 C
19'’4r56"N/75°59'14"W

Shallow, brackish, Ruppia beds

1/18/97

23.9

11.1

8.4

10.5

AP

Grape Tree Pond, LC

Shallow, brackish, mangtove/sea grape tree

1/18/97

■n

■1

■9

AP

19°4r51'‘N/80‘’03’10"W

(Coccoloba) fringe

6/5/97

■B

AP

Jackson’s Pond, LC

Permanent, mangrove/mixed woodland

1/19/97

22.1

10.8

13.0

9.9

AP

19°4l'26"N/B0°03'54”W

fringe

52

COPEPODS OF THE CaYMAN ISLANDS

TABLE 1 (Continued)

Tanp.

Salinity

DO.

Copepod

Site

Habitat

Date

“C

%o

mg/1

pH

species

Lighthouse Pond, LC

Seasonal hypersaline, connected to

1/19/97

23.7

31.8

9.6

10.2

AP

19"39'34''M/80"06'32"W

underground cave system

6/4/97

27.7

1.1

2.9

9.2

AP

McCoy’s Pond, LC

Shallow, brackish, mangrove fringe

1/19/97

Hi

AP

19M0'26"N/80 '’05*49“ W

6/4/97

WM

AP

Salt Rock Cave, LC

Mouth of anchialine cavesystem

6/6/97

NT

NT

NT

AP, MA

Sandy Point Pond, LC

Shallow, brackish, eutrophic

1/18/97

21.4

14.2

10.2

AP

19M2’05“N/79'*57'53“W

6/5/97

8.7

8.4

9.8

AP

Tarpon Lake, LC

Seasonal, brackish-hypersaline, old-

1/18/97

23.4

8.5

4.2

9.8

AP

19»40>4J'*n/80'’02'27”W

growth mangrove swamp

6/3/97

25.9

5.2

4.9

8.3

AP

Westerly Pond -east site, CB

Narrow brackish inlet from main pond.

1/21/97

23.0

11.1

11.2

10.5

AP

19'’4ri2"N/79‘’52‘49’'W

mangrove fringe

6/8/97

27.8

2.8

1.3

8.7

AP

Westerly PoikI -west site, CB

Shallow hypersaline, mangrove fringe

1/21/97

33.8

4.6

10.0

AP,MN

19'’41’03'’N/79'’53'18"W

6/8/97

mm

3.4

4.4

9.3

AP,ML

Mangrove Wreck Pond, CB

Brackish, dredged canal adjacent to old

1/21/97

23.4

16.3

7.1

10.3

AP

19'’4ri4'’N/79'’52'10’'W

growth mangrove swamp

6/7/97

28.4

2.8

6.2

9.2

AP

Red Shrimp Hole, CB

Marshland, ironshore rock pools, mangrove

6/8/97

27.3

0.6

1.9

8.5

AP

19°4138"N/79^50*52''W

fringe, sinkhole connection to cave system

Salt Pond, CB

Shallow, brackish-hypersaline, man-made

1/21/97

25.5

28.2

8.8

10.5

AP

l9°4l’16'’N/79“5r49"W

levee on one edge

6/8/97

27.2

8.1

5.5

9.8

AP,TT

The Split-s, CB
19'’4r39’'N/79'’52’13"W

Interior brackish, karstic bluff formation

1/22/97

23.2

7.2

2.5

9.5

AP

occurrences of the predominantly freshwater cyclopoids,
Macrocyclops albidus (Jurine, 1820), Mesocyclops
longisetus (Thiebaud, 1914), Thermocyclops tenuis
(Marsh, 1909), Tropocyclops extensus (Kiefer, 1931),
Tropocyclops prasinus cf. aztequei Lindberg, 1955, and
Mesocylops ogunnus Onabamiro, 1 957. Two species with
greater tolerance for higher salinities, the harpacticoid
Metis jousseaumei (Richard, 1 892) and the calanoid /4car/w
toma Dana, 1852, were limited to single occurrences at
Vulgunner^s Pond and Sea Pond, sites on Grand Cayman
with direct marine influence.

Most of these species have been previously recorded
from Grand Cayman (Reid 1990), and the overall
biogeographic affinities of the local copepod community
are clearly tropical. The most noteworthy record is that of
Mesocyclops ogunnus, an apparently introduced Afro-
Asian species, found at Least Grebe, Grand Cayman, and
Betty Bay Pond, Grand Cayman, 2 nearly freshwater sites.
It can be distinguished from the known American species
of Mesocyclops by the presence of a row of spines on the
maxillular palp, a character shared only with the African M.
salinus Onabamiro, 1957, Other diagnostic characters of
M. ogunnus include: pediger 5 with several lateral and a

few dorsal spines, seminal receptacle with broad lateral
arms and a long curved pore-canal, caudal ramus with
naked medial surface and with spines at the bases of the
lateral and lateral most terminal caudal setae (Van de Velde
1984, Reid andPinto-Coelho 1994).

Mesocyclops ogunnus is distributed in Nigeria,
Subsaharan Africa, the Near East, South and Southeast
Asia, and Brazil This species inhabits a wide variety of
freshwater environments, and is one of the mosteurytopic
species of Mesocyclops in the Afro-Asian region (Van de
Velde 1984, Jeje and Fernando 1992, Reid and Pinto-
Coelho 1994). This adaptive capacity would explain the
success of this species when introduced into a new
environment. In the Cayman Island system, M. ogunnus
is not widely distributed, nor present in a variety of
environments. This suggests that the invasion of M.
ogunnus in the Cayman Islands is quite recent, since, like
many other introduced copepods, M. ogunnus is a very
efficient competitor and can exploit different types of
environments (Reid and Pinto-Coelho 1994). Were this
species long established in the Caymans, we would expect
it to be common and abundant, A more thorough
investigation into similar sites throughout the year would

53

Suarez-Morales et al.

Figure 1. Cayman Islands, British West Indies showing the location of coastal saline ponds where copepods were collected.
Inset shows relative location in the Caribbean Sea and distances between the 3 islands.

54

COPEPODS OF THE CaYMAN ISLANDS

likely better define the extent of the M ogunnus invasion
into the Cayman Islands. The adaptability of A/, ogunnus
to differing environmental conditions leads us to anticipate
that it will successfully compete with and probably displace
some of the native species and may become a dominant
zooplankier in the area.

