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Full text of "A monograph of the British naked-eyed Medus© : with figures of all the species"

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Alex. Agassiz. 

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COMPARATIVE ZOOLOGY, 

AT HARVARD COlLEfiE, CAMBRIDGE, MASS. 
JFounSfB bu ptitatt subsniptioii, in 1861. 

Deposited by ALEX. AGASSIZ. 

No. 3 C, S'3 t- 



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RAY SOCIETY 



INSTITUTED MDCCCXLIV. 



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LONDON 



MDCCCXLVIII. 



MONOGRAPH 



BRITISH NAKED-EYED MEDUSAE 



JTigurrs of all tftf 0mit^* 



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EDWARD FORBES, F.R.S.; L.S.; and G.S. 

PROFESSOR OF BOTANY IN KING's COLLEGE, LONDON ; 
AND PALAEONTOLOGIST TO THE GEOLOGICAL SURVEY OP THE UNITED KINGDOM. 



LONDON : 

PRINTED FOR THE RAY SOCIETY. 



MDCCCXLVm. 



rWNTED BY C. AND J. ADLAKD, BAKTHOLOMEW CLOSE. 






PREFACE. 



This short preface is Intended both as an explanatory notice and as a 
dedication. Without the aid of my friend Mr. M' Andrew, the following- 
Monograph could not have been drawn up. The mass of the materials 
employed in its construction was the produce of several delightful scientific 
cruises, when I accom]}anied him in his Yacht, the Osprey. Nine 
years ago this history was commenced : were it not for the ardent friend 
of science, whose name I wish to associate with my work, as many more 
would be required for its completion. Even now, I can offer only an 
outline of a most curious and interesting, though neglected, department of 
British Zoology. The greater part of the matter in this essay is new. With 
one exception — kindly communicated by Mr. Alder — every species has been 
examined by myself. Every figure is original. Any defects in the 
engravings must be laid to my charge ; their merits are due to my friends 
Mr. W. Baily, and Mr. C. R. Bone, for whose exertions I have to return 
many thanks, 

Edward Forbes. 

London ; June 25, 1848. 



CONTENTS. 



PAGE 

Subject of the Work . . . . • ■ -1 

General Structure of the Naked-eyed Medusae . . . . . 2 

Muscular System . . . . ■ . .3 

Nutritive System . . . . . . . 4 

Reproductive System . . . . . . .6 

Organs of Sense . . . . • . . . 8 

Power of Stinging . . . . . .10 

Phosphorescence . . . . . . . 11 

Development . . . . . .14 

Table of Genera . . . . . . . 17 



Account of Genera and Species ..... 19-74 



Eaumeration of higher British Discophorae . . . 75 

Classification of Medusae . . . . . .78 

How to Observe and Preserve them . . ... 89 



Bibliographical Appendix . . . . .91 

Index of Genera and Species described . . . . . . 103 



PLATES. 



I. Species of Willsia and Cikce. 
II. Species of Oceania and Saphenia. 

III. Species of Oceania and Turris. 

IV. Species of Stomobeachium and Polyxenia.* 
V. Species of Tima and Geryonia. 

VI. Species of Slabberia and Sarsia. 
VII. Species of Modeeria and Sarsia. 
VIII. Species of Thaumantias. 
IX. Species of Geryonopsis and TflAUMANTiAs.f 
X. Species of Thaumantias. 
XI. Species of Thaumantias. 
XII. Species of Bougainvillea and Lizzia. 
XIII. Species of Euphysa and Steenstrupia. 



* 111 this plate for Polyxenia cyanostyhi, read Polyxenia Alileri. 

t 111 this plate for Thaumantias cymbaloidea, read Geryonopsis delicatula. 



BRITISH NAKED-EYED MEDUSA. 



The creatures which I am about to describe and dehneate in the following monograph 
are animals of very simple organization and beautiful form. They are members of the lowest 
section of the Animal Kingdom, and are intimately allied to the polypes, as we shall see when we 
come to consider their classification, which will be best understood after we have examined their 
structure. They are mostly minute, often microscopic, though many of their nearest relations, 
such as the great stinging Medusae, grow to a considerable bulk. They are active in their 
habits, graceful in their motions, gay in their colouring, delicate as the finest membrane, 
transparent as the purest crystal. They abound in the sea, but are not equally plentiful at 
all seasons. They have the power of emitting light, and when on a summer's evening the 
waves flash fire as they break upon the shore, or glow with myriads of sparks as they curl 
and froth around the prow of the moving ship or under the blade of the striking oar, it is to 
delicate and almost invisible Medusae that they chiefly owe their phosphorescence. 

They belong to that section of Acalephae termed by Eschscholtz Discophora : the upper 
portion of the body being formed in the shape of a hemispheric disk. All the Discophorae 
may be conveniently arranged in two great groups : the first consists of those which have the 
eye-like bodies or ocelU of their margin protected by more or less complicated membranous 
hoods or lobed coverings, a character which accompanies one of great importance, viz. their 
possession of a much ramified and anastomosing series of vessels. This section I propose to 
name Steganojythalmata {ari-yavoQ, covered). 

The second division includes all those which have the ocelli naked, often aborted, and 
which possess a very simple vascular system, the circulating canals proceeding to the margin 
either altogether unbranched, or if divided, not anastomosing with each other. These I term 
the naked-eyed Medusje, or G ijmnopthuhiuta (yvfxvog, naked). 

It is to the history of the British species of the second division that this monograph is 
devoted. The observations embodied in it are the fruit of several years' research, having 
been commenced in the year 1839, and continued every summer, either in the British seas or 
abroad, until the autumn of 1846, when an account of them was read, for the first time, at the 
Southampton Meeting of the British Association. That year and the previous summer were 



2 ACCOUNT OF THEIR STRUCTURE. 

by far the most prolific in results ; the voyages which I had the pleasure of making with my 
friend Mr. M'Andrew aroimd the British coasts having afforded admirable opportunities for 
the study of our Medusae. These creatures are of so very delicate and often unpreser\-able 
a nature, that casual circumstances usually determined the extent to which the examination of 
their structure and habits could be pursued, and as most of my observations had necessarily 
to be made at sea, those circumstances were not always most favorable. In the present 
very unsatisfactory state of this branch of zoology, however, I do not think it necessary to 
apologise for unavoidable imperfections, for having often experienced the difficulty of con- 
ducting inquiries into tribes of which the species had as yet been but vaguely defined and 
rarely figured, I trust this account of an important and beautiful tribe of animals, of which so 
far as the British seas are concerned, only a few very fragmentary notices are accessible, may 
serve as a basis for future and more extensive researches. They offer a fresh and but little 
explored field for discovery. Their organization is but partially understood, and much requires 
to be done before the signification of their several parts be fully made out ; of their habits 
we possess but very slight knowledge. Their development is a subject of the greatest interest, 
seeing that upon its clearing up will probably depend the future classification of the zoophytes. 
On most of these points I can scarcely pretend to speak ; what I offer are the rudiments only 
of an extensive subject. It is for naturalists expert in physiological and anatomical investi- 
gations, skilled in the use of the microscope, and not too trustful in its revelations, free in 
their movements, and with time untrammelled at their disposal, to carry out this most interesting 
branch of research, to which, if my imperfect monograph give an impetus, I am content — 

Quod pofui, feci ; quod restat suppleat alter 
Doctior, et nostris faveat nan invidus ausis. 

Before commencing a detached description of the species, it is best to examine the 
features of organization common to the tribe. 

The parts presented by these animals are the following : 

A. The disk or umbrella. This forms the greater portion of the animal's body. It is 
hemispheric, but varies from being extremely depressed and almost plane, as in certain species 
of Thaumantias and Mquorca, to a nearly cylindrical form, as in Turris. One of its commonest 
shapes is that of a round glass shade, such as is placed over ornaments or statuettes to 
protect them from dust. It is usually smooth, rarely pilose. Its under surface, on which in 
certain tribes the reproductive bodies are placed, is called the suh-umhrella. Around its 
margin internally there is in many species a projecting ledge of membrane called the veil 
(velum). The margin itself is usually provided with more or less numerous tentacles (cirri 
mar finales), of variable structure, the bases of which are often swollen into a bulb, and 
deeply coloured or marked with a brilliant spot {ocellus). In the substance of the disk are 
the vessels, often conspicuously visible. 

B. From the centre of the sub-umbrella hangs a more or less produced proboscis-like 
body {pedunculus) , of variable form and dimensions. In this is the stomach, and, in certain 
genera, the ovaries. At its extremity is the mouth, surrounded by variously-formed contractile 
lips, occasionally furnished with produced tentacula. 

Such arc the characters visible at a glance. A more minute examination makes us 
acquainted with the structures they include. 



