THE OCCURRENCE OF SEA WASPS (CUBOMEDUSAE)

IN MISSISSIPPI SOUND AND THE NORTHERN

GULF OF MEXICO

PHILIP J. PHILLIPSl AND W. DAVID BURKE

Gulf Coast Research Laboratory, Ocean Springs, Mississippi

ABSTRACT
Two species of Cubomedusae, Chiropsalmus quadrumanus and Tamoya
haplonema, are reported from the northern Gulf of Mexico in Mississippi
coastal waters. C. quadrumanus was commonly taken in Mississippi
Sound in August and September. Two specimens of Tamoya haplonema
taken at a depth of 50 fathoms constitute the first record of this species jn
the Gulf of Mexico. Conditions of high salinity and temperature in Mis-
sissippi Sound probably allowed for the development of populations of these
bottom-dwelling medusae in the estuary. Specimens of Chiropsalmus with
intact gonads were found to be sexually mature, indicating a breeding pop-
ulation in northern Gulf waters. The morphology of Chiropsalmus and
Tamoya is also considered.

INTRODUCTION
During August, September, ~f 1968, sea wasps, Chirop-
and December
salmus quadrumanus L. Agassiz (Family Chiropodidae) were encountered
in Mississippi Sound. Two specimens of the cubomedusan Tamoya haplo-
nema F. Muller (Family Carybdeidae) were taken in a shrimp trawl off the
Mississippi coast at a depth of 50 fathoms. One additional specimen of T.
haplonema in the Gulf Coast Research Laboratory museum was collected by
Buena Ballard on 31 October 1959 in a shrimp trawl in the vicinity of Gulf
Shores, Alabama. Sea wasps (Cubomedusae) are commonly found in warm
and tropical waters. Guest (1959) stated that C. quadrumanus constituted
a major component of the invertebrate fauna of Matagorda Bay, Texas,
during the summer and fall months. Mr. J. Y. Christmas of the Gulf Coast
Research Laboratory informed us that C. quadrumanus has been taken
during the summer months in Mississippi Sound in previous years; however,
no notes were made of its occurrence. Owing to the lack of pigmentation,
transparency of the bell, and damage in collection, this medusa is probably
often overlooked in trawl samples.
During August and September several cases of coelenterate stinging,
severe enough to warrant medical attention, were reported to us. Whether
these stings were caused by sea nettles, Chrysaora quinquecirrha (DeSor),
or cubomedusae is uncertain; however, sea wasps, being amongst the most

1 Present address: Department of Biology, Texas A&M University, College Station, Texas 77843.
854 Bulletin of Marine Science [20(4 )

noxious of cnidaria, do constitute a potential hazard. In these waters, col-

lecting Chiropsalmus and handling live specimens required rubber gloves
and considerable care.
DISTRIBUTION

Guest (1959) discussed the distribution of C. quadrumanus. On the

Atlantic coast it is found as far north as Beaufort, North Carolina, and has
been reported as far south as the coast of Brazil. Kramp (1961) listed this
form as occurring also off the coasts of Sumatra, Australia, and West Africa,
as well as in the Indian Ocean. This is the first record of this species for
Mississippi Sound. In the course of our sampling program, Chiropsalmus
was taken only in the estuary and not in open Gulf waters. Local fishermen,
however, have brought us specimens of Chiropsalmus taken in the northern
gulf both east and west of the mouth of the Mississippi River.
As summarized by Kramp (1961), T. haplonema has been reported pre-
viously in the western hemisphere as far north as New England and as far
south as Brazil, as well as in the West Indies and Bahamas. Kramp also
listed this form as occurring off the coasts of the Congo, French Guinea,
Sengambia, and Ludenitz Bay, S. W. Africa. This is the first reported occur-
rence of Tamoya in the Gulf of Mexico. Hedgpeth (1954) listed the sea
wasp Carybdea aurifera Mayer (Family Carybdeidae) as occurring in the
Gulf of Mexico. C. aurifera is, however, a rare form and is listed by Kramp
(1961) as a "young medusa, probably = C. sivickisi Stiasny 1926." Juve-
nile or young medusae of Carybdea and Tamoya are easily confused, and it
is quite possible that the specimen of Carybdea reported by Hedgpeth is a
young specimen of Tamoya.
MORPHOLOGY

Morphologically there was relatively little difference among the various

size-classes of C. quadrumanus. Subumbrellar sacs were relatively larger in
larger specimens; their length ranged up to one-half the height of the bell
cavity. In the smallest specimens examined (28.0- to 48.0-mm bell diame-
ter) , subumbrellar sacs were relatively undeveloped, ranging from one-
eighth to one-quarter the height of the bell cavity. The ratio of bell diameter
to bell height is approximately I : 1 and is expressed in the equation Y =
-10.1139 + 1.1552 X; the correlation coefficient is 0.90. The relationship
of bell diameter to bell height is shown graphically in Figure 1. The mean
bell diameter and mean bell height for all specimens measured are 50.1 mm
and 52.1 mm, respectively. There are no significant differences in size
between sea wasps taken on different dates.
The oral arms are one-half to two-thirds as long as the diameter of the
bell. The normal number of pedalial branches is seven; however, pedalia
with six or eight branches are common. Different pedalia on the same
1970] Phillips & Burke: Sea Wasps in Mississippi Sound 855