It is probable that A/, ogunnus has been transported
along with aquaculture organisms to other parts of the
world, since it has been recorded from aquaculture ponds
in the Ivory Coast. Aquacultural activities have apparently
effected the introduction of several species of copepods.
For example, the Asiatic calanoid, BoeckellairiarticulaiOj
was apparently introduced to Italy together with Chinese
carp. Pseudodiapiomus marinust another Asiatic calanoid,
was possibly introduced in a similar manner into the
United States. Pseudodiapiomus trihamatus of the Indo-
Pacific may have been introduced to Brazil with the shrimp
Penaeus monodon. Finally, Mesocyclops rultnet-i, an
East-Asian cyclopoid was perhaps introduced to the
Southern U.S. by rice culture (reviewed by Reid and Pinto-
Coelhol994).

The other copepods found in the Cayman Island
ponds we sampled (species of Tropocyclops and
Apocyclops panamensis) have different ecological niches
and may not be competitors of M ogunnus. Apocyclops
panamensis, the most abundant species in the Cayman
Island ponds sampled, was introduced to the Ivory Coast
from Western Atlantic coasts (Dumont and Maas 1988).
The only calanoid found in the Cayman Island ponds is
Mastigodiaptomus nesus\ however, the specimens
recorded during this survey lack the characteristic dorsal
keel described by Bowman (1986) for this species.

Thermocyclops tenuis had previously been recorded
only from Grand Cayman (Reid 1 990), and the new records
from Little Cayman and Cayman Brae represent a modest
range extension for this cyclopoid. Specimens from this
area have been deposited at the National Museum of
Natural History, Smithsonian Institution (USNM-268059).

Acknowledgments

We are grateful to the Cayman Island National Trust
who funded the project and to the Cayman Department of
the Environment who cooperated in logistics on Grand
Cayman. Logistic and field assistance was provided by
Fred Burton and Patricia Bradley. Richard Heard, Chet
Rakocinski, Sara LeCroy, Wayne Price, and Mike Abney
provided field assistance and comments on early drafts of
this manuscript.

Literature Cited

Bowman, T.E. 1986. Freshwater calanoid copepods from the
West Indies. Syllogeus 58:237-246.

Campos-Hernindez, A. and E. Sufirez-Morales. 1994.
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I. BiologiA y Sistem^lica. CIQRO, Chetumal, Mexico.

Dumont, H.J. and S. Maas. 1988. Copepods of the lagune Ebri6
(CdtedTvoirc). Revue d’HydrobiologieTropicale 21 iS-?.

Jeje, C.Y and C.H. Fernando. 1992. Zooplankton associations
in the Middle Niger-Sokoto Basin (Nigeria: West Africa).
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Hydrographic 77:237-253.

Ketelaars, H.A.M. andL. W. Van Breemen. 1 993. The invasion
of the predatory cladoceran Bythotrephes longimanus
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Biesbosch reservoirs. Verhandlungen der Internationale
Vereinigung fiir Theoretische und Angewandte Limnologic
25:1168-1175.

Reid, J.W. 1990. Continental and coastal free- livingCopepoda
(Crustacea) of Mexico, Central America and the Caribbean
region. In: D. Navarro and J.G. Robinson, cds., Diversidad
Bioldgica en la Reserva de la Biosfera de Sian Ka’an,
CIQRO/University of Florida, Quintana Roo, Mexico,
p. 175-213.

Reid, J.W. and R.M, Pinto-Coelho. 1994. An Afro-Asian
continental Mesocyclops ogunnus, found in Brazil;

with a new key to the species of Mesocyclops in South
America and a review of intercontinental introductions of
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Sewell, R.B.S. 1940. Copepoda, Marpacticoida. The John
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Museum (Natural History) 8:1-382.

SuSrez-Morales, E., J.W. Reid, T.M. Iliffe and F. Fiers. 1996.
Caialogo de los Cop6podos (Crustacea) Continentalcs de
la Peninsula de Yucatan, Mexico. CONABlOand ECOSUR,
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Velde, I, Van de. 1984. Revision of the African species of the
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55

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

Variations in the Ventral Ciliature of the Crustacean Symbiont Hyalophysa (Ciliophora^
Apostomatida) from Mobile Bay and Dauphin Island^ Alabama

Stephen C. Landers

Troy State University

Michael A. Zimlich

Troy State University

Tom Coate

Troy State University

DOI: 10.18785/grr.ll01.08

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

Part of the Marine Biology Commons

Recommended Citation

LanderS; S. C., M. A. Zimlich and T. Coate. 1999. Variations in the Ventral Ciliature of the Crustacean Symhiont Hyalophysa
(Ciliophora; Apostomatida) from Mobile Bay and Dauphin Island; Alabama. Gulf Research Reports 11 ( 1 ) ; 57-63.
Retrieved from http:// aquila.usm.edu/gcr/voll l/issl/8

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 editor ofThe Aquila Digital Community. For more information, please contact Joshua.Cromwell^usm.edu.

Gulf Research Reports Vol. 11, 57-63, 1999

Manuscript received July 24, 1998; accepted November 17, 1998

VARIATIONS IN THE VENTRAL CILIATURE OF THE CRUSTACEAN
SYMBIONT HYALOPHYSA (CILIOPHORA, APOSTOMATIDA) FROM
MOBILE BAY AND DAUPHIN ISLAND, ALABAMA

Stephen C. Landers, Michael A. Zimlich and Tom Coate

Departmeni of Biological Sciences, Troy State University, Troy, Alabama 36082, USA

ABSTRACT Apostome ciliates arc symbiotic organisms whose 1 ife cycles are complex and involve specific feeding,
divisional, migratory, and phoretic stages. In this study we examined apostome irophonts (the diagnostic stage)
from a variety of crustacean hosts in the Mobile Bay and Dauphin Island, Alabama, area. The hosts were grass
shrimp {Palaemoneles pugio and P. pahtdosus\ striped hermit crab (Clibanarius vittatus), blue crab (Callinectes
sapidus), and pink shrimp {Farfanlepenaeus (=Penaeus) duorarum). A number of similar but distinct morphoty pes
of apostomes were present, those corresponding to descriptions of species of well as variant forms.
The morpho types observed in this study had the following characteristics; variations in the formation of the anterior
ventral field ofkinctosomes from falciform field 9, variations in the degree to which ciliary row I (kinety 1) was
separated into 2 segments; and variations in the development of kinety a . A record of the variant morphotypes
that do not correspond exactly to an established species should prove useful to biologists attempting to identify
apostomes from crustacean molts. We choose not to name the variant forms as new species because they exist as
different morphotypes within a population of cells, because some of these types occur in low frequency, and
because one of the variant forms changes from one morphotype to another.