MUSCULAR SYSTEM. 3 

Substance. — When we examine a naked-eyed Medusae by polarized light, we see that at 
least two distinct tissues enter into its composition ; these are equally distinguishable by the 
naked eye, if the creature be sufficiently large, when we see the one presenting the aspect of 
a transparent, and almost always colourless gelatinous membrane, the other a translucent, 
and as if granular, substance. The former constitutes the mass of the body; the latter 
forms the margin of the mouth, the edge of the umbrella, and the tentacula. The first 
is immoveable and uncontractile, elastic, but not extensile ; the second is highly contractile 
and active. They are both composed of cells, those forming the active tissue differing in being 
nucleated cells of the fibrous order, and intermingled with granular corpuscles. The former 
are covered with a fine amorphous smooth epidermis, beneath which, in the higher forms and 
in the so-called Oceania cruciata — possibly in most species — are cells containing a spiral 
thread. Such cells are also present on the surface of the tentacula in many species, exactly 
as in the hydroid polypes. Will described the cells beneath the epidermis of Geryonia as 
round, transparent, and lobed. A ciliated epithelium has been observed by Will in the inner 
surface of the lip-ring in Geryonia, also in the tentacular canals. It lines (as I have seen) 
the gastro-vascular canals in Tliamnantias, and probably in all the genera. Several of the 
higher Medusae (conspicuously those of the genus Cyanaa) have the power of stinging 
severely. The power resides in the skin, and, especially in some of the appendages of the 
sub-umbrella, appears to be always connected with the second or granular tissue. Wagner 
has attributed it to the fihferous vesicles, which, in some species if not in all, have the 
power of projecting the contained thread with its barbed extremity, even as the hydroid 
polypes and the Actinese do. But as many Meduste and Actineae provided with these curious 
organs do not sting, such explanation is doubtfid. I have never found any of the naked-eyed 
species to sting. 

Muscular System. — The motor tissue in these Medusae is of the simplest kind, and 
consists, in most cases, simply of bands of the granular substance just described. In certain 
genera, especially in Tiirris, the motor bands exhibit a distinct fibrous arrangement (Plate III, 
f. \, e, and f. 2, /), and Professor R. Wagner has stated that distinct muscular fibres with 
transverse striae are present in the " Oceania cruciata" (a Thaumantias ?) of the 
Mediterranean.* Will has observed a few longitudinal fibres in the motor ring of a 
Geryonia. In the higher Medusae the muscular system is much more developed, especially in 
Rhizostoma, the movements of which may be shown experimentally to depend on the mus- 
cular bands hning the sub-umbrella. I have paralysed one side of a lihixostoma Jldrovandi, 
whose disk measured more than a foot across, by removing with a scalpel the bands of that 
half, whilst the other side contracted and expanded as usual, though with more rapidity, as if 
the animal was alarmed or suffering. All the Medusae when irritated become much more 
rapid in their movements, and contract and expand their disks or bodies in a hurried and 
irregular manner, as if endeavouring to escape from their persecutors. In the naked-eyed 
species, the muscular system usually consists of a marginal motor ring, the tissue of which is 
continuous with the tissue of the marginal tentacula ; concentric rings of motor tissue forming 
the walls of the tentacles themselves, and a ring of similar tissue forming the margin of the 

* Ueber den Bau der Pelagia noctiluca und der organization der Medusen. 1841. 



4 NUTRITIVE SYSTEM. 

lips. To these there is superadded, in the genus Turris, longitudinal, highly-developed muscular 
bands, running from the base of the peduncle to the marginal band. Whatever be the 
arrangement, the movement is the same. The animal swims in an oblique position, con- 
tracting and expanding alternately its umbrella; occasionally pausing as if to rest, but capable 
of continuing its motions for an indefinite time. The lips can be expanded or contracted as 
occasion may require to seize its prey. The tentacles in many species are capable of wonderful 
extension, and can be retracted suddenly into a very small compass, often into a mere tubercle ; 
but there are many naked-eyed Medusae which vary their tentacles at an almost uniform 
length. Each of these organs may be extended or contracted singly, or in concert with its 
fellows, evidently obeying promptly the will of the animal of which they form part. They 
guide the Medusa through the sea, and can anchor it. I have seen a Geryonia anchor itself 
by means of its lips, clasping a coralline with them, and remaining tranquil so fixed for a 
considerable time. 

Nutritive System. — This consists of a stomachal cavity^ excavated in a more or less 
produced proboscis, depending from the summit of the sub-umbrella, opening externally by a 
more or less expanded mouth, margined by variously-formed contractile lips, and superiorly 
communicating with a system of radiating canals, which run to a common marginal canal. 
The orifices of these canals probably in every case open into a common cavity or intestinal 
reservoir superior to the stomach, though sometimes stated to open directly into the latter. 
The true position of the stomach in these animals has been a subject of much dispute, which 
is not to be wondered at, considering the extreme variations presented by the central peduncle. 
It was indeed for a long time supposed that several of the Dlscophone had no true mouths, 
but absorbed, as if by suckers or roots, their nourishment from without, a view, however, 
which all the more recent researches have tended to disprove. By some natui-alists, the 
cavity above the cavity of the peduncle has been regarded as tiie stomach, and the latter as a 
pharynx, a view which has been partially supported by Milne Edwards. Eschscholtz made 
the mistake of supposing the ovaries in the naked-eyed species to be stomachs. Will, and 
more recently Frey and Leuckhart, regard the peduncular cavity only as the stomach, a view 
which, certainly among the gymnopthalmatous Discophoraj, I hold from my own observations, 
for I have observed that the process of digestion goes on wholly in that cavity. Its dimen- 
sions vary greatly ; among our British forms, it is especially large in Stomobrachium, a genus 
which approaches nearly JEquorea, where it is almost an open space surrounded by a slight 
veil of membrane. In Tun-is and Oceania, it is also large and well defined. In Willsia and 
Thaumantias, it is campanulate, and occupies the greater part of the peduncle. In Geryonia 
and Tima, it is small, in comparison with the peduncle, and confined to its extremity. 
In Slahheria, Sarsia, and Steenstriqna, it is tubular, but can assume a bell-shape. In 
Bougainvillea and Lix^ia, it is a conical cavity, with singularly branched lips. The commu- 
nication of the stomach with the gastro-vascular canals is not clearly made out in all the 
genera. Will, in his description of Geryonia ijellucida, states that at the fundus of the 
stomach there are four small obtuse prominences, each of which presents a small aperture 
which is the orifice of one of the water-canals. In another species, the base of the stomach 
into which the vessels opened seemed to be separated from the remainder. In Thaumantias 
leucostyla, he found a distinct cavity separated from the stomach at its base, the walls of 



VASCULAR SYSTEM. 5 

which were Uned with \'ibratile ciha ; from this cavity the vessels sprang. In Mquorea, 
Milne Edwards describes the canals as opening directly into the large and gaping stomachal 
cavity. My own observations accord with those of Will, to the effect that, in most cases 
(among the Gymnopthalmata), there is either a well-defined cavity at the base of the stomach 
into which the vessels open, or an indication of such a cavity. This I regard as homologous 
with the sac so distinctly separated fi-om the digestive tube in the CiUograda, and into which 
the vessels from beneath the rows of cilia open. The superior cavity in both cases may be 
regarded as an effort towards a specialization of the respiratory system — a view first suggested 
by Will. From it the circulating fluids flow into the gastro-vascular canals, which all run 
without dividing, except in the case of Willsia, into a common marginal vessel, caecal 
projections of which, in several instances, appear to be prolonged into the marginal tentacula. 
Will, however, observes, and I can confirm his remark, that the canals of the tentacles in 
ThmmmnUus do not communicate with the vessels. The walls of the gastro-vascular canals 
are ciliated. The fluid within becomes coloured, according to the food taken by the animal. 
I have seen it in a Thaiimantias fed upon small Crustacea turn completely yellow. 

The system of vessels, partly nutritive, partly respiratory, proceeding directly from the 
stomach, or from a cavity opening directly into it, may be regarded as a good instance of 
phlebenterism. Dr. Will, however, regards it as an aquiferous system, and describes a 
circulatory system distinct from it. He asserts that in Geryonia all the water-vessels are 
accompanied by blood-vessels, which spring from the sides of the stomach, and proceed to its 
base, there to run alongside of the water-vessels. He states that they are distinctly to be 
recognised on both sides of the latter, especially when they contract ; then the blood-vessels 
remain expanded, and appear much thicker. At the circular marginal water-vessel the blood- 
vessel is usually observed only on one side, and that at the lower. Sometimes there appears 
a narrow margin, filled with blood-corpuscles at the upper edge. " The contents of the 
blood-vessels usually consist of a clear fluid, in which a great number of finely granular 
corpuscles, of a diameter from 1-400 — 1-500'", are floating." He observed similar blood- 
vessels in Thaumantias. 

Aware of these obsen-ations, and of the accuracy of the observer, I made every endeavour 
to satisfy myself on the matter with the species of several genera. But though I sometimes 
fancied I saw such vessels, in the end I came to the conclusion that the appearances were 
deceptive. No such vessels appear to have been noticed by Wagner or Milne Edwards. 
Will described similar vessels distinguished by this red colouring in Beroe. I have seen the 
appearances to which he alludes, but could not satisfy myself of their vascular nature. Frey 
and Leuckhart, also, with Will's observations before them, have sought in vain ; their remarks 
upon this subject are so much to the point that I quote them verbatim : " Our attention whilst 
investigating was hkewise directed to this point, but without discovering the characters 
mentioned (i. e. by Will). Neither in Cyd'qype, nor in Geryonia, nor in Cyauceu, did we 
succeed in discovering particular blood-vessels in addition to the canals of the abdominal 
ca\ity. W^e may assert, with particular distinctness, that the species (new ?) of Geryonia 
obsel^^ed by us is altogether deprived of a peculiar system of blood-vessels, although Will 
has recognised such in Geryonia pclliicidu, described by him, and from which our species 
is principally distinguished only by having marginal tentacles of equal length. A result. 