100

90

80

70

60

'?
EO
-'
.......
L 50

.•
a
E
40

-;; 30
<!l

20

10

o 10 30 40 50 60 70 80 90 100

Bell Height (•••on)

FIGURE 1. The relationship of bell height to bell diameter in Chiropsalmus

quadrumanus.

medusa occasionally have varying numbers of branches. Tentacles are

milky white and are about one-half to two-thirds as long as the height of the
bell in preserved specimens.
Specimens of T. haplonema are one-third higher than wide. As can be
seen from Table 2, pedalia are about 2.75 times as long as broad, and the
height of the bell is approximately three times the pedalial length. The four
perradial sense organs are located about one-seventh the distance from the
margin to the apex of the bell. Tentacles have a distinct pinkish hue.
Tamoya and Chiropsalmus are easily distinguished, as Tamoya has un-
branched pedalia.
856 Bulletin of Marine Science [20(4)

TABLE 1
DATA FOR SPECIMENS OF Chiropsalmus quadrumanus COLLECTED
IN MISSISSIPPI SOUND IN 1968
---
Water Mean and range Mean and range of
Depth Salinity temp. of bell height bell diameter
Date (ft) Number (%0) ( °C) (mm) (mm)
---
21 Aug. 27 4 27.8 30.1 39.5 34.0
(38.0-41.0) (30.0-41.0)
28 Aug. 18 3 27.8 25.0 37.3 36.7
(28.0-53.0) (27.0-51.0)
30 Aug. 13 16+ 28.5 26.6 56.1 52.6
(36.0-74.0) (34.0-64.0)
27 Sept. 13 4 27.7 26.8 not measured not measured

5 Dec. Just below 3 34.0 15.0 58.7 70.0

low-water (38.0-80.0) (45-100)
mark on north
beach of bar-
rier island

ECOLOGY
C. quadrumanus is taken predominantly in bottom trawls. On one occa-
sion in December, three specimens were found stranded in very shallow
water on the north beach of Horn Island. Data for the collection of C.
quadrumanus are presented in Table 1. When picked up in trawls, these
cubomedusae were often taken along with large numbers of sea nettles,
Chrysaora quinquecirrha (DeSor). As was reported by Guest (1959),
these medusae were taken on soft muddy bottoms. Large volumes of ecto-
procts and hydroids were also taken. Analyses of the stomach contents of
C. quadrumanus revealed large numbers of benthic microcrustacea (Lucifer
sp.) as well as larvae of Squilla sp., crab zoeae, and amphipods. We believe
that these sea wasps were grazing on the hydroids and ectoprocts and cap-
turing the associated microcrustaceans.
As seen in Table 1, depths at which individuals of Chiropsalmus were
taken ranged from 13 to 27 ft, and salinity from 27.8%0 to 34.0%0. During
the summer months, salinities in Mississippi Sound were relatively high,
approaching that of open Gulf waters. Guest (1959) reported a similar
condition for Matagorda Bay. During July and August, oceanic sipho-
nophores, including Nanomia bijuga (Chiaje), Agalma okeni (Eschscholtz),
Muggiaea kochi (Will), and Diphyes dispar (Chamisso & Eysenhardt), were
taken in the estuary. The high salinity regime of Mississippi Sound during
1970] Phillips & Burke: Sea Wasps in Mississippi Sound 857
TABLE 2
MORPHOMETRICS OF SPECIMENS OF Tamoya haplonema FROM THE
GULF OF MEXICO

Pedalia Height of sensory

Bell Bell organs above
height diameter Width Length Ratio, length bell margin Source of
(mm) (mm) (mm) (mm) to width (mm) specimens
120 90 16 44 2.64 15 Mississippi
15 46 3.07 coast
16 44 2.64
16 44 2.64
130 80 9 43 4.78 18 Mississippi
13 34 2.62 coast
36 13 2.77
missing missing
93 75 15 34 2.27 13 Alabama
15 33 2.20 coast
14 33 2.36
15 32 2.13
Sum 30.12
Mean 2.76
Variance 0.5350
Standard deviation 0.73