Introduction

Bradbury (1966) established the genus Hyalophysa
in 1966 for the organism H. chattoni, a common
apostomatous ciliatc associated with crustaceans in North
America, This symbiont spends most of its life cycle
encysted on a host such as a shrimp or crab, waiting for
a chemical signal to indicate that the host will soon molt.
After receiving the signal, the ciliate metamorphoses from
a quiescent phoretic cell to a trophont (macrostome) that
will excyst upon eedysis of the crustacean (Figure 1 ). The
trophont then swims to the inside of the exoskeleton and
feeds by pinocytosis on the exuvium contained within.
Following this single opportunity to feed, the ciliate
settles on a substrate, encysts, and produces daughter
tomites. The tomites (microstomes) are migratory cells
with a non-functional mouth thatseltle on a crab or shrimp
to encyst and begin the cycle again.

Exuviotrophic apostome ciliates are ubiquitous
organisms, reported from a wide variety of crustaceans in
North America including members of the genera
Pagurus, Clibanarius, Palaemoneles, Cambarus, Uca,
Vpogebia, CallinecteSi Sesarma, Penaeus, Alpheus,
Lophopanopeus. Cancer, Panopeus, and Carcinides
(Bradbury 1966, Bradbury and Clamp 1973, Grimes 1976,
Johnson 1 978). Only one report exists in the recent literature
that surveys apostomes from a number of hosts from the
same locale (Grimes 1976). The present study was
undertaken to belter understand the apostomes of the
Dauphin Island and Mobile Bay region in Alabama by
sampling the apostome trophonts feeding in the molts of

a variety of crustaceans. The hosts examined in this study
were Palaemoneles pugio, P. paludosus, Clibanarius
vittatus, Callinectes sapidus, and Farfanlepenaeus
{=Penaeus) duorarum. Penaeid shrimp names are based
on Perez Fartante and Kensley ( 1 997).

We report many different apostome morphotypes
including H. chationi (Bradbury 1966), a number of
variants similar to FI. chattoni, as well as variant forms
that do not exactly match published species descriptions.
These morphotypes illustrate the variation that occurs in
the ciliature within apostome species from one host to
another, and provide insights to the transformation from
the phoront to the trophont.

Materials and Methods

Grass shrimp (P. pugio), blue crabs (C. sapidus), and
striped hermit crabs (C. vittatus) were collected with a dip
net or by hand in the airport road marsh, Dauphin Island,
Alabama (30“1 5'N, 88“07'W). Pink shrimp (F. duorarum)
were collected by throw net from the eastern end of
Dauphin Island (30“ 15.03* N, 88“ 04.60' W), and the grass
shrimp P. paludosus was collected by dip net at Meaher
State Park in Baldwin County, Alabama (30“39'N, S7“55' W)
between the mouths of the Apalachee and Blakeley rivers.
The animals were kept at the main campus of Troy State
University in filtered water obtained at the collection site
and were fed flaked or pelleted fish food every other day.
Their water was changed approximately once a week.

57

Landers et al.

Figure 1. The life cycle of the apostomatous ciliate
Hyalopkysa. Clockwise from the top: trophonts within the
exoskeleton, tomonts undergoing division while encysted
on the substrate, the swimming infestive tomite, the
encysted phoront. Line drawings of the cells are based on
silver nitrate impregnation. Adapted from Landers et al.
1996.

Grass shrimp were housed in large groups and only
isolated in glass bowls prior to molting. The prcmolt
shrimp were identified by the presence of the developing
setae visible under the old exoskeleton in the uropods
(Freeman and Bartcll 1 975). Crabs and prawns were kept
in isolation at all times due to the difficulty in identifying
premolt organisms. Following eedysis, the apostomes
swimming in the exoskeleton were pipetted directly out of
the molt for fixation and silver impregnation.

The ci Hates were fixed in2.5-5%glutaraldehydefor5- 15
minutes. After a thorough washing in distilled water, the cells

were enrobed in warmed gelatin and impregnated with silver
nitrate followingamodificationoftheChatton-Lwoffmethod
(Bradbury and Clamp 1973). Followingsilver impregnation
the cover slips were immersed in cold 70% ethanol,
dehydrated, cleared in xylene, and mounted with resin.

Results

A variety of different apostome morphotypes were
observed (Figures 2-10) which had the following 3
characteristics: variations in the dissolution of falciform
field 9 (FF9) to form an anterior ventral field of kinetosomes
(AVF); variations in the degree to which ciliary row 1
(kinety 1 or Kl) was separated into 2 segments; and
variations in the development of kinety a (K^ from FF9.
During this study we did not observe variations in the
dorsal or the posterior ventral ciliature of the trophont
stage, but only differences involving the above named
characteristics. Though a gradation of morphotypes exists,
the cells that are most representative of the data are
illustrated in Figures 2-10. The numbers of each cell type
are referenced by the host crustacean in Table 1 .

Apostomes from Clibanarius viiiatus

Few trophonts (5) were identified from the striped
hermit crab, though all exhibited the typeci liature originally
described for//, chattoni (Figures 2 and 8). This ciliature
has been described previously (Bradbury 1966). A brief
description of the cell follows: The cell is oval to reniform
and measures approximately 55 x 30 mm (the size is variable
depending upon the amount of ingested food). Nine
kinetics spiral dextrally around the cell from the anterior
to the posterior end. Kinety 1 extends posteriorly along
the anterior third of the cell, then bends sharply to the
right and continues around the cell. Kinety 2 is divided,

TABLE 1

Listing of all apostome ciiiates and their hosts (#observed/^ examined). The ciliates are referenced by Figure number
from this articleand by host ’^Data from morphotype #4 and #5 combined.