6 REPEODUCTIVE SYSTEM. 

entirely correspondent with ours, was likewise obtained by our respected friend Professor 
Bergmann, during numerous observations which he had an opportunity of instituting, during 
a residence at Iceland, on many different Medusae. Will himself does not appear to have dis- 
covered any particular system of blood-vessels in all Acalephse . . . We believe ourselves justified 
in pronouncing an opposite opinion, ^'iz., that no special blood-fluid, or any special vascular 
system, exists in many of the Acalephis, except the nourishing fluid contained in the abdominal 
cavity. Some Medusae, however, may possess such. This can be the less objected to d priori, 
since we know that the development of the vascular system in the different classes is subject to 
very considerable differences. The statements of Will are moreover, in many respects, too 
decided, as not to allow us altogether to doubt the correctness of his observations, although it is 
the very peculiar behaviour of the blood-vessels, as described by him, which justifies us in 
believing in the possibility of an error ; in addition to which, we must mention that our own as 
well as Bergmann's observations have furnished quite a different result from those of Will. 
Indeed the question can only be decided by new and careful observations."* 

With these judicious remarks I entirely agree, and hope the suggestion of further inquiry 
will be taken up by some of our expert microscopical observers, qualified for such an inquiry 
by possessing the requisite physiological knowledge, without which microscopical researches 
must always, and justly, be received with distrust. 

Reproductive St/stem.- — The majority of the naked-eyed Medusae have very distinct 
reproductive glands. These are placed either on the surface of the sub-umbrella, or on the 
inner and upper part of the peduncular cavity. In each case their position has a distinct 
relation to the arrangements of the gastro-vascular canals. Instances of the former 
arrangement are seen in Stomohrachium, Geryonia, Thaiimantias, Circe, and Slabheria ; 
of the latter, in Turris, Saphenia, Oceania, and Willsia. They are not so definitely marked 
in Sarsia, Steenstrupia, and Modeeria, in which genera the whole of the substance of the walls 
of the peduncle seems to be composed of a germ-producing tissue. In Bougainvillia and 
Lizxia, the condition of the reproductive glands is intermediate between the two modes just 
described. In Euphysa, the ovary appears to depend from the centre of the peduncular cavity. 

The organs of generation in these Medusae were long misunderstood. Peron and Lesueur 
recognised their true position in most of their " monostomous gastric Medusae," in which the 
genera Oceania and JEquorea, according to their view of the extent of those groups, were 
placed along with Pelagia, and other forms having no true immediate affinity. In the case 
of ^qtiorea, however, they did not recognise the ovaries. In their group of " agastric 
Medusae," of which Geryonia maybe cited, they seem altogether to have misunderstood these 
organs. Their importance and meaning were equally lost sight of by Lamarck. Eschscholtz 
still further lost the clue to their signification ; for, founding his system upon the supposed 
manifestation or obscurity of the reproductive glands, he divided all the Biscophora into 
PhanerocarjJCB (exactly equivalent to my Steganopthalmata) and Cryptocarptp, the latter 
group including the naked-eyed forms, describing the generative organs in the latter as 
stomachs or appendages of the stomach. Cuvier, misled probably by Peron, does not seem to 
have clearly comprehended the signification of the parts in this section of the Medusae, though 

* Frej' and Leuckhavt. 



REPRODUCTIVE SYSTEM. 7 

practically versed in the higher group. Blainville recognised the ovaries in Oceania (following 
Peron and Lesueur), hut mistook them (following Eschscholtz) for stomachal appendages in 
Thaumantias, Geryonia, and allied genera. Lesson, confused throughout, repeats the same 
mistake. Ehrenberg recognised their true position in his " McUcertmn campamdatum'' (i. e. 
Stomobrachium octocostatuni) and Oceania pileata. Sars first described them in Thaumantias 
and Stomobrachium as dilatations of the gastro-vascular canals ; but more lately recognised 
their true office. Milne Edwards demonstrated their true nature in JEquorea violacea, and 
inferred their office by analogy in all the so-called Cryptocarpce. Brandt appears to have 
followed Eschscholtz. Will perfectly comprehended their true nature, and demonstrated their 
structure. Frey and Leuckhart take the same view. Indeed, it seems strange that such great 
diversity of opinion and so much error should ever have prevailed respecting the position of 
the glands of generation in the naked-eyed Medusee, especially when their true nature in the 
steganopthalmatous species was recognised by all. 

In Turris, the genus which may be regarded as the highest in organization of the order 
under consideration, the ovaries are highly developed, and line the upper part or chamber of 
the stomachal cavity in the form of convoluted tubular and fimbriated membranes, conspicuous 
from their brilliant colour. Such an arrangement closely approaches that met with in the 
Steganopthalmata. In Oceania, a similar arrangement, though not so perfectly made out, 
prevails. In Geryonia, Thaumantias, and allied genera, the ovaries are more or less clavate 
or leaf-shaped, and are either expanded on the under surface of the sub-umbrella in the course 
of the gastro-vascular canals, or depend from it as membranous sacs or laminse, the latter form 
being that seen in Stomobrachium and JEquorea. In Willsia, Bougainvillia, and Lixzia, 
they present the appearance of lobes on the sides of the stomach ; but those of the first-named 
genus are much more regular and normal than those of the two latter. Their number, when 
well defined, may be very considerable (as in ^quorea and Mesonema) ; but in our British 
forms the greatest number is that seen in Stomobrachium and Circe, where there are eight ; 
next, Willsia, which has si.x ; the remaining genera have four ovaries, each of which, in 
several instances, is composed of two equal and similar parts. 

Though I have used the word ovaries for these bodies, as if the animals were unisexual, 
I have done so only as a convenient form of speech. There is every reason to believe that the 
majority of the Medusae are bisexual, though the two sexes appear to be united, but maintained 
by distinct organs in certain forms, especially in the higher group. The dioecious character 
of the naked-eyed forms has been demonstrated by Milne Edwards, Wagner, and Will. The ' 
first-named naturalist showed that some individuals of JEquorea violacea were females, having 
eggs in their generative organs, others on which there were no trace of eggs, but abundance 
of spermatozoa, being males. 

Will describes the sexual organs of Geryonia pelliicida as lying in the course of the 
radiating gastro-vascular canals, their further extremities rounded, the inner ends pointed. At 
the latter he found ducts of emission which reach the base of the peduncle. Each gland 
consists of two lancet-shaped halves ; each half is provided with a special duct of emission, so 
that there are consequently eight ovaries in the female, and eight testicles in the male, of this 
genus. " The ovaries are twisted sacs in which the eggs lie close to each other, the largest 
towards the margin of the disc, the smallest towards the peduncle. The perfectly-developed 
eggs are of a whitish colour, opake, and measure 1-8'"; the germinal spot is round and 



8 ORGANS OF SENSE. 

1-200'" in size. The males cannot be distinguished from the females either by shape or size 
of the body, or by the form of the sexual glands." Will fancied these glands had a greenish 
glitter in the male, which was not present in the female. " The testicles are likewise twisted 
sacs filled with spermatozoa. The latter consist of a thick oblong body, measuring 1-800'" 
and an extremely slender, long tail, which is only visible during vibration." Will found at all 
times as many males as females. 

Organs of Sense. — The lips and their appendages, the marginal tentacula, and the bulbs 
at their bases, may be enumerated under this head. The lips and the tentacula are instruments 
of touch and prehension, the former chiefly for the purpose of seizing the animal's prey, and 
sometimes, as I have seen in the case of a Geryonla, for anchoring the body. The lips vary 
much in form. They are sometimes (as in Circe, most species of TJiaumaiitias, and 
Polijxenia) simple lobes ; in other cases (as in Tnrris, Geryonia, and Oceania) fimbriated lobes ; 
in BouguinviUea and Zizxia, they are furnished with single or branched tentacular processes, 
reminding us of the curious gland-tipped cirrhi, which are so conspicuous in the genus 
Cassiopeia among the higher Discop/iora, and which were long supposed, and are usually 
still described to be roots or suckers for the purpose of absorbing nourishment. In Sarsia, 
Slabberia, and Steenstrujjia, the lip is a simple ring around the orifice of the tubular digestive 
cavity. The tentacula in all our British examples of the naked-eyed Medusae, are simple and 
usually filiform, though highly contractile, and in some species often reduced almost to a 
point. In Slabberia we have an abnormal form of these organs, their termination presenting 
the appearance of a bulb. In Euphysa, the single tentacle is clavate and different in structure 
from that of any other British genus. In the same curious form all the tentacles except one are 
aborted, a remarkable modification seen also in Steenstrupia. In a new species of Geryonia, 
here figured, alternate tentacles are glanduliferous. In not a few species there are two 
varieties of tentacles placed in a single series round the margin, but the majority have the 
tentacles only of one kind. 

