Life History of the Cannonball Jellyfish, Stomolophus meleagris L.

Agassiz, 1860
(Scyphozoa, Rhizostomida)
Authors(s): Dale R. Calder
Source: Biological Bulletin, Vol. 162, No. 2 (Apr., 1982), pp. 149-162
Published by: Marine Biological Laboratory
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Reference: Biol. Bull. 162: 149-162. (April, 1982)

LIFE HISTORY OF THE CANNONBALL JELLYFISH, STOMOLOPHUS

MELEAGRIS L. AGASSIZ, 1860 (SCYPHOZOA, RHIZOSTOMIDA)

DALE R. CALDER

Department of Invertebrate Zoology, Royal Ontario Museum, 100 Queen's Park, Toronto, Ontario,

Canada M5S 2C6

ABSTRACT

Stages in the life history of the scyphozoan Stomolophus meleagris from the

planula to the newly liberated ephyra were raised in the laboratory and are described

for the first time. After swimming actively for 2-5 days, the ciliated planula larvae

settled and scyphistoma morphogenesis occurred. Fully developed scyphistomae

were cone-shaped and bore a whorl of about 16 tentacles around a dome- or knob-

shaped proboscis. Podocyst formation was the only observed method of asexual

reproduction in cultures of scyphistomae maintained for one month. Strobilation

began as soon as nine days after scyphistoma morphogenesis and occurred in scy-

phistomae with as few as eight tentacles. The strobilation process, completed in

about 3.5 days at 25?C, was not accompanied by any noteworthy color changes.

Most strobilae produced two ephyrae each, although the number varied from one

to three. Some scyphistomae began to strobilate a second time within a week after

completion of an initial round of strobilation. Newly liberated ephyrae possessed

a normal complement of eight lappet pairs and eight rhopalia. They were mor-

phologically similar to, yet distinguishable from, ephyrae of the related species

Rhopilema verrilli. None of the examined stages of S. meleagris contained algal

symbionts.

INTRODUCTION

Biologists did not recognize the relationship between polyp and medusa stages

in the Cnidaria until life history studies were undertaken on scyphozoans by Sars

(1829, 1835, 1841), Dalyell (1836), and Siebold (1839). Polypoid and medusoid

forms, previously regarded as separate taxa, were both found to occur in the life

cycles of Aurelia aurita and Cyanea capillata. Sars, in the course of his studies,

also demonstrated that Scyphistoma, Strobila, and Ephyra were stages in the life

cycle of scyphozoans rather than being separate genera.

In the years since these early studies, complete life histories have been described

for few of the 200 known species of Scyphozoa. In the Hydrozoa, such investigations

have become frequent to resolve problems of classification and synonymy; separate

classification systems for polyps and medusae have been used in the class, and

polypoid and medusoid stages of a species have often been known by different

names. Comparable problems in scyphozoan systematics have been relatively minor

because the conspicuous and relatively divergent species of scyphomedusae have

been named and classified while their small, obscure, and morphologically indistinct

polyps have not. The one notable exception is the genus Stephanoscyphus Allman,

1874, whose polyps are protected by a well-developed sheath of perisarc. Originally

thought to be a hydrozoan, Stephanoscyphus gives rise to scyphomedusae of the

order Coronatae.

Received 9 December 1981; accepted 18 January 1982.

149

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 DALE R. CALDER
150

With the dearth of life history investigations in the Scyphozoa, stages other

than the medusa are often unknown even for some of the more common species.

Strobilation, a process in which the medusoid form is derived from the polypoid

form, likewise remains undescribed for most scyphozoans. So it is with Stomolophus

meleagris L. Agassiz, 1860, one of the most abundant species of scyphomedusae

along the southeastern and Gulf coasts of the United States (Mayer, 1910; Kraeuter

and Setzler, 1975; Burke, 1976; Calder and Hester, 1978). Various phases in the

development of the medusa only have been described in this species (Mayer, 1910;

Stiasny, 1922). The purpose of this paper is to describe the life history of S.

meleagris from planula to newly liberated ephyra stages, including observations

on the strobilation process.