Figure #

Host

2

3

4

5

6

7

Clibanarius vittatus

5/5

Callinectes sapidus

1/15

11/15

3/15

Farfantepenaeus {-Penaeus) duorarum

1/27

nil

\im

2/27

Palaemonetes pugio

18/95

3/95

65/95*

65/95*

1/95

8/95

Palaemonetes paludosus

5/17

12/17

58

ApOSTOME ClLlATES OF CRUSTACEA

Figures 2-7. The ventral ciliature of trophonts of Hyalophysa. Line drawings based on silver nitrate impregnation. Solid
lines indicate ciliary rows (kinetics). Individual dots represent kinetosomes. K = kinety, CVP = contractile vacuole pore,
FF = falciform Field, AVF = anterior ventral field, Ka = kinety a, xyz= kineties x and z, T = kinetosomal tail. Figure
2. Hyalophysa chattoni type morphology. Figure 3. //. chattoni variant with a poorly developed AVF. FF9 has divided into
two rows but has not broken into an AVF. Figure 4. H. chattoni \ nr x^nX with an altered Kl and AVF. Note the kinetosomal
tail, derived from FF9, at the lower right corner of the AVF. Figure 5. //. chattoni variant with an altered Kl and AVF.
Note the kinetosomal tail, derived from FF9, at the lower right corner of the AVF. Figure 6. H. chattoni variant. Note the

separation of Kl to form a Kla and Klb and the absence of
Note the large AVF, kinetosomal tail, and altered Kl.

forming a K2a and K2b. Kinety 2a runs along the left of K3.
Kinety 4 has a crook at the anterior end and extends
around the cell to the posterior. Kinety 5 is divided into
K5a, a short Z-shaped fragment, and K5b, which bends
around the celland terminates on the mid-ventral surface.
Kinety 6 and K7 spiral from the anterior pole to the
posterior pole. The posterior portion of K8 is similarto K6
and K7, but anteriorly it is a double row of kinetosomes
termed the Falciform Field (FF8). Kinety 9 parallels K8 on
the right. Anteriorly K9 is broken into a field of scattered
kinetosomes termed the Anterior Ventral Field (AVF).
Three short kineties (x, y, and ^ are located to the left of
the contractile vacuole pore between K9 and K I . Kinety
a is a short kinety located anterior to xyz .

Apostomes from Farfantepenaeus{-Peiiaeus)duorarum
Hyalophysa spp. trophonts from F. duorarum molts
were variable in many respects. In 7 of 27 cells the FF9 did not

kinetosomal tail on the AVF. Figure 7. H. chattoni variant.

break apart to form an AVF but instead formed one to 3
doubled rows of kinetosomes that occupied the area between
FF8 and K 1 a (Figure 3). Additionally, K I was divided into a
K 1 a and K 1 b, with K 1 a completely separated from its lower
segment and aligned along the left side of K2a. Kinety awas
not observed in these trophonts. This morphology is an
intermediate form between Hyalophysa and Gym nodi nioides
(Bradbury I966,ChattonandLwoff 1935).

The majority (17 of 27) of the trophonts from F.
duorarum were similar to the H. chattoni variant illustrated
in Figure 4. In this type, FF9 divided into scattered groups
of2 to4 kinetosomes to form an AVF and possessed a tail
of doubled kinetosomesin the lower right corner, derived
from the remnant of FF9. Kinety a was observed in this
type. Kinety 1 was either divided into a separate K 1 a and
Klb, separated by a few scattered kinetosomes, or Kla
was connected to Kl b but appeared to be stretched away
from its lower fragment. In addition to this cell type, 2 of

59

Landers et al.

Figures 8-10. Photomicrographs of selected silver-stained apostomes. Figure 8. Hyalophysa chattoni specimen from
Palaemonetes pugio. The cell is approximately 81 |im wide. Figure 9. H. chattoni variant from P. pugio. Note the
kinetosomal tail (arrowhead). The cell is approximately 59 ^m wide. Figure 10. H. chationi variant from Callinectes
sapidus. Note the break in K1 (arrowhead). The cell is approximately 75 i^m wide.

27 cells possessed no tail (Figure 6) and one cell was a
type specimen (Figure 2).

Apostomes from Callinectes sapidus

Most of the trophonts (11 of 1 5) observed from the
blue crab had a morphology similar to the trophont that
was most common on F. duorarum (Figure 4). K 1 was
either stretched to the point of separation or was divided
into a K I a and K 1 b and separated by a short gap occupied
by 3 to 4 kinetosomes. An AVF was fully formed, with a
tail of kinetosomes present in the lower right corner that
varied from short (4 kinetosomes) to much more defined
(8 kinetosomes). Kinety a was present in these cells, either
attached to the tail of kinetosomes or separate from it. In
addition to this cell type, 3 cells from C. sapidus had no
tail (Figure 6) and one was similar to the type morphology
of H. chattoni (Figure 2).

Apostomes from Palaemonetes pugio

A large number of cells from P. pugio were examined
with the majority of the cells (65 of 95) similar to the
morphologies illustrated in figures 4 and 5. In these cells
a tail of kinetosomes is found at the posterior right comer
of the AVF, varying in size from 6 kinetosomes ( Figure 4)
to 36 (Figure 5). The average number of kinetosomes in
thetailwas 14 (N = 33). Ka had usually not yet separated
from the kinetosomal tail of the AVF. The 30 remaining
cells represented a variety of morphologies. Eighteen of
the cells were the type morphology (Figure 2), 3 cells had
a FF9 that was divided into 2 or 3 fragments rather than an

AVF (Figure 3), and one cell had a type AVF but a broken
K1 (Figure 6). Finally, 8 cells possessed a large AVF in
which individual kinetosomes were spread out into a large
shield-shaped field (Figure 7), A tail of kinetosomes was
present and Kla was shortened, connected to Klb by
scattered kinetosomes. The AVF of this apostome is
similar to that of//. frager/(Grimes 1976).