At the base of the marginal tentacula or cirrhi there are present in a great many of these 
animals coloured spots or bulbs. In some species (as in Thaumantias pilosella, Slabberia 
halterata, Willsia stellata, Lixzia octo-punctata, &c.) these points are very strongly coloured, 
and from their magnitude indicate the course of the animal when in motion, appearing like 
a circle of gems in the water. Where some of the tentacula are aborted (as in Steenstrupia 
and Euphysa), they are not aborted with these organs, but are all conspicuously developed ; 
in many forms only certain tentacles have bulbs at their bases. In other forms, the tentacula 
are present and highly developed, but no coloured spots or bulbs are seen at their bases, as in 
certain kinds of Geryonia and Circe. When these bulbs are examined under the microscope, 
we find their organization more complicated than at first glance it would seem to be. In the 
majority of species, perhaps in all, these bulbs, whether conspicuous from colouring or not, 
contain a small cavity quite distinct from any coloured spot which may be present. The 
former is the otoUtic vesicle, the latter the ocellus. 

The ofolitic vesicle, which, from analogy and its peculiar structure, is considered an 
organ of hearing, is a small spherical sac developed in the midst of the granular substance of 
the bulb, and containing more or fewer minute vibrating bodies. Will has described the 
otolitic vesicle and its contents in a Geryonia as follows : " The auditory vesicles are seated 



OCELLI. 9 

in the course of the marginal circular vessel in very uncertain number, usually, however, one 
at each side of the larger marginal cirrhi, and beside the smaller one, only at one side. They 
are round, measuring l-40thofa line in diameter, and consist of a tolerably thick membrane : 
they contain from one to nine, and even more, round globules. If there is only one, it is situated 
exactly in the centre of the vesicle, but if there are several, they are lying together either in 
two groups or separately joined to each other at the wall of the vesicle. Their size varies 
from 1-300 — 1-150'". I have never observed them move. Muriatic acid dissolves them, and 
causes the vesicle to burst." In his Thaumantias leucosUjla, he describes the auditory vesicle 
as "measuring 1-60'", and containing globules of the dimensions 1-200'". They are seated 
beneath the basis of the marginal fibres on a small projection. They arc not present, however, 
beneath all the marginal fibres." Milne Edwards obsci-ved, in his yEquorea molacea* two 
hemispheric or oval vesicles on each side of each marginal tubercle, and containing two or 
three spherical corpuscles. Kolliker observed that the otolitic cavities or vesicles in Oceania 
(as well as in higher forms) were lined with vibratile cilia, and that the otolites vibrated. Frey 
and Leuckhart, whilst they saw the otolites vibrate distinctly in certain Ciliograda, found them 
perfectly motionless in Genjonia, even as Will had observed. I have observed the vibration 
of the otolites distinctly in more than one species of Thaumantias ; so has my friend Dr. 
Melville. I have seen them also vibrating in their cavities at the bases of the tentacles of 
more than one species of Oceania, a genus in which they are highly developed. 

The ocelli, from analogy, are regarded as rudimentary eyes, or rather light-perceiving 
organs. In the gymnopthalmatous Medusae they are very rudimentary, and in most species 
consist only of an assemblage of pigment- cells more or less symmetrically disposed. They 
vary much in colour, different species of Thaumantias, for instance, presenting purple, orange, 
yellow, black, and even variegated ocelli. Yellow, with a red dot, is a common appearance. 
This dot indicates a higher or more concentrated condition of the organ. It is especially 
defined in Oceania, and in Turris neglecta, forms at the head of the tribe. In Slabberia, 
the resemblance of the ocelli to the coloured bulbs which terminate the tentacula is very 
striking, but when minutely examined, they are easily distinguished from the latter organs 
by the presence of a small black dot. In some forms of Sarsia and in Euphijsa we have 
curiously particoloured ocelli ; also in Willsia, though not so defined. In Zixzia, and 
especially in BougainvUlia, we have compound ocelli, formed out of several united, and 
variously colovn-ed, either entirely black, or entirely yellow, or piebald, black and yellow, or 
yellow and bright red. In Circe, and some other forms, no ocelli can be observed. 

That these bodies are the eye-spots, there can be no doubt, when we compare them 
with similar bodies in the higher Medusse. In them crystals are present, as was first 
pointed out by Gaede. These crystals were shown by Rosenthal to be silicious, a character 
by which they are strikingly distinguished from otolitic crystals, which are always 
calcareous. 

Though, as we have seen, there are well-marked organs of sense in these animals, the 
presence of a nervous system has not been clearly made out. For my part, I have not been 
able to satisfy myself as to the existence of either ganglia or nervous filaments in any of the 

* Ann. Sc. Nat. (2d Ser.) t. xvi, p. 195. 



10 POWER OP STINGING. 

naked-eyed Medusae, though I have seen appearances, both in the higher Discophorce and in 
the Ciliograda, which would induce me to admit their presence in some Acalephce. Will 
has observed that in Geryonia there is a small cavity beside the otolitic vesicle, which is 
filled with a yellowish-green matter, in which the vesicle itself is bedded to a third of its 
circumference, and he considers this a ganglion, whilst he admits it cannot be proved to 
be so histologically. I believe I have seen a similar appearance in several species, but 
not so constantly as to permit of the assignment of so important an office as the duty of 
a nervous ganglion to the tissue. Frey and Leuckhart recognised the same bodies in 
Geryonia, but doubt their nervous nature, and remark that the individual masses in 
this instance did not seem to be sufficiently distinctly separated from the neighbouring 
parenchyma, as to warrant their concluding with certainty that such bodies are peculiar 
isolated formations. 

Power of Stinging. — In the minds of most people who have been at the sea-side 
the notion of a Medusa naturally associates itself with that of a nettle, since both the animal 
and the plant enjoy an equal reputation for their stinging powers, and for the production of 
an extremely similar, though not the less unpleasant sensation, when incautiously handled or 
inadvertently touched. The term Acalephce, so frequently applied to the whole of the 
Medusa tribe, is significant of their nettle-like nature. Yet it is not improbable that this 
offensive faculty of stinging is possessed by only a small minority of the sea-jellies — a 
minority chiefly, if not wholly, composed of the steganopthalmatous species. Among them 
the Cyancea capillata of our seas is a most formidable creature, and the terror of tender- 
skinned bathers. With its broad, tawny, festooned, and scalloped disk, often a full foot or 
even more across, it flaps its way through the yielding waters, and drags after it a long train 
of riband-like arms, and seemingly interminable tails, marking its course when the body is far 
away from us. Once tangled in its trailing " hair," the unfortunate who has recklessly 
ventured across the graceful monster's path, too soon writhes in prickly torture. Every 
struggle but binds the poisonous threads more firmly round his body, and then there is no 
escape ; for when the winder of the fatal net finds his course impeded by the terrified human 
wrestling in its coils, he, seeking no combat with the mightier biped, casts loose his enve- 
nomed arms and swims away. The amputated weapons severed from their parent body vent 
vengeance on the cause of their destruction, and sting as fiercely as if their original proprietor 
itself gave the word of attack. The Cyancea Lamarcldi possesses a like dangerous power, 
and Pelagia cyanella also, though very faintly, as I have experienced. But, unless Chrysaora 
hysoscella sting, no other Meduste of our seas besides those mentioned, have been observed, at 
least by me, or naturalists known to me, to possess this noxious property. I have in vain 
endeavoured to elicit such nettling proofs of rage in any of the naked-eyed species, though I 
have stirred, and grasped, and rubbed together hundreds of them belonging to many genera. 
It is right, however, to notice this matter, for it may yet be found that either at particular 
seasons, or under peculiar circumstances, more than one species can sting. Dicquemare has 
stated that certain species of Oceania sting, though very slightly, and only when they come 
in contact with very sensitive parts, such as the eyes. Not being ambitious of sufi'ering 
stone-blindness by playing too closely with even the smallest gorgon's head, I have never 
ventured to repeat the worthy Abbe's experiment, and prefer keeping my eyes intact to 



PHOSPHORESCENCE. 11 

poking Medusae into them. For such rash experiments, Ben Jonson's song might be 

paraphrased — 

" O do not wanton with those eyes, 
Lest you be sick with seeing," 

— and not bad advice either. 

In such Medusae as do sting, the power has been beheved by Dicquemare, Eysenhardt, 
and others who have practically looked to the subject, to reside in a mucus which can be 
thrown off by the animal. Certainly such mucus, as I have often experienced, retains its 
urticating properties for some time after being detached from its producer. If the view which 
has been of late mentioned, that the filiferous capsules with their barbed projectiles are the 
causes of the stinging sensation, the power of the mucus to sting does not contradict it, for 
usually in it numbers of filiferous capsules may be perceived under the microscope. 