MATERIALS AND METHODS

Stages in the life history of Stomolophus meleagris were raised in the laboratory

from planula larvae obtained from identified medusae. Planulae were first isolated

in cultures on 5 March 1974 from a holding tank containing medusae of S. me-

leagris collected in the Cooper River at Charleston, South Carolina. Scyphistomae

were observed in these cultures two days later, but all died within a week. Planulae

were obtained a second time from medusae collected 14 July 1981 about 1.5 km

offshore from the entrance of Murrells Inlet, South Carolina. A bucket containing

one male and one female medusa was placed in a mesh bag and suspended overnight

just below the surface of the water. The following morning, water, debris, and

planulae from the bottom of the bucket were poured into large preparation dishes.

Planulae present in these dishes were isolated in a fingerbowl containing 200 ml

of filtered seawater of 35.7%o salinity. The fingerbowl was covered, and the culture

was maintained at room temperature (~27?C) in the laboratory.

Developing scyphistomae were fed pieces of newly hatched Artemia several

times a day until they were large enough to ingest entire nauplii. Undigested ma-

terials were removed daily from the cultures using a pipette, and the water was

changed at least once a week. Strobilation was observed in polyps maintained both

at room temperature and at 25?C in a constant temperature cabinet.

Cultures were lost, apparently due to a bacterial infection, one month after

being established.

DESCRIPTIONS

Planula

Planulae, somewhat flattened in cross-section, varied in outline from elongate-

cylindrical to slipper-shaped to irregularly oval (Fig. la) and measured 120-390

gm long and 60-130 Atm wide. The anterior and posterior ends were rounded. They

were mouthless, and a solid, inner, endodermal mass was enclosed by an outer,

ciliated ectoderm. Living planulae were translucent-whitish in color.

Scyphistoma

At metamorphosis, planulae attached by the anterior end to the substrate and

gradually became flask-shaped as the narrowing stalk of the developing scyphistoma

differentiated from the expanding calyx. Newly metamorphosed scyphistomae

(Figs. lb,c) varied from 200-430 ,m in height from pedal disk to mouth. A thin

cuticle enveloped the slender stalk. Tentacles, usually four in number but sometimes

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 151
LIFE HISTORY OF STOMOLOPHUS

db

FIGURE 1. Planula and scyphistoma stages of Stomolophus meleagris. a. Planula. b. Newly met-

amorphosed scyphistoma. c. Young scyphistoma. d. Intermediate, 8-tentacled scyphistoma. e. Fully

developed scyphistoma. Scale bars = 250 um.

as few as two and as many as six, appeared near the distal end of the bulbous calyx.

They were filiform with scattered nematocyst batteries, and occurred in one whorl.

The oral disk was largely occupied by a prominent, dome-shaped proboscis. The

mouth, round or irregular in shape, was capable of considerable dilation. The color

was translucent-whitish.

Intermediate scyphistomae (Fig. Id) were 0.5-1.0 mm high and bore eight very

contractile tentacles. These were filiform with scattered nematocyst batteries, and

occurred in one whorl. The stalk was slender, and the cuticle originally enclosing

the stalk had become vestigial. The calyx was cone-shaped. A large, flexible, dome-

shaped proboscis occupied most of the oral disk, which bore four peristomial pits.

The expansible mouth was round, irregular, or quadrate, and the pharynx was

quadrate. The color was translucent-whitish, but the endoderm became light orange

after digestion of Artemia.

Fully developed scyphistomae (Fig. le) attained about 2 mm in height from

pedal disk to mouth and bore about 16 filiform tentacles. Tentacles were contractile,

in one whorl, and had scattered nematocyst batteries. The moderately thick stalk

was somewhat variable in length and merged almost imperceptibly with the cone-

shaped calyx. The proboscis was large, flexible, and dome- or knob-shaped. The

mouth continued to be expansible, and was round, irregular, or quadrate in shape.

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 152 DALE R. CALDER

The pharynx was quadrate, and the oral disk bore four prominent peristomial pits.

The color was whitish, with the endoderm becoming light orange after digestion

of Artemia.