Apostomes from Palaemonetes paludosus

Trophonts from the molts of P. paludosus were similar
to one of 2 morphologies. Five of 17 cells had a short
kinetosomal tail and a bend or break in Kl, as seen in
apostomes from C. sapidus^ P. pugio ^ or F. duorarum
(Figure 4). The remaining cells ( 1 2 of 1 7) had no kinetosomal
tail and a separated or bent Kl (Figure 6). Of the last group
of cells, 2 had a Kla that did not curve towards Klb but
instead was aligned close to K2a. Those 2 cells were most
similar to the freshwater apostome H. bradburyae (Landers
etal.1996).

Discussion

In this study we have demonstrated a number of
apo.stome variants. Particular variants are not restricted to
specific species of hosts, but rather, are found in mixed
populations on a number of crustaceans. All of the
variations result from subtle differences that occur in the
cell during the transformation of the phoront stage to the
trophont (Figures 1 1-1 3). Of all of the changes that take
place during this transformation, the formation of the AVF

60

ApOSTOMB ClLlATES OF CrUSTACHA

Figures 11-13. Line drawings illustrating the metamorphosis of the phoront to the trophont during the premolt period on
the host (adapted from Landers 1986). Note the formation of the AVF from FF9. K = kinety, FF = falciform field,
AVF = anterior ventral field, Ka = kinety a, xyz = kinetics i and z, EC = developing extended cytostome.

from FF9 and the bend in K1 are the most variable. The 4
nominal species of Hyalophysa are differentiated by
characteristics of the AVF and K1 , among other features
(Bradbury i 966, Bradbury and Clamp 1 973, Grimes 1 976,
Landers et al. 1 996). We report variations in the trophont
ciliature that involve 3 key characteristics, the AVF, Ka,
and K1 .

The dissolution of FF9 is a process that occurs
normally during the phoretic stage of Hyalophysa to form
the AVF (Bradbury and Trager 1967). Landers (1986)
described this metamorphosis using protargol silver
impregnation (see Figures 11-13) and suggested that Ka
is a derivative from the posterior fragment of FF9. This
hypothesis is confirmed by the present data. Variant
fonns in which a tail of kinetosomes exists clearly show Ka
connected to the posterior tip of the AVF tail.

Kinety I is a variable structure among the
species. In the //, chattoni type morphology, not often
seen in this study, K 1 has a sharp 90‘’ bend to the right as
it extends posteriorly along the right border of the cytostome.
This bend is also found in H. trageri. In H. Iwoffi and H.
bradburyae K1 is divided, though the position of the
anterior segment differs. In the present study K1 was most
often stretched into either 2 kinetics that were barely
connected or they were separated by a gap occupied by
scattered kinetosomes. Conversely, a wide separation was
observed between K 1 a and K 1 b in some apostomes fromP.
paludosus, a characteristic more similar to the freshwater
form H. bradburyae than to H. chattoni. A wide separation
between K 1 a and K 1 b was also present on apostomes with
an undeveloped AVF (Figure 3).

The morphotypes described in this report were chosen
as representatives to reflect the many variations we
observed. One morphotype matches that of a described
species (Figure 2) whereas other forms have characteristics

that do not correspond to established species. For example,
the cell illiustrated in Figure 3 is intermediate between
Gymnodinioides and Hyalophysa. We think this form
should currently be considered a variant of H. chattoni,
and not a species of Gymnodinioides because the later
genus possesses an unbroken Kl, and FF9, if present, is
unbroken (Chatton and Lwoff 193 5, Bradbury etal. 1996).
The cells illustrated in Figures 4 and 5 are similar to H.
chattoni though in these forms the posterior tip of FF9 has
not completed its transformation and remains as a tail of
kinetosomes on the ventral surface. The cell in Figure 6 is
similar to H. chattoni ifKl a points posteriorly towards
K 1 b, as illustrated. However, if K I a is more closely aligned
next to K2a, the cell is similar to//, bradburyae, a fre.sh water
form (note: this form on P paludosus is not surprising,
because the shrimp were caught near the Apalachee and
Blakeley rivers where a freshwater apostome might be
expected). The cell in Figure 7 is similar to the H. chattoni
variants in Figures 4 and 5 as well as to //. trageri (a
species known only from the genera Sesarma and Uca). It
is similar to H. trageri because of the large shield shaped
AVF, but differs from that species in having a kinetosomal
tail on the AVF and havinga separated Kl . At this time we
are reluctant to assign the variants illustrated in
Figures 2-7 to new taxa because they exist as different
morphological types within the same population of cells
and because of the low frequency of some of the variant
types. Additionally, we have observed that the cells
illustrated in Figures 4 and 5 transform into the H. chattoni
type morphology after feeding has ended (Zimlich,
manuscript in preparation), suggesting that some of the
variants represent a lag in the development of the H.
chattoni trophont.

It should be pointed out that some of these variant
types are not restricted to Alabama, though they, and not

61

Landers et al.

The eslablished taxa, represent the dominant types from
the Mobile Bay area. Neptun (1988) reported the variant
itiustrated in Figure 5 from P. pugio in North Carolina,
though it was rarely seen there. Also, the variant described
in Figure 3 from F. duorarum was found (rarely) in molls
of P. pugio in North Carolina (S. Neptun, personal
communication).

Although different species of apostome trophonts
are morphoiogically di.stinct, other stages in the life cycle
such as the tomont and tomite are remarkably similar to
one another (Chatt on and Lwo('Tl935). In the trophontthe
cilia are apparently not involved in feeding and can vary
in position without affecting the cell. Our data support
this hypothesis, since cells of all morphologies bloated
nonnaliy as they fed within the host’s exoskeleton.