Phosphorescence. — Whatever doubt there may be about the stinging faculty of the 
naked-eyed sea-jellies, there can be none about their capability of emitting light in the dark. 
This wondrous power, possessed by comparatively few inhabitants of the air, is a gift bestowed 
on many of the dwellers in the waters, and is especially possessed by creatures of the order 
,Radiata, as if, to use a fanciful analogy, their star-like forms had given them star-like 
properties. The true polypes exhibit the phenomenon of phosphorescence in great perfection, 
but as the majority of them are fixed, at least in their supposed most perfect condition, they 
can play but little part in producing the luminosity in the sea, as seen by ordinary observers. 
Many annelides and other articulata are phosphorescent, and even starfishes of the Ophiura 
tribe, as has recently been shown by Quatrefages, but for similar reasons they are not likely 
to be chief producers of the light. It is mainly by the Arachnodermata and minute animals 
closely allied to them that the waves at night are — 

" Spangled with phosphoric fire 
As though the lightnings there had spent their shafts. 
And left the fragments glittering in the field."* 

The phenomenon of the luminosity of the sea was known to be produced by Medusae 
as long ago as the time of Pliny,t and has attracted much attention in connexion with the 
more ordinaiy species of Acalephae ever since. The first observations of importance on this 
subject in modern times were those of the accurate Forskal, who described the phospho- 
rescence of the Pelagia and ^5'?/or^«, observed by him during his voyage to Egypt, in 1762. 
Since his time many voyagers and travelling naturalists, including Banks, Humboldt, Chamisso, 
Peron, Lesueur, SpLx, Mertens, and Baird, have published valuable observations on this 
interesting subject. On our own coasts, attention has been called to the subject, more 
especially by MaccuUoch and Macartney. The valuable observations of Suriray in France 
were chiefly confined to some minute animals allied to Medusae. In Germany, Tilesius and, 
above all, Ehrenberg have pubhshed important and original essays on this subject ; and in 
Italy, the experiments of the indeftitigable Spallanzani still furnish some of our best modem 
data. Many more authors might be cited who have treated the matter in greater or less detail, 

* James Montgomery, Pelican Island. Canto I. 
t Hist. Nat., lib. xxxii, c. 10. 



12 PHOSPHORESCENCE. 

and those of our readers who may wish for fuller information on the general question, would 
do well to consult the excellent essay of Brandt, in the 'Transactions of the Imperial Academy 
of St. Petersburgh' for 1838, and the article "Luminous Animals," in Dr. Todd's 'Cyclopfedia 
of Anatomy,' by Dr. Coldstream, a gentleman well versed in such inquiries. Several very 
interesting observations on the phosphorescence of the Cilograda* have also been published 
by authors, both British and foreign ; but these are foreign to my subject at present, as I 
wish to confine the discussion mainly to the naked-eyed Medusae, and intend on a future 
opportunity to treat of the Beroe tribe in full, having accumulated abundant new materials 
for an essay on the British Ciliogracla. 

Most of the accounts of the phosphorescence of Medusae have reference to the higher or 
steganopthalmatous species only ; nevertheless, at an early period the light given out by naked- 
eyed species attracted attention, for we find in the middle of the last century the phospho- 
rescence of one of them (" Medusa microscopica" — probably a young state of the Saphenia 
dinema of modern authors) attracted the attention of the observant Slabber. Previously, the 
phosphorescence of " Medusa cequorea" {^quorea ForsJicdina, Lamarck) was noticed by 
Forskal : " Rasa hgno, parum adeo in tenebris splendet."t Peron and Lesueur afterwards 
mentioned their JEquorea phosphorifcra as possessing the luminous power. Tilesius observed 
it in Charybdea marsupialis ; Rathke, in his " Oceania Blumenbachii."^ Macartney announced 
the Thaumantias hemi splicer ica to be an active cause of the luminosity of the sea on our own 
coasts, and detailed some interesting experiments which I shall presently have to cite. Among 
the twenty phosphorescent Medusae mentioned by MaccuUoch, it is probable that several were 
species of this division. One of the forms figured by Baird as a cause of phosphorescence in 
tropical seas§ appears to be a Sarsia. Ehrenberg observed this phenomenon in Oceania 
pileata, " Melicertum campannlatimf {Stomohrachium octocostatum), " Oceania micro- 
scojnca," and Thaumantias lenticula and hemisphcerica.\\ 

The British naked-eyed Medusae in which I have observed phosphorescence are species 
of Turris, Oceania, Diancea, and Thaumantias. Li no case have I seen it continuous or 
constant in any one species. La every instance the light has been given out only under 
circumstances of irritation, and not always even then. Thus, on the 27th of July, 1845, when 
in the Zetland Islands with Mr. M'Andrew, we collected myriads of small Medusae, and 
placing great numbers in a basin of sea-water, irritated them by many annoying devices, 
but though the vessels were charged with individuals of Thaumantias, Steenstrupia, 
and Lixxia, active and in good health, no light was given out ; nor could this arise from any 
peculiarity of conditions in the vessel or contained water, for individuals of the ciliograde 
Mnemia norvegica {Bolina hibernica of Patterson) gave out flashes of vivid bluish light, so 
as at times to illuminate the whole vessel. Tliis experiment was often repeated with the 
same result, though at the same season in the following year, the not giving out light was the 

* The essays of Mr. Robert Patterson, in the ' Transactions of the Royal Irish Academy,' should 
especially be consulted. I may here mention that such members of the Ray Society as have not 
attended much previously to the subject of this essay, could not prepare themselves better for a prac- 
tical study of it, than by consulting the admirable httle ' Zoology for Schools,' by the valued friend just 
mentioned. 

t Fauna Arabica, p. 110. J Philosophical Transactions, 1810. 

§ Loudon's Mag. Nat. Hist., vol. iii, 1830. || Berhn Transactions, 1832. 



PHOSPHORESCENCE. 13 

exception, and not the rule with similar Medusse, on the coast of Cornwall. Spallanzani 
fancied that the higher Medusa, Pelagia phosphorea, always emits light more or less, but his 
own experiments go far to disprove the notion. My friend and countryman Dr. I. Heywood 
Thompson, R.N., has, during his voyages on the western coast of Africa, and elsewhere in 
tropical and in southern seas, paid careful .attention to the phenomenon of phosphorescence of 
the sea, and has never observed it, when the product of animal life, to occur, except under 
circumstances of irritation.* 

Macartney narrates the following experiments, instituted on a species of Thaumantias, 
with the view of ascertaining the effects of various irritating conditions : — 

" 1. Some hemispherical Medusae were put into a spoon, containing a small quantity of 
sea-water, and held over a burning candle. As soon as the water became heated 
the Medusce appeared like illuminated wheels, the spots at the margin and centre 
alone emitting light, in which manner they shone vividly and permanently for about 
20 seconds, when they shrunk and died, after which they were no longer luminous. 
" 2. Some of the same species were put into spirits : a strong and unremitting light was 
instantly given out, which issued from the central and marginal parts, as in the 
preceding experiment, and continued until they died. 
" 3. Some of the scintillating and hemispherical species of Medusae, contained in a small 
glass jar, were introduced into the receiver of an air-pump, and the air being 
exhausted, they shone as usual when shaken ; if any difference could be perceived, 
the light was more easily excited, and continued longer in vacuum. 
" 4. A Medusa hcmisp1i(er\ca was placed in a small glass dish, containing a quantity of 
water, merely sufficient to allow the animal to preserve its figure ; being insulated, 
it was electrified and sparks drawn from it, which had not the slightest effect ; the 
experiment was repeated several times with different individuals, but without exciting 
the animals to throw out light. 
" 5. Some hemispherical Medusae were placed in contact with the two ends of an 
interrupted chain, and slight electric shocks passed through them. During the very 
moment of their receiving the shock no light was visible, but immediately 
afterwards the Medusae shone like illuminated wheels, which appearance remained 
for some seconds. Upon the closest inspection with a magnifying glass, no motion 
could be perceived to accompany the exhibition of light. The application of 
electricity in this instance seems to have acted as a strong mechanic shock. "f 
Ehrenberg placed the same species in spirits, with a view to observe the effect, and found 
the phosphorescence brilliantly revived when it had ceased to be exhibited by ordinary modes 
of irritation. 

Spallanzani had, previous to either, instituted similar experiments with those described 
by Macartney, but on one of the higher Medusce, the Pelagia only. He found raising the 
temperature revive the phosphorescence, when it had otherwise ceased to appear. 

I have found the sudden plunging of Thaumantias into fresh water or spirits call forth 
the dormant phosphorescence suddenly, and with extreme vividness. It gradually fades away 

* His notes will appear in the account of tlie Niger Expedition by Capt, Allen and Dr. Thompson, 
t PhU. Trans., 1810. 