Strobila

Elongation of the calyx occurred in the early strobila, but the first clear external

indication of strobilation was the development of a small marginal lobe at the base

of each rhopalar tentacle (Fig. 2a). About six hours later, segmentation began as

a faint circular incision proximal to the tentacular ring; this incision became pro-

gressively deeper and more pronounced (Fig. 2b). A second or even a third incision

d^efU

FIGURE 2. Strobilae of Stomolophus meleagris. a. Early strobila with tentacular lobes. b. Early

strobila with incision. c. Early strobila with second incision. d. Early strobila with developing segments.

e. Mid-strobila, with regressing tentacles and developing ephyral segments. f. Late strobila with well

developed ephyra segments and basal polyp. Scale bar = 500 Am.

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 153
LIFE HISTORY OF STOMOLOPHUS

subsequently appeared proximal to the first in polydisk strobilae, each forming a

segment representing an incipient ephyra (Fig. 2c). As strobilation proceeded, the

proboscis became enlarged, tentacular lobes on the distal segment became more

pronounced, and marginal lobes became progressively better defined about the

periphery of any proximal segments (Figs. 2c,d).

About 36 hours into the strobilation process, tentacles began to undergo regres-

sion, contracting and expanding periodically. The eight rhopalar tentacles were

resorbed, some more rapidly than others (Fig. 2e), into the cleft between the de-

veloping lappets on each marginal lobe. If tentacles in addition to the eight rhopalar

ones were present, they were resorbed into the ocular clefts between the marginal

lobes. Simultaneous with tentacular regression, rhopalia with statoliths became

apparent, lappets elongated, incisions constricting the segments deepened, manu-

brium development progressed on any proximal ephyral segments, a new proboscis

began to form on the basal polyp, and the polyp began to regenerate new tentacles.

Nascent ephyrae, beginning with the one at the distal end, became capable of

contractions. These movements were weak at first, but became progressively

stronger and more frequent. Tentacles of the original scyphistoma were completely

resorbed about 54 hours into strobilation.

In the late strobila (Fig. 2f), incisions continued to deepen and separate the

developing ephyrae. Ephyrae increased markedly in size and underwent rapid de-

velopment prior to release. Pulsations became stronger and occurred more fre-

quently. Development of the manubria in proximal ephyrae and proboscis in the

polyp proceeded. Gastric cirri were visible in the developing ephyra prior to lib-

eration. Ephyrae were liberated about 3.5 days after strobilation began.

Ephyra

Newly liberated ephyrae (Figs. 3a,b) were about 1.5-2.0 mm wide from lappet-

tip to lappet-tip. There were typically eight marginal lobes, eight rhopalia, and

eight pairs of slender, distally pointed lappets. Rhopalar clefts were U-shaped and

slightly more than half as deep as the large ocular clefts separating the marginal

lobes. The manubrium was small ind cruciform in cross-section. Four lips were

often present about the mouth, but oral arms were lacking and no papillae were

present. The stomach portion of the gastrovascular cavity was nearly circular, and

1-2 gastric cirri were present in each quadrant. Eight blunt-ended rhopalar canals

and eight small adradial bulges extended peripherally from the central stomach.

Coronal muscles were barely evident in unstained specimens, but the radial muscles

were visible extending to each lappet from the marginal lobes. The exumbrella was

marked by a ring of small nematocyst batteries about the periphery of the stomach,

and a large, elongate battery was present on each marginal lobe. The mesoglea

was thin. The ectoderm was pale straw colored, while the remainder of the ephyra

was translucent.

GENERAL OBSERVATIONS

Planula larvae of S. meleagris are apparently not incubated by the adult me-

dusae. Medusae of both sexes were dissected and examined microscopically for

incubated planulae during studies in 1974 and 1981, but none were found. Planulae

obtained both years were found free in containers of water in which sexually mature

medusae had been held overnight.

Planulae swam actively by ciliary propulsion, rotating counterclockwise around

the anterior-posterior axis. Within two to five days they attached by the anterior

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 DALE R. CALDER
154

II

bv

FIGURE 3. Newly liberated ephyra of Stomolophus meleagris. a. Subumbrellar view. b. Exum-

brellar view. Scale bar = 500 ,m.

end and metamorphosed directly into scyphistomae, without an intervening larval

cyst stage.