Many hypotheses and future experiments can be
designed to address the nuestion of why these variants
exist and whether the variation in the ventral ciliature has
a functional or developmenia! significance. As the ventral
ciliature does not appear to affect the feeding process it
is possible that this variation has evolved within the
species because there are few selective pressures to
restrict the patterning of this ciliature. All of the .species
of P/yaiophysa revert to a common morphology as they
encyst and produce daughter tomites, suggesting that
developmental restraints exist during lomitogenesis that
do not allow for as much morphological variation in later
stages. There are many factors that could play a role in
determining the subtle morphological differences of the
trophonf s ventral ciliature, such as diet, host animal,
water temperature, season, and pollution effects. It is also
possible that the morphotypes exist as a result of genetic
variations within the population that are not immediately
influenced by environmental factors. Future avenues of
research are plemifulm this area. For example, apostomes
from one host could be used to infect other cmstaceams
to see if the proportion of the variant types changes with
the host, .Also, a clonal population of cells could be
produced from one trophont and carried through many
molt cycles on cleaned shrimp to see if morphological
variations are present. Many other experimental variables
could be tested in the laboratory to furthcranalysepossible
cause.s of variations in the trophont,

in theirhisloricraonograph, Chatton and Lwoff (1935)
separated the apostomes into a number of distinct groups
based on their diet and lifecycles. Thfs .study has focused
on only one group, the exuviotvopbs, w hose diet consists
of exuvial fluid from crustacean exaskejetons. Earlier
reports (Chatton and Lwoff 1 935, Bradbury 1966, Grime.s
1 976, Lindley 1 978) leave little doubt that exuviotrophic
apostomes exist on probably all crustaceans ranging from

decapods to amphipods to barnacles. While previous
reports acknowledge exuviotrophic apostomes, probably
of the genus ffyalophysa, from the shrimp,
Farfaniepenaeus aztecus, F. duorarum, F. brasiliemis,
Lilopenaeus {-Penaeus) setiferm, and 1. vannamei,
(Johnson 1978, Lotz and Overstreet 1990), our study
confirms the presence of Hyalophysa chattoni variants
on the pink shrimp, F. duoraritm, and extends the known
record of the genus Hyalophysa to a variety of Crustacea
from the Mobile Bay region. This record establishes the
variability present in the apostome population of this
region.

Additionally, we have observed apostome trophonts
within molts of ibcmole crab Fonerda spp. from Dauphin
Island but were not able to obtain satisfactory silver
stains. Future studies of apostomes will attempt to
determine the exuviotroph fauna of Crustacea from the
high energy beach zones.

ACKNOW! cEDGMEN rs

The authors would like to thank Di\ B J. Bateman for
h elp digitizing I in e drawings on th e compu te r . Th i s pro] ec t
was supported in pan by a grant from the honor society,
Beta Bela Beta, aw'arded to M. Zim licit.

Literaturk Cjtf.d

Bradbury, P.C. 1966, The life cycle and morphology of the
npostoinatous ciliate Hyalophysa ohattoni n.g,. ri, sp.
Journal orPrutozoology l3*.20U-225.

Bradbury, P.C. and PC. Clamp. J973, Hyalophysa IwoffU sp, n.
fjoiti the ficshwal&f shrimp PalcKmonetes paludosus and
rcvisionofthc genus Hyalophysa. Journal olTrotozoology
70:210-213.

Bradbury, P.C. ai^d W/l'ragcr, 1967. The meiamorpho.sis from
ihc pluMOut to the trophont in tiyalophy.^a. Journal of
Protozoology H:307-312.

Bradbury, P.C., L.-M. Zhang and X.-B, Shi. 1996. A
vedescriplion of Gymnodinioides caridinae (Miyashita
1933) from Pahemonetes smsnsis (Soilaud 191 1) in the
SonghuaRiver. Journal of Eukaryotic Microbiology 43 :404-
40S.

Chatton, T. and A. Lwoff. 1935. Lc.s CiliiJs Apostomes. 1.
Aper^u hi.storiquc et general; elude monographique des
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Freeman. J, and C- Bartell. 1975. Characterization of the molt
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(Decapoda, Caridea). General and Comparative
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Grime.s, B.H. 1976. Notes on the distribution of Hyalophysa
and Gyrnnodinioidus on. crustacean hosts in coa.stal North
Carolina and a description o( Hyalophysa trageri sp. n.
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62

Apostome Ciliates of Crustacea

Johnson, S.K. 1 978. Handbook of shrimp diseases. Texas A&M
Sea Grant Publication. TAMU-SG-75-603.

Landers, S.C. 1986. Studies of the phoront of Hyalophysa
chattoni (Ciliophora, Apostomatida) encysted on grass
shrimp. Journal of Protozoology 33:546-552.

Landers, S.C., A. Confusione and D. Defee. 1996. Hyalophysa
bradburyae, a new species of apostome ciliate from the
grass shnmp Palaemonetes kadiakensis. European Journal
of Protistology 32:372-379.

Lindley, J. A. 1 978. Continuous plankton records: the occurrence
of apostome ciliates (Protozoa) on Euphausiacea in the
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63

Gulf Research Reports

Volume 1 1 I Issue 1

January 1999

Gordon Pennington Gunter^ 1909-1998

W David Burke

Gulf Coast Research Laboratory

DOI: 10.18785/grr.ll01.09

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

Part of the Marine Biology Commons

Recommended Citation

Burkc; W. 1999. Gordon Pennington Gunter, 1909-1998. Gulf Research Reports 11 (l): 65-67.
Retrieved from http;//aquila.usm.edu/gcr/voll l/issl/9

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GORDON PENNINGTON GUNTER
1909-1998

W. David Burke

Gulf Coast Research Laboratory, Institute of Marine Sciences, The University of Southern
Mississippi, 703 East Beach Drive, Ocean Springs, Mississippi 39564, USA

... if you are interested in marine science or any other science, you run along as fast as you can go. Other things
are just an interference, they just take up your time. (Gordon Pennington Gunter)

Gordon Pennington Gunter was bom in the Red River
country of north Louisiana, Natchitoches Parish, in the
townofGoldonna, on August 1 8, 1909, or “about 44 years
after the death throes of the
Confederacy”, as Gunter described his
birth year. Gunter also recorded that his
father, John Osbon Gunter, had been bom
in Creston, Louisiana, in 1 876, or “about
the year the last of the Yankee soldiers
left.” Gordon Gunter’s grandfather, Miles
Osbon Gunter, served as a cavalryman
under Fighting Joe Wheeler. Gunter
recalled his mother saying his great-
grandmother died during the War
because, “She was old and tired and sick
and did not have enough to eat.” Dr.