14 DEVELOPMENT. 

as life departs. The same phenomenon is exhibited by the hydroid, heUanthoid, and asteroid 
polypes. If a bunch of one of the bushy corallines, such as Sertularia abietina, be plunged 
when alive and active into fresh water or spirits, a gorgeous display of living stars is instan- 
taneously produced. So also with Pennatulu jihospJtorea* 

The light of the Medusre, as Spallanzani observed, is not given out by all parts of the 
body indifferently, but only by certain structures. Spallanzani states that when he cut off 
the margins of the Pelagia fliosphorea to a depth of from 5 to 6'", the border continues to 
shine, which is not the case with the disk, and he attributes the phenomenon to the produc- 
tion of a phosphorizing mucus by the light-producing parts. Ehrenberg regards the light as 
an act of organic life. He observes that " the active organic phosphorescence appears 
frecpiently periodically, jDroduced either spontaneously, or by excitement, frequentl)^ as 
rapidly produced sparks, resembling small electric discharges. This repeated sparkling 
converts a mucous, gelatinous fluid, which is discharged more abundantly during the operation, 
into a secondary state of phosphorescence, which continues for a time, even after the death of 
the organism, or after the severing of its parts." He considers the mucus enveloping the 
ovaries as particularly susceptible, when in a fresh state, to this imparted phosphorescence. 
In Oceania pileata he observed the light emanate from the locality of the ovaries, which, 
being pendant in the centre of the sub-umbrella, illuminated the animal as an argand lamp 
illuminates its glass shade. In Thaumantias JiemisphcBrica he observed the light to be given 
out by the bulbous bases of the tentacula, w-hich formed a garland of sparks of fire around 
the circumference of the umbrella. Macartney had previously noticed, as we have seen, that 
in this naked-eyed species, the light was given out from the same spots, and, he adds, from 
the centre also. I have observed that in Thawnantias lucida the light was invariably given 
out by the bulbs of the tentacles, and so also in other species of the same genus. In the 
Bianea appendiculatu, which is a beautifully luminous species, the phosphorescence is of a 
greenish hue, and appears to radiate from the reproductive glands. In the Mediterranean I 
have seen a large Mesonema give out rich flashes of flame from the bases of its numerous 
marginal tentacles. Both Spallanzani and Tilesius have noticed that the light in the higher 
Medusce shone most vividly during the contractions of the umbrella. 

Taking one fact with another, it would seem that the phosphorescence in the naked-eyed 
Medusae is developed by the reproductive and motor systems : how, we cannot say. 
Ehrenberg has concluded that the production of the light is " a periodical vital act dependent 
on the nervous system, and similar to the development of electricity." But this can only be 
regarded as an hypothesis. We have no clear evidence yet of the presence of a nervous 
system in these animals. 

Development. — When treating of the reproductive organs of these animals, I discussed 
the ovarian and spermatic glands; but the naked-eyed Medusae do not reproduce their 
species only in the normal fashion, i. e. by fecundated eggs ; several of them are now known 
to multiply their kind by gemmation, little ones springing out almost ready made from the 
substance of their parents, as Minerva budded on the creative brain of Jupiter. This mode 

* See a notice of the plienomena exhibited by this zoophyte when phosphorescing, in the second 
edition of Dr. Johnston's British Zoophytes. 



DEVELOPMENT. 15 

of propagation by gemmation was long supposed among Radiata to be an especial privilege 
and distinction of the true zoophyte ; but the march of discovery and the revolutions of 
science, do away with such artificial distinctions, though the recognition of them in their time 
gave no small impulse to the onward progress which was eventually to destroy them. The 
discoveries of Sars, Dalyell, Loven, Wagner, Van Beneden, Dujardin, and Steenstrup have 
changed the face of this section of creation seriously, and prophetically indicate many coming 
changes. It is the duty of the philosophical zoologist to keep pace with the railroad sjieed 
of modern research, and whilst conservative of all past statements, as yet insufficiently 
combated, never to hesitate to cast away preconceived notions and old teachings the moment 
they are clearly shown to be untrue. " Free and unprejudiced spirits will neither antiquate 
truth for the oldncsse of the notion, nor slight her for looking young, or bearing the face of 
novelty."* 

The polypes of the genus Coryne and its allies, of Tuhdaria and Eiidendrium, and of 
the beautiful Corymorpha, send forth at certain times bud-like bodies, more or less sym- 
metrically arranged around their heads. These bodies have long been recognised as 
young animals, though not until very recently was it known that the creatures so produced 
bear no resemblance to their parents, but were indeed true Medusa, and not polypes, which, 
however, in their turn produced eggs capable of producing polypes. Such appears to be the 
true interpretation of the phenomenon — a part of the justly celebrated theory of alternation of 
generations which has originated in the imaginative mind of Steenstrup. On the bearing of 
such discoveries on the better classification of the Radiata, I shall have to make a few remarks 
presently. Now, among the Acaleplue, no such reproduction by gemmation in the manner 
of the Coryne was known until discovered in 1 836 by Sars, who had previously been the 
first to announce the surprising fact of the intermediate Strohila condition of the higher 
Medusae, a discovery made independently and simultaneously in Scotland by Sir John Graham 
Dalyell. 

The discovery made by Sars was that certain forms of naked-eyed Medusae multiply 
their species by means of gemmation, the buds being produced either from the walls of the 
peduncle or stomachal prol:)oscis, or from the surface of the ovaries, the former mode occurring 
in the " Cytm'is octoimnctatd' {Li%%ia octopunctata, Mihi), the latter in Thamnantias midti- 
cirrata. In both cases, the new individuals were not different from, but similar to, their 
parents, and, in one instance, provision seemed to be already made in the new-formed individuals 
for continuing to propagate by the same mode other individuals similar to itself. The full 
account of these remarkable and highly important observations, illustrated by excellent figures, 
is contained in the lately published ' Fauna Norvegica,' a most admirable work, by one of the 
greatest of li^^ng investigators of life in the sea. I shall e.xtract it hereafter when describing 
the Lix%ia, in the second part of this essa)'. At present I shall cite the summing up given by 
Sars, and his notice of the phenomenon as it occurs in the Thaumantias. 

" We now recognise a mode of procreation and development hitherto unknown among the 
Acalephae. From a certain part of the body (in this instance the tubular stomach, hanging 
independently in the cavity of the disk) romidish knobs grow forth from the upper towards 
the lower part, which gradually assume the shape of a bell, by opening themselves at the free 

* Henrv More. 



16 DEVELOPMENT. 

end ; on tlie margin of these openings dark granules (marginal granules) make their appearance, 
being the nuclei or first beginnings of the marginal fibres, which latter gradually grow forth, 
and the stomach exhibits itself at the base of the cavity of the bell-formed disk, together 
with the mouth and mouth-tentacle ; from the stomach vessels radiate towards the margin of 
the disk ; — in short, the young Acalephis, being merely attached to the mother by means of 
a short peduncle issuing from the back of the disk, develops itself in all essential organs, 
whilst it is still attached to the mother like the bud of a plant. Finally, after a certain 
expiration of time, it severs itself from the mother, and now swims about as an independent 
individual. 

"I likewise found quite the same mode of perpetuation, on the 9th of May, 1837, in 
Tliaumuntias multicirrata, in an Acalephis of more than one inch in diameter. In the 
four narrow, folded together, so-called ovaries that issue from the stomach, and extend along 
the margin of the disk, there were blossoming forth, even as described in Cytms, some 
globular bell-shaped gemmules (I observed from five to six towards the external extremity of 
the ovarium), the smallest of them furnished with four, the largest with eight black marginal 
granules, and short marginal fibres growing forth from the former. Perpetuation by means 
of prolification has hitherto been chiefly observed among the poly|3es, in which indeed it is the 
prevalent mode, but it was latterly also observed among the Infusoria (Vorticelles), the Tunicata 
(the compound Ascidians), and, lastly, also in some of the Annelides (the Naicles and Syllis 
prolifera, to which I may add the FUograna hnplexci). We now likewise recognise this 
mode of perpetuation in an animal which will undoubtedly be declared by all classifiers to be 
an Acalephis, against the assertion of Ehrenberg, that ' a bud-bearing or self-dividing 
Acalephis is a contradlctio in adjecto' Thus are our speculations and inferences not 
unfrequently frustrated by a boundless and ever-varying nature." 

I am not aware of an)^ naturalists ha\'ing confirmed the observations of Sars. With very 
great pleasure, therefore, am I able not only to bear out, by personal observation, the remark- 
able statements of the Norwegian naturalist with respect to the species he mentions, but also 
to extend them to other species and other genera. 

I have observed four modes of propagation by gemmation among the naked-eyed 
Medusae. 1st. Gemmation from the ovaries, as noticed by Sars in Thawnantias imdticirrata, 
and which I have seen, though not in an advanced stage, in my Thmimantias lucida, a nearly 
allied species. 2d. Gemmation sub-symmetrically from the peduncular stomach, as described 
by Sars in his accomit of Lixxia octojmnctata. This I have seen in all its stages, exactly as 
he narrates, in the same species, a very abundant animal in the Zetland seas. I call it sub- 
symmetrical gemmation, for whilst the four gemmae are symmetrically arranged around the 
peduncle, one of them is constantly in a more advanced condition of development than the 
other three. This appears to be a generic habit, for I find the same phenomenon in Z/zzia 
hlondina, a new and very distinct species of the same genus, in which gemmiparous repro- 
duction is equally conspicuous. 3d. Gemmation irregularly from the walls of a tubular 
proboscis. This I have discovered in a new Sars'ia, which I have named Sarsia 'prollferu. 
From the sides of the long peduncle many gemmules are seen springing in all states of develop- 
ment, and presenting an indistinct spiral arrangement. There is no order of development 
with respect to position, individuals variously advanced springing indifferently from various 
parts of the peduncle. [See the account of the species in the Second Part.] The fourth mode of 



GENERA. 17 

gemmation, and a very remarkable and quite novel one, I have discovered in another form of 
Sarsia, taken abundantly in 1 836 on the coast of Cornwall, and named by me Sarsia prollfera. 
In this extraordinary animal the buds are produced at the bases or tubercles of the four marginal 
tentacles, and hang from them in bunches, like grapes. The degree of development is not 
equal in all four bunches, and in each case buds are seen in very various stages of develop- 
ment, from embryo wart-like sproutings to miniature Medusae, simulating in their essential 
characters the parent animal. [See figures and description of Sarsia prolifera in the synopsis 
of the species.] 