Recently metamorphosed scyphistomae were sessile, but if dislodged were ca-

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 LIFE HISTORY OF STOMOLOPHUS 155

pable of slow locomotion along the bottom of the container using the cilia. Dislodged

polyps eventually reattached through the formation of a pedal stolon, which became

anchored to the substrate. Little is known about asexual reproduction in the polyp

of S. meleagris. However, one scyphistoma with 16 tentacles was forming a

podocyst shortly before loss of the culture.

Strobilae were first observed in the cultures only nine days after polyp mor-

phogenesis occurred. Scyphistomae as small as 1 mm or less, and bearing as few

as eight tentacles, became strobilae. The strobilation process was neither preceded

nor accompanied by any noteworthy changes from the translucent, pale straw color

assumed by the developing ephyrae. In most cases, two ephyrae were formed by

each strobila, although as few as one and as many as three were produced on

occasion. Some of the scyphistomae of S. meleagris strobilated a second time, as

soon as a week after completion of the initial strobilation.

Ephyrae produced during strobilation swam actively and appeared to be normal

morphologically. No attempt was made to raise the ephyrae, and most were pre-

served shortly after liberation.

None of the life stages of S. meleagris examined during this study bore algal

symbionts.

DISCUSSION

Agassiz (1860, 1862) described and illustrated the medusa of Stomolophus

meleagris from specimens collected at Wassaw Island, Georgia, and Charleston,

South Carolina. The species has subsequently been reported from New England

to Brazil in the western Atlantic (Kramp, 1961; Larson, 1976), but its occurrence

north of Cape Hatteras is probably due largely or entirely to transport in water

currents. Elsewhere, it has been recorded from southern California to Ecuador in

the eastern Pacific and from the Sea of Japan to the South China Sea in the western

Pacific (Kramp, 1961; Omori, 1978). In the orient, S. meleagris is one of several

rhizostome medusae used for human consumption (Omori, 1978, 1981).

Medusae of S. meleagris are frequent in coastal waters from North Carolina

to Florida and in the northern Gulf of Mexico (Mayer, 1910). Kraeuter and Setzler

(1975) reported finding medusae of this species from March through October in

Georgia, and Burke (1976) noted that S. meleagris was almost always present in

Mississippi Sound. In South Carolina, they occur sporadically throughout the year

and at times are a hinderance to commercial shrimp trawling because of their

abundance (Calder and Hester, 1978). As with many species of scyphomedusae,

abundances vary greatly from year to year as well as from season to season. Despite

their frequency of occurrence in waters of the southeast and Gulf coasts of the

United States, medusae of this species are an insignificant public health hazard

because the toxin of their nematocysts is relatively innocuous to humans (Toom

and Chan, 1972).

The life history of S. meleagris (Fig. 4) resembles that described for other

species of neritic Scyphozoa (Naumov, 1961; Russell, 1970). The fertilized egg

develops into a tiny, motile planula larva. After swimming freely in the water for

several days, the planula attaches to a suitable substrate and transforms into a

sessile polyp or scyphistoma. Scyphistomae feed and grow, attaining a maximum

size of a few millimeters. They reproduce asexually a number of ways, including

the formation of podocysts and motile or non-motile buds, but only podocyst for-

mation was observed in S. meleagris. In addition to their function in asexual re-

production, podocysts in the Scyphozoa are resistant to adverse environmental

conditions (Cargo and Schultz, 1966). Under favorable conditions the scyphistoma

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 DALE R. CALDER
156

Medusa

tI
r

FJ- Fertilized

Egg

=-) ?

Planula

Young

Scyphistoma

Scyphistoma

Strobila (Fully Developed)

FIGURE 4. General life history of the scyphozoan Stomolophus meleagris. See text for explanation.

undergoes strobilation; during this process the polyp is known as a strobila. Two

separate developmental phenomena, namely segmentation and metamorphosis, are

involved in the strobilation process (Thiel, 1938; Spangenberg, 1968). Strobilation

results in the derivation of several organisms from a single individual. One or more

free-swimming ephyrae may be produced, and the basal portion of the strobila is

left as a small scyphistoma after release of the ephyrae. This small scyphistoma

rapidly returns to normal size and is capable of repeated strobilation. Ephyrae

develop into medusae, completing the life cycle.

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 LIFE HISTORY OF STOMOLOPHUS 157

Scyphopolyps have been described for relatively few species of Scyphozoa.