Gunter attributed her death to the result of
Sherman’s March to the Sea. Gunter
remained somewhat cool toward the
memory of William Tecumseh Sherman
and usually spoke pejoratively about the General. Gunter
described seeing an old Confederate Veteran hurrying
along on New Orleans’ Royal Street in 1931. The old
veteran was dressed in the old butternut uniform of the
Confederacy, and Gunter hurried along to overtake him,
just to touch him. Gunter could not overtake the old man
in the crowd and that was the last Confederate soldier he
was ever to see. It was inevitable that with these
sensitivities Dr. Gunter should find himself involved with
The Sons of Confederate Veterans and The Order of the
Stars and Bars, organizations devoted to the preservation
of respect and honor for those men who had served as
Confederate Soldiers. Dr. Gunter served that organization
long and faithfully and rose to become Commandant of the
organization at the state level. In keeping with his
ecumenical view, it should be pointed out that Dr. Gunter
was also a member of The Sons of the American Revolution.

Gordon Gunter had gone off to Louisiana State Normal
College with the idea that he might become an attorney,
like his father, or perhaps become a French scholar. He
abandoned both those ambitions immediately after being

exposed to his first biology course, which interestingly
enough was mandatory, rather than elective. That course
seemed to have been a turning point in Gunter’s life as he
proceeded to earn a B.A. in zoology,
securing that degree in 1929. With that
degree in hand he went to the University
of Texas with the intention of becoming a
bacteriologist and earned the M.A. degree
in 1 93 1 . Upon completion of the master’s
degree, Gunter worked on shrimp and
oysters in Louisiana, Florida and Texas,
and on fishes in California, during the
Debris Dam Fisheries Survey for the U. S.
Engineers Office. Dr. Gunter was always
nattily dressed and he did not go about
during business hours without a jacket
and necktie. Years earlier Gunter had been
admonished by his mentor, Professor
Williamson of Louisiana State Normal
College, for going about the campus
improperly dressed, that is to say sans necktie. He seemed
never to have forgotten the instruction in dress and at
some level it might have embarrassed him. It could be
pointed out that the omission of the necktie could have
been due to youthful exuberance and just sheer excitement
associated with being at school, because Gunter also
recalled that his father had bought him a fine red gelding
to go back and forth to school on, and in the excitement
at his first day of matriculation, young Gunter clanked
about in the college halls throughout most of the first day,
oblivious to the fact that he was still wearing his roweled
riding spurs.

In 1932 Gunter married his firstwife,Carlotla “Lottie”
Gertrude La Cour. They produced a daughter, Charlotte
Anne Gunter Wood Evans of Galveston, Texas, and two
sons. Miles Gordon Gunter and Forrest Patrick Gunter of
Austin, Texas. Dr. Gunter took measureless pride in these
children. For many years the single bit of decoration in
Gunter’s office was a big photograph of his son, Gordon,
in his Marine dress whites. The younger Gordon Gunter
barely survived injuries sustained in a fiery helicopter

65

Burke

crash in the Philippines, en route to Marine duties in
Vietnam. He is today a successful attorney in Austin,
Texas.

Gunter had returned to the University of Texas in 1 939
as an instructor in physiology and had a concurrent
appointment as a marine biologist to the Texas Game, Fish
and Oyster Comm ission. During this time he was lured into
the study of physiology and zoology by Professor Elmer
Julius Lund, and Gunter completed his doctoral work in
those disciplines in 1 945 . After a great deal of work by Dr.
Lund, the University of Texas founded the Institute of
Marine Science at Port Aransas in 1945. Gunter, after
receiving his Ph.D., conducted research there, becoming
acting director of the Institute from 1949 to 1954, then
director until he left in 1955 to come to Mississippi. Lund
had also established Publications of the Institute of
Marine Science in 1945 and Gunter served as editor of that
journal from 1950 to 1955.

In 1955, Dr. Gunter accepted the appointment as
Director of the then eight-year-old Gulf Coast Research
Laboratory in Ocean Springs, Mississippi. That same year
he married the former Miss Frances Hudgins of Kosciusko,
Mississippi. They produced two sons, Edmund Osbon
Gunter, bom in 1960, and Harry Allen Gunter, bom in 1964.
Dr. Gunter doted on these sons and almost always referred
to them as his ‘Tittle boys’", I suppose in contradistinction
to his older children who would have been pretty well
grown up at the time. In his memoirs. Dr. Gunter has
referred to his older children as his “brood of littleTexans”.
Dr. Gunter was indulgent of his ‘Tittle boys*” vitality and
encouraged them in some practices that I suppose must
have been unsettling to Mrs. Gunter, who usually went
along with the program cheerfully enough. One activity
that seemed to amuse Dr. Gunter very much involved
asking red-haired Harry, the younger boy, to “Climb the
walls, Harry; show our visitor how you do it!” At which
point Harry would dash across the room, propel himself
against the wall and take two or three steps up the vertical
wall. This effort would take him along pretty well toward
the ceiling, at which point he would somersault and land
on the floor with a resounding thump, sometimes on his
feet, sometimes not.

Mrs. Frances Gunter is now retired after a
distinguished career as an elementary school teacher;
Harry is a medical investigator and lives in Purvis,
Mississippi, with his family. Edmund has for several years
now worked with technical aspects of production with
educational television in Mississippi and seems to have
retained some of his father’s interest in things natural,

Gordon Gunter, during the course of his directorship
at the Gulf Coast Research Laboratory, took the place from

a part-time summer school teaching facility to a full-time
year-round research facility, and much of the significant
early research in the northern Gulf of Mexico took place
here under his direct supervision. Dr, Gunter started out
with one full-time scientist and two part-time support
personnel. At the time of his retirement, GCRL programs
were conducted by about 100 senior marine scientists,
technical staff, and support personnel. Dr. Gunter was a
50-year member of the American Fisheries Society, a
charter member and president of the World Mariculturc
Society, later named the World Aquaculture Society, and
a member and president of the Mississippi Academy of
Sciences. His lifetime body of work is represented by over
330 scientific papers and articles, both scholarly and
popular. His earlier works regarding the relationships of
salinity and temperature of the northern Gulf to marine life
have been required university readings to an entire
generation of marine biology students (see Selected
Bibliography). He was singlehandedly responsible for
establishing and developing GCRL’s library, which may
well be the premier marine library on the Gulf Coast and
today bears his name. In the early 1960s, Dr. Gunter
developed the concept of Gulf Research Reports as a
mechanism . . devoted primarily to publication of the
data of the Marine Sciences, chiefly of the Gulf of Mexico
and adjacent waters.”