I look upon this last discovery as very important in its bearing on the history of this 
phenomenon of gemmation among these creatures, seeing that in the case under consideration the 
seat of reproduction is not in the peduncle, where in Sarsia the true ovaries might be supposed 
to be seated, since they are not manifest elsewhere, but in a portion of the animal quite apart. 
It would appear, indeed, that gemmation can occur anywhere in the course of the granular 
motor tissue, or from the true ovaries, but not from other than the motor, and especially 
reproductive, tissues. The power of the motor tissue to produce germs occurs also in other 
orders of Medusae : for I have seen the same phenomenon in the Beroe. When treating of 
the phosphorescence of the naked-eyed Medusae, we have seen that that phenomenon was 
exhibited either by the ovaries, or centre (peduncle) or bases of the tentacles and rim ; all, as 
I have just proved, seats of reproductive power. This goes far to connect that phenomenon 
with the generative functions. 

The development of the oviim in the naked-eyed Medusae has still to be observed. Not 
until the phenomena attending it have been made known, can we hope to ascertain the history 
of the metamorphoses which they possibly, indeed most probably, undergo before arriving at 
the state in which we visually find them. 

The preceding outline of the structure and physiology of the naked-eyed Medusae, so far 
as known, will enable the reader to understand the detailed account of our British genera and 
species, which I shall now proceed to give, reserving remarks on their zoological affinities as 
a group, for a review of the subject at the conclusion of the synopsis, when we shall have 
the necessary evidence before us. 

There are forty-three species of Gymnopthalmatous Pulmograda known to the author 
as inhabiting the British seas. The greater number of these are undescribed forms. They 
may all be arranged under eighteen Genera, grouped together as in the following table : 

I. Vessels branched. (Willsiad^e.) 

1 . Willsia. 

II. Vessels simple; ovaries convoluted, and lining the pedunculated stomach. 

(OCEANrDiG.) 

2. Turris. 

3. Saphenia. 

4. Oceania. 

III. Vessels simple, eight or more ; ovaries linear, in the course of the vessels on the 

sub-umbrella. (/EquoreaDjE.) 

5. StomohracJiium. 

6. Polyxenia. 

3 



18 GENERA. 

IV. Vessels simple, eight ; ovaries as many as the vessels, small, on the course of the 

siib-umbrella. (Circead^.) 

7. Circe. 

V. Vessels simple, four ; ovaries four, in the course of the vessels on the sub-umbrella. 

(GERYONIAD.ffi.) 

8. Geryo7iia. 

9. Tim a. 

1 0. Geryonopsis. 

1 1 . Thaumantias. 

12. Slabber ia. 

VI. Vessels simple, four ; ovaries in the substance of the peduncle. (Sarsiadj!.) 

13. Sarsia. 

14. Bougainvillea. 

15. Li%%ia. 

16. Modeeria. 

17. Eupliysa. 

18. Steenstrupiu. 



WILLSIADiE. 

Genus I. Willsia, Forbes (1846). 

Char. Gen. Umbrella globose ; ovaries six, radiating around the base of the 
short, campanuiate, four-lipped stomach ; vessels six, twice dichotomously dividing 
before they reach the marginal vessel; a marginal tentacle opposite each branch: 
ocelli conspicuous. 

Willsia stellata, Forbes. 
(Plate I, Fig. 1.) 

I have constituted this genus for one of the most elegant of our naked-eyed Medusas, 
strikingly distinct in its characters from all recorded forms, and quite new. I have dedicated 
it to my friend Dr. Will, of Erlangen, whose work on the ' Medusas of the Adriatic,' ah-eady 
often quoted, is one of the most valuable and original contributions to this department of 
zoology ever printed. 

This beautiful little creature is, when full grown, about a quarter of an inch in diameter. 
The umbrella is nearly globular, quite smooth, and colourless. The sub-umbrella is small in 
comparison, and hemispheric, or slightly cyhndrical in form. The margin bears twenty-four 
extensile, pale yellowish tentacula, of a minutely granular tissue, and moniliform structure. 
At the base of each is a bulb or ocellus, of a deep purple red and tawny yellow colour, the 
darker hues being disposed in an arrangement partly crescentic, and partly eye-like. When 
highly magnified, the coloured parts are seen to include a cavity containing a central body, 
which, though not obsei-ved to vibrate, is probably the homologue of an otolitic mass. Some 
specimens were observed which had only twenty tentacula. 

The central peduncle, or stomach, is campanuiate, and opens widely by four scarcely 
undulated lips.* Sometimes the orifice presents a six-lipped appearance, and it is very pro- 
bable that it may contract itself indifferently into four or six divisions. Around the base of 
the stomach, partly attached to its side, and partly to the sub-umbrella, are the six double 
ovaries, or reproductive bodies, each of an oblong, somewhat spoke-like shape, and coloured 
of a tawny yellow. The colour is due to two fulvous masses on each side the gastro-vascular 
canal, one of which traverses the centre of each ovary. The fulvous masses are bordered by a 

* In tlie fig-ure there is an appearance as if of a fifth lip, \iliith is a mistake. 



20 WILLSIA STELLATA. 

nearly colourless margin. The six ovaries together form a beautiful star around the base of 
the stomach. Each gastro-vascular canal runs without dividing through the corresponding 
ovary, but when it reaches half way down the sub-umbrella, it divides into two, and at a 
fourth of the distance between the margin and the centre, each branch again divides into two, 
which is the final division, for these last two branches run directly into the marginal canal, 
each opposite the origin of a tentacula. 

The Willsia stellata was first observed by Mr. M'Andrew and myself, in September, 
1845, in the Bay of Oban. Many specimens occurred, and its appearance in the vessel of sea- 
water, was very striking, the star-like ovaries conspicuously distinguishing it from Medusae 
of other genera. It is so transparent, that usually only the reproductive star and the mar- 
ginal circle of brilliant ocelli — like a mimic sun with its surrounding planets — could be 
perceived by the naked eye. When placed in a watch-glass, however, the singidar arrange- 
ment of its vessels, and the other details of its structure, may easily be made out vnthout the 
use of a high magnifying power. 

In August, 1836, when visiting Penzance Bay, we took great numbers of the same 
species, so that it would appear to be rather widely distributed. It was not observed at 
Zetland, nor further west than the Lizard on the south coast of England. 

In the Plate, fig. 1 represents the natural size ; 1 , «, a magnified view as seen from the 
side; 1, h, the creature seen from above; 1, c, the mode of division of one of the gastro- 
vascular trunks ; \,(l, an ovary ; and 1, e, a tentacle, both greatly magnified. 



OCEANIDvE. 

Genus II. Turris, Lesson (1837). 
CoNis, Brandt (1838).? 

Umbrella sub-cylindrical or mitrate ; ovaries four, double, dense, convoluted, 
lining the cavity of the peduncle ; vessels simple ; margin of the umbrella with 
numerous tentacula ; muscles of the disk conspicuous and highly developed ; mouth 
of the peduncle fimbriated. 

This well-marked genus was constituted by Lesson for three species of naked-eyed 
Medusae, having an organization which seemed to indicate a high position in their order, 
though not so high as the founder of the generic group would seem to place it. It forms part 
of the tribe Niicleiferre in the arrangement of Lesson, where it is strangely associated with 
Pandea, Circe, and Bougainvillea. The high development of the muscular system in the 
species of Turris, indicate their superiority to the Oceanim proper, which otherwise they 
closely resemble. The only species known besides the following, is the Turris papua of the 
author cited, who discovered it near the island of Waigiou. 

The T. papua has only eight tentacles, whilst our native species are remarkable for the 
number of those organs fringing their margin. This variation in number of tentacles seems 
a good source of specific distinction in the genus. 

1. Turris digitalis (sp.), Miiller. 

Plate in, Fig. I. 

Synonyms. Medusa digitate. O. F. Muller, Prod. Z. D., p. 233, No. 2824 (1766). 

O. Fabricius, Fauna Groenlandica, p. 366 (1780). 
MeUcerta digitale. Peron, Ann. du Mus., t. .\iv, p. 352 (1809). 
Dianea digitale. Lamarck, Ann. Sans. Vert. (1817). 
Eirene digitale. Eschscholtz, Syst., p. 95 (1829). 
Turris horealis. Lesson, Prod. (1837), and Hist., p. 284 (1843). 

This beautiful species is an inhabitant of the northern seas. It was observed on the 
coast of Greenland by Otho Fabricius, who, in his ' Fauna Groenlandica,' one of the most 
remarkable zoological works of the last century, has given a short but expressive description 
of it. " Hsec omnium minima digitale tam figura, quam magnitudine refert. Corpus conicum 
hyalinum, vix in aqua observaretur, si non motu ejus margo coloratus in conspectum veniret. 
Striae multse vi.\ notabiles longitudinal iter in verticem concurrunt. Margo ciliatus est ciliis 



2-2 TURRIS DIGITALIS. 

intus liamatis flavis cum albis mixtis. In cavitate infera, quse profunda, pistillum apice, ut 
videtur penicillato dependet, in aliis album, in aliis penicillo flavo. Habitat in mari ad oras 
exteriores. Est vivida satis ; margine flexo in aqua salit." 