Known polyps of the order Coronatae are distinctive in being elongate forms en-

veloped in a chitinous tube (Russell, 1970). They resemble, and were believed by

Chapman (1966) and Werner (1966, 1967a,b, 1970, 1973) to be related to, the

fossil Conulata. Chapman (1973) observed behavioral as well as morphological

differences between coronate and semaeostome polyps. Semaeostome and rhizo-

stome scyphistomae display considerable morphological similarity from species to

species within and between orders. The basic form is that of a goblet or cone,

attached aborally by a pedal disk, and bearing an oral whorl of tentacles (Russell,

1970). A cuticular theca, if present at all, is vestigial (Chapman, 1966, 1968).

Among the rhizostomes, scyphistomae are known for Cassiopea andromeda,

Cephea cephea, Cotylorhiza tuberculata, Mastigias papua, Rhizostoma pulmo,

Rhopilema verrilli, and now Stomolophus meleagris (Table I). Of these, S. me-

leagris most closely resembles R. verrilli in its cone-shaped calyx and long proboscis.

The only reliable way to distinguish polyps of these two species at present would

be to induce and observe strobilation. Red and orange pigments appear during

strobilation in the developing ephyrae of R. verrilli that are lacking in S. meleagris.

Studies on the nematocysts of scyphistomae and other stages of S. meleagris,

underway as part of another investigation, may provide a means of separating the

two species. Diagnostic differences in the nematocyst complement were detected

in scyphistomae of the semaeostomes Aurelia aurita, Chrysaora quinquecirrha,

and Cyanea capillata (Calder, 1971).

Scyphistomae of S. meleagris began to strobilate prior to attaining full devel-

opment. Polyps as young as nine days old, measuring 1 mm or less in height and

bearing as few as eight tentacles, were observed undergoing strobilation. Ephyrae

produced by these strobilae appeared normal in all respects. Strobilation so soon

after scyphistoma morphogenesis was unexpected but may not be unusual in the

Scyphozoa, given optimal conditions. Strobilation has also been observed, under

field conditions, in polyps of Chrysaora quinquecirrha that were less than two

weeks old (D. G. Cargo, Chesapeake Biological Laboratory, personal communi-

cation).

Strobilation in the Scyphozoa may be monodisk, in which one ephyra is pro-

duced, or polydisk, in which two or more ephyrae are produced. Although monodisk

strobilation appears to be common in the rhizostomes (Table I), it can no longer

be regarded as a fixed characteristic of the group, as suggested by Sugiura (1966).

Polydisk strobilation was the norm in S. meleagris, and has been observed in three

other species of rhizostomes (Table I). Conversely, Uchida and Sugiura (1978)

demonstrated that strobilation in the semaeostome Sanderia malayensis was

monodisk. Strobilation in semaeostomes such as Cyanea capillata, Aurelia aurita,

and Chrysaora quinquecirrha is usually, but not always, of the polydisk type (Ber-

rill, 1949; Spangenberg, 1968; Calder, 1974). Berrill (1949) concluded that the

type of strobilation depended upon the size and shape of the scyphistoma. Monodisk

strobilation usually occurs in polyps having a short calyx, while polydisk strobilation

normally occurs in polyps having an elongate, columnar calyx. The type of stro-

bilation thus appears to have little or no phylogenetic significance within the Scy-

phozoa.

Several polyps underwent strobilation twice during the month that cultures of

S. meleagris were maintained in the laboratory. However, strobilation several times

in succession is well known in the Scyphozoa, having been reported in species such

as Chrysaora quinquecirrha (Cargo and Schultz, 1967; Loeb, 1972; Calder, 1974;

Cargo and Rabenold, 1980), Aurelia aurita (Thiel, 1962), Cassiopea andromeda

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 TABLE I

A summary of some attributes of seven known species of rhizostome polyps.