these months undoubtedly allowed for the incursion of these forms. The
advent of cold weather in October was probably responsible for the sudden
disappearance of Chiropsalmus from the estuary. The reappearance of
viable individuals of Chiropsalmus in December on the north beach of Horn
Island near Dog Keys Pass was probably due to south-southwest winds that
then switched to a northerly direction. North winds also will commonly
strand Stomolophus meleagris Agassiz on the north beaches of the barrier
islands. The occurrence of C. quadrumanus in the estuary is due to the
fortuitous combination of high salinity, high water temperatures, and suit-
able wind conditions. This species may be much more common in the more
southerly portions of the Gulf of Mexico, as witnessed by the fact that
individuals were much more numerous in Matagorda Bay (Guest, 1969).
Intensive sampling of the plankton in Mississippi Sound and adjacent
offshore waters has revealed no planktonic cubomedusoids. The early stages
of these forms remain a mystery. We attempted to maintain individuals of
Chiropsalmus in aquaria; however, they did not remain viable for more than
3 days. All of the individuals of Chiropsalmus with intact gonads were
examined and found to be sexually mature; the smallest of these had a
27-mm bell diameter. This species is dioecious; about equal numbers of
858 Bulletin of Marine Science [20( 4)

males and females were present. The sperm packets and ova look re-
markably like those of S. meleagris. Ova were predominantly unfertilized,
although a small percentage (less than 1 per cent) of ova were in early
cleavage. The three specimens taken in December were sexually mature,
indicating that these medusae may spawn year round.
Tamoya haplonema is a rare form in northern Gulf waters and little can
be said regarding its ecology. As is the case with Chiropsalmus, Tamoya is
a bottom dweller. Two specimens taken 9 October 1968 off the Mississippi
coast at 50 fathoms (29°19'N, 88°14'W) were in poor condition, having
been damaged by the trawl. One specimen taken in 1959 off the Alabama
coast was a mature female having mostly unfertilized ova along with a very
small number of fertilized ova in early cleavage.
Although cubomedusae are not a "normal" component of the estuarine
fauna, high salinities along with high water temperatures will allow for the
temporary establishment of large populations in Mississippi Sound. Follow-
ing a newspaper article concerning sea wasps, numerous specimens of
Chiropsalmus were brought to us by local shrimp fishermen. Although in
our sampling program we capture Chiropsalmus on relatively few occasions,
it is probably much more common in these waters during the warmer
months of the year than is indicated by our data. Guest (1959) reported
this cubomedusa to be as abundant as Chrysaora and Stomolophus in Mata-
gorda Bay; however, such was not the case in Mississippi Sound.
The occurrence of sexually mature specimens of C. quadrumanus in
winter as well as summer indicates that this form may breed on a year-
round basis and that the medusoid stage may have a relatively long life-span,
at least longer than that of the semaeostome Chrysaora, which, on the
Mississippi coast, does not survive past October.

ACKNOWLEDGMENTS
We would like to thank Mr. Edward Donovan for his help in collecting
C. quadrumanus. We also wish to express appreciation to Mr. Charles K.
Eleuterius for his help in preparing the statistics in this paper. This study
was conducted in cooperation with the U.S. Department of the Interior,
Bureau of Commercial Fisheries, under Grant No. PL 89-720 (Project
JF2-1) .
ADDED NOTE
It has come to the attention of the senior author that, during the spring
and summer of 1970, Chiropsalmus quadrumanus was responsible for
numerous cases of stinging on the Galveston, Texas, bathing beaches.
Numerous specimens of C. quadrumanus were collected by Mr. Howard
Armstrong and Mr. Bill Wardle on the front beach and in Galveston Bay
during August.
1970] Phillips & Burke: Sea Wasps in Mississippi Sound 859
SUMARIO

INSlDENCIA DE CUBOMEDUSAE EN MISSISSIPPI SOUND Y LA

PARTE NORTE DEL GOLFO DE MEXICO

Se reportan dos especies de Cubomedusae, Chiropsalmus quadrumanus y

Tamoya haplonema. de la parte norte del Golfo de Mexico en las aguas
costeras de Mississippi. C. quadrumanus se cogi6 comunmente en Missis-
sippi Sound en agosto y septiembre. Dos ejemplares de Tamoya haplonema
constituyen el primer reporte de esta especie en el Golfo de Mexico. Con-
diciones de alta salinidad y temperatura en Mississippi Sound probable mente
hacen posible el desarrollo en el estuario de poblaciones de estas medusas
que habitan en el fondo. Se encontr6 que ejemplares de Chiropsalmus con
g6nadas intactas estaban sexualmente maduros. indicando la existencia de
una poblaci6n en estado de reproducci6n en las aguas del norte del Golfo.
Se consider6 tambien la morfologia de Chiropsalmus y Tamaya.

LITERATURE CITED
GUEST, W. C.
1959. The occurrence of the jellyfish Chiropsalmus quadrumanus in Mata-
gorda Bay, Texas. Bull. Mar. Sci. Gulf & Carib., 9( 1): 79-83.
HEDGPETH, J. W.
1954. Scyphozoa. Pp. 277-278, in The Gulf of Mexico, its origin, waters,
and marine life. Fishery Bull. Fish Wildl. Serv., U. S., 55 (89), xiv +
604 pp.
KRAMP, P. L.
1961. Synopsis of the medusae of the world. J. mar. bioI. Ass. U. K., 40:
1-469.