As early as 1968, Dr. Gunter was working with a
handpicked staff of physiologists to formulate an artificial
diet for raising shrimp. Even though no particularly high
level of technology existed for culturing shrimp at that
time, it is apparent that Gunter understood the inevitability
of such development, which was, of course a burgeoning
industry by the mid-1980s. Gunter always believed that
one ofthe major needs inthenorth central Gulfof Mexico
was a large, long-term effort to discover the full effects of
the Mississippi River on the biology of the fisheries
resources in the area. “We have learned much but there
are still too many things unknown about the River’s
influence,” he said. “This work alone is enough to keep a
multi-disciplinary team of workers busy for 20-25 years,
and that would be quite an accomplishment.” Gunter
frequently conjectured as to what the “real natural history”
of the Mi.ssissippi River would be if the Army Corps of
Engineers would stop tinkering with it. Most competent
hydrologists concur that without control efforts, the
natural tendency would be for the Atchafalaya to
“capture” the flow of the Mississippi River. In other
words the Mississippi River, instead of flowing pastNew
Orleans, would turn westward and enterthe Gulf of Mexico
near Morgan City, Louisiana. On one occasion he spent
many days at his desk, clucking and scribbling and calling

66

Gordon Pennington Gunter

and harassing various libraries for historical river flow
data of the Mississippi River proper as contrasted to
flows down the Atchafalaya River. He concluded that the
tendency was for the Atchafalaya to grow and the
Mississippi to diminish in such a manner that by the year
2038 these two rivers would be of equivalent size.

Gunter’s career as a marine biologist and leader in
marine research and education spanned more than 60
years. After stepping down as Director of GCRL, he
continued his association with the Laboratory as professor
of zoology and director emeritus until his retirement from
active service to the State of Mississippi in 1 979 at the age
of 70. “He was one of thepioneers,” retired GCRL Director
Thomas D. Mcllwain, said. Mcllwain, now a National
Marine Fisheries Service administrator, was a leader in
nominating Gunter’s name for a National Oceanic and
Atmospheric Administration (NOAA) research vessel in
recognition of the marine scientist’s fisheries work in the
Gulf of Mexico. The NOAA ship was moved to
the Gulf of Mexico and commissioned as the Gordon
Gwrt/e/* on August 28, 1998, with Dr. Gunter in attendance
at the ceremonies.

About 1 977, 1 was invited to accompany Dr. Gunter on
a trip to Texas and we found ourselves in Goldonna,
Louisiana, where he wanted to show me his boyhood
home. We spent part of that afternoon wandering about
in the old Goldonna Cemetery, where Dr. Gunter would
point out where his parents were buried and the markers
of cousins, uncles and other kin. On December 1 9, 1 998,
Gordon Pennington Gunter joined them, and I will miss
him. No more will 1 have a traveling companion whose
standard traveling accouterment consisted of a handgun,
an Authorized King James Version of the Bible, and a quart
of bourbon.

Acknowledgments

I gratefully acknowledge the use of Gunter Archives
No. 1 , 2, 6, 7, 1 0, and 1 1 , located at Gunter Library, Gulf
Coast Research Laboratory, Ocean Springs, Mississippi,
and the article, “ Serendipity and science: The life of
Gordon Gunter,” by James Tighe, found in Coast January-
February 1996.

Selected Publications

1 938. Notes on invasion of fresh water by fishes of the Gulf of
Mexico, with special reference to the Mississippi-
Atchafalaya river system. Copeia 1938(2):69“72.

1 938. Seasonal variations in abundance of certain estuarine and
marine fishes in Louisiana, with particular reference to life
histories. Ecological Monographs 8:3 1 3-346.

1941. Relative numbers of shallow water fishes of the northern
Gulf of Mexico, with some records of rare fishes from the
Texa.s coast. The American Midland Naturali.st 26:194-
200 .

1945. Studies of marine fishes of Texas. Publications of the
Institute of Marine Science, University of Texas 1 : 1-190.

1950. Seasonal population changes and distributions as related
to salinity, of certain invertebrates of the Texas coast,
including the commercial shrimp. Publications of the
Institute of Marine Science, University of Texas I;7-51.

1950. Correlation between temperature of water and size of
marine fishes on the Atlantic and Gulf coasts of the United
States. Copefa l950(4);298-304.

1952. Historical changes in the Mississippi River and the
adjacent marine environment. Publications of the Institute
of Marine Science, University of Texas 2:1 19-139.

1957. Predominance of the young among fishes found in fresh
water. Copeia 1957(I):I3-l6.

1957, Salinity. Chapter7. In: Treatise on Marine Ecology and
Paleoccology. Vol. 1 Ecology. Memoir 67, Geological
Society of America, p. 1 29- 157. (A.S. Pearse and Gunter).

1957. Temperature. Chapter 8. In: Treatise on Marine Ecology
and Paleoccology. Vol. 1 Ecology, Memoir 67, Geological
Society of America, p. 1 59- 1 84.

1961. Some relations of estuarine organisms to salinity.
Limnology and Oceanography 6: 1 82- 1 90.

1961. Salinity and size in marine fishes. Copeia I961(2):234-
235.

1963. Biological invesligationsoflheSt. Lucie Estuary (Florida)
in connection with Lake Okeechobee discharges through
the St. Lucie Canal. Gulf Research Reports, 1:189-307.
(Gunter and G.E. Hall).

1964. Some relations of salinity to population distributions of
motile estuarine organisms, with special reference to penaeid
shrimp. Ecology 45:181-185, (with J.Y. Christmas and R.
Ki Hebrew).

1965. A biological investigation oftheCaloosahatchee Estuary
of Florida. Gulf Research Reports 2:1-71.

1 967. Some relationships of estuaries to the fisheries of the Gulf
of Mexico. Part IX Fisheries, In: G.H. Lauff, ed„ Estuaries,
Publication No. 83. American Association for the
Advancement of Science, Washington, DC. p. 621-638.

1974. A review of salinity problems of organisms in United
States coastal areas subject to the effects of engineering
works. Gulf Research Reports 4:380-475. (Gunter, B.S.
Ballard and A. Vekataramiah).

67