No.

tenta- Asexual repro-

Species Shape cles Color Algal symbionts duction Strobilation Strobila color References

Cassiopea an- 32+ greenish present (some- motile buds monodisk (infre- greenish brown
goblet-shaped,

dromeda stalk long, brown times absent quently poly-

(Forskal, in young scy- disk)

1775)* phistomae)

Cephea cephea goblet-shaped, 16 white absent motile buds monodisk

(Forskal, stalk long,

1775) oral disk

width moder-

ate

16 present motile buds monodisk

berculata stalk long,

(Macri, 1778) oral disk

width moder-

ate

goblet-shaped, 16 white absent (zoox- motile buds monodisk

(Lesson, stalk long, anthellae

1830) oral disk could be in-

width moder- troduced)

ate

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 TABLE I-Continued

No.

tenta- Asexual repro-

Species Shape cles Color Algal symbionts duction Strobilation Strobila color References

Rhizostoma cone-shaped, 32 absent motile buds, polydisk Paspaleff (1938)

pulmo (Macri, stalk length polyp buds,

1778) moderate, oral stolon buds,

disk wide podocysts,

strobila buds

Rhopilema ver- cone-shaped, 16+ whitish absent podocysts monodisk (occa- whitish, develop- Cargo (1971)

rilli (Fewkes, stalk length sionally poly- ing ephyrae Calder (1973)

1887) moderate, oral disk) with red and

disk width orange pig-

moderate, pro- ments

boscis clavate

Stomolophus cone-shaped, 16 whitish absent podocysts polydisk (occa- whitish

meleagris L. stalk length sionally mon-

Agassiz, 1860 moderate, oral odisk)

disk width

moderate, pro-

boscis dome-

or knob-

shaped

* Gohar and Eisawy (1960a) and Neumann (1979) have been followed in regarding Cassiopea xamachana Bigelow, 1892 as a synonym of C. andromeda

(Forskal, 1775).

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 DALE R. CALDER
160

(Gohar and Eisawy, 1960a), Mastigias papua (Sugiura, 1963), and Cephea cephea

(Sugiura, 1966).

The ephyra is the least differentiated and most difficult stage to identify in the

development of the medusa of scyphozoans. Russell (1970) distinguished ephyrae

of Aurelia aurita, Chrysaora hysoscella, Cyanea spp., and Rhizostoma octopus

from Britain using morphological differences in radial canal shape, number of

gastric filaments, marginal tentacle development, and arrangement of nematocyst

batteries on the exumbrella. Ephyrae of Aurelia aurita, Chrysaora quinquecirrha,

Cyanea capillata, and Rhopilema verrilli from the east coast of the United States

have been distinguished on the basis of morphology (Larson, 1976) and nematocyst

complement (Calder, 1977). Of these four species, newly liberated ephyrae of S.

meleagris most closely resemble R. verrilli. Unlike R. verrilli, they have (1) faint

instead of prominent adradial bulges in the gastrovascular cavity; (2) a small ma-

nubrium with no papillae; (3) no rose or orange pigments in the stomach and ma-

nubrium endoderm; (4) a different arrangement of nematocyst batteries on the

exumbrella. Ephyrae of S. meleagris were also smaller at liberation than those of

R. verrilli described by Calder (1973), but this may have been due in part to the

small size of the strobilae.

ACKNOWLEDGMENTS

Thanks are due to John Miglarese, David Knott, and Robert Van Dolah of the

Marine Resources Research Institute, Charleston, South Carolina, for assistance

in the collection of medusae from which planulae were obtained. I am indebted to

Renate Carson for translating some papers from German, to Susan Pasch for typing

the manuscript, and to Dr. David Barr for reviewing an early draft. Medusae were

collected during field studies funded by the Coastal Plains Regional Commission

(Contract No. 10340031) and the Coastal Engineering Research Center (Agree-

ment No. W74RDV CERC 77-59).

LITERATURE CITED

AGASSIZ, L. 1860. Contributions to the natural history of the United States of America. Vol. 3. Little,

Brown and Co., Boston. 301 pp.

AGASSIZ, L. 1862. Contributions to the natural history of the United States of America. Vol. 4. Little,

Brown and Co., Boston. 380 pp.

BERRILL, N. J. 1949. Developmental analysis of Scyphomedusae. Biol. Rev. 24: 393-410.

BIGELOW, R. P. 1900. The anatomy and development of Cassiopea xamachana. Mem. Boston Soc.

Natur. Hist. 5: 191-236.

BURKE, W. D. 1976. Biology and distribution of the macrocoelenterates of Mississippi Sound and

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