This account of its characters and habit seems to have furnished all subsequent authors 
with the brief notices which they give of the species, indeed it is doubtful whether it has 
been examined by any since the days of the author of the 'Fauna Groenlandica.' Miiller probably 
intended to give a representation of it in his invaluable ' Fauna Danica,' but, as it is, we have 
no figure extant. 

When enjoying a delightful cruise with my friend Mr. Smith, of Jordanhill, in his yacht 
the Amethyst, during the summer of 1839, I paid particular attention to the Medusae of the 
Clyde, and laid the foundation of the present monograph. Not so well versed then, as now, 
in the art of securing these fragile and floating creatures, it was a source of not unfrequent 
vexation to behold many of them, either apparently new or doubtful, pass by our vessel beyond 
our reach, or when we endeavoured to secure them, sinking slowly in the sea depths. Among 
those which we failed altogether in catching, was one of considerable size — three inches or so 
in length — and conspicuously distinguished from all others of my acquaintance by its singular 
cylindrical umbrella, and the dense, brilliantly-coloured nucleus. A rude and rapid memo- 
randum of its aspect in the water, as seen over the vessel's side, was all the record which we 
could bring away. 

In the autumn of 1845, when on a voyage of research with Mr. M'Andrew, in his yacht 
the Osprey, we procured numerous Medusae in the sound of Brassay, among the Zetland Isles. 
It was with no small pleasure that, on emptying the tow-net one morning in August, I 
found in it what at first glance appeared to be a floating Actinea, but which, on closer 
examination, was evidently identical with the creature I had seen in the Frith of Clyde six 
years before, and which I had often anxiously looked for since, but in vain. A further 
inspection, and a comparison of it with published descriptions, soon con^^nced me that we 
had secured the finger-shaped Medusa, so pithily described by Otho Fabricius. 

The umbrella is sub-cylindrical and mitrate, swelling gently into a bell-shape in the 
centre, somewhat apiculated at the apex. Its substance is highly transparent and colourless, 
but firm. Its margin is fringed by fifty or more long annulated, and as if granulated, ten- 
tacula, which are white, with orange bases, and when contracted, are curled or rather hooked 
at their extremities. The bases of the marginal tentacles are covered by an external veil-like 
prolongation of the margins. The sub-umbrella occupies slightly more than two thirds of 
the length of the body, and is cylindrical in form. Along its sides are seen to run eight 
conspicuous longitudinal bands of muscular tissue, which have a furbelowed appearance, in 
consequence of habitually contracting at fixed intervals. The substance of these bands is 
composed of distinct fibres. From the centre of the sub-un;brella depends a capacious and 
cyhndrical peduncle, the whole of whose interior is occupied by the stomach and ovaries, or 
reproductive glands. The latter bodies consist of four quadrate groups of foliated and convo- 
luted masses, of a bright reddish-brown or brownish-red hue, arranged in the course of four 
simple gastric vessels. Each of these masses consists of a double series of fimbriated organs, each 
series being placed on one side of a vessel. Below the ovaries the peduncle contracts, though 
not greatly, and expands again to form the much divided fimbriated lips, which fringe and border 
the wide oral orifice, and which are parted into four somewhat indistinct lobes. The lips are 



TURRIS NEGLECTA. 23 

of a fine rose colour. They reach to a level with the margin of the umlirella. The specimen 
I have described was a little more than an inch in length, exclusive of its tentacula. It was 
sluggish, possibly owing to fatigue, as it had probably been in the tow-net during a greater 
part of the night. I have represented it in Plate III, fig. 1, a, of the natural size ; fig. 1, b, is an 
outline showing the arrangement of the muscular bands ; fig. 1 , c, represents a portion of the 
peduncle laid open, with two of the ovarian masses, and the fimbriated lips, also a part 
of the border of the umbrella with its tentacula ; fig. 1, d, is a portion of one of the tentacles, 
magnified ; and fig. 1, ^, a portion of the tissue of one of the muscular bands, as seen under a 
high power. 

Eschscholtz has strangely placed this Medusa as an associate of Diantsa viridula, in his 
genus Eirene. 

2. Tunis neglecta, Lesson. 

Plate III, Fig. 2. 

Synonyms. Turris neglecta. Lesson, Prod., No. 38 (1837), and Acal. Hist., 
p. 284 (1843). 
CyancBu cocchiea. Davis, Ann. Nat. Hist., t. vii, p. 234, pi. 2, f. 

12, 13(1841). 
Carm'mrothen beroe. Slabber, Ph. Vet., p. 59, t. xiii.f. 3? (1781). 
Oceania sungidnolenta. Peron, Ann. de Mus., p. 347? (1809). 
Oceania tetranema. Peron, Ann. de Mus., loc. cit. ? (1809). 

A beautiful little species, which, when in its native element, is brilliant as a bead of 
brightest coral. It appears to be not uncommon in the Solent and around the Isle of Wight. I 
first caught it on the south coast of that island in 1844, when geologising there with my friend 
Captain Ibbetson. Since then it was taken in the west bay of Portland, just before the 
Southampton meeting of the British Association, by Mr. M'Andrew and myself, and again off 
the mouth of Southampton harbour during the week of the meeting, in a memorable day's 
cruising, when a small band of British naturalists fraternised with Agassiz and Middendorf, 
and enjoyed themselves as true students of Nature only know how, when " dredging the 
waters under." 

The umbrella is transparent, smooth, and sub-hemispheric, inclining to conical. Its 
summit is slightly pointed. The sub-umbrella is small in proportion, reaching to a little more 
than half the height of the former. It is slightly pyramidal, with a truncated summit. The 
muscular bands are distinctly seen striping its sides. The peduncle is large, and has a 
singularly substantial aspect, in consequence of the compact masses of rich crimson or vermilion 
convoluted and fimbriated ovaries which occupy its broader and upper half. This bright red 
nucleus causes the animal, small as it is, for it reaches scarcely more than one fourth of an 
inch in height, to be very conspicuous in the water. The substance of the lining of the sub- 
umbrella is also very firm. The brilliant colouring of the ovaries is due to the presence of 
large red ova. I met with individuals, which at first sight seemed as if belonging to a different 
species, in which the reproductive glands were dull pink. These may have been males. 
The peduncle terminates in four lips, which are fimbriated at their edges, and highly muscular. 



24 TURRIS NEGLECTA. 

The radiating vessels are four, and all simple. The margin of the umbrella is studded 
with a close-set circle of tentacula, more than sixty in number (16x4+4), very contractile, 
being sometimes elongated into hair-like filaments, and at others contracted into little knobs. 
Each tentacle has a large bulbous base, the upper part of which bears a brilliant ocellus, 
consisting of crimson pigment-cells, and there is another speck, probably the site of an otolitic 
mass below. 

A very interesting account of this species, under the name of Cyanaa coccinea, was given 
by Dr. J. F. Davis, in the seventh volume of the 'Annals of Natural History.' " Amongst 
the variety of animals," writes that gentleman, " which we had opportunities of seeing, 
during our stay at this charming marine watering-place (Tenby), none afforded greater interest 
than a small Medusa belonging to the genus Cyanma, Cuvier Having been dis- 
covered by Mrs. Davis, who had likewise the best opportunity of watching its motions during 
several weeks that she kept it in a glass of sea-water at Tenby, and afterwards here (Bath), 
whither it was conveyed in a phial of the same, and lived three weeks after its arrival, I will 
state the history of this ' thing of light and life' in her own words. ' One morning, while 
pouring some sea- water into the vessel containing my Actinice, I observed two small objects, 
which I took for the young of these animals, and as quickly as possible raised them in a spoon 
out of the basin, and placed them in a tumbler of clean sea-water. They resembled tiny bell- 
glasses. Four transverse rays were perceptible on their sides, and a minute red body, with 
four white arms forming a cross, was suspended in the water. Around the edge of the bell 
or disk appeared a delicate white fringe, which was lengthened or shortened at the pleasure 
of the animal. The contraction was sometimes so great, as to give the fringe the appearance 
of being knotted up to the edge of the bell or disk. It was highly interesting to watch their 
movements in the water as they ascended from the bottom, the bell or disk conti'acting and 
dilating alternately, until the animal arrived near the surface of the water. This motion was 
particularly conspicuous at the edge of the disk, and the fringe or tentacula became shortened 
as the animal rose in the water ; but when they descended again the tentacula lengthened, 
sometimes to a great degree, after which the animals sunk gradually, and without any visible 
effort. At the end of a fortnight one of my pets turned itself inside outwards, and remained 
in this state for some time, when it died, and left only a few flocculent particles at the bottom 
of the vessel. The other lived more than two months longer, and even bore a voyage to Bath 
in a closed phial of sea-water, and remained active and vigorous during the space of three 
weeks, when it likewise shrunk, died, and disappeared like the former, but without the previous 
eversion.' " 

Plate in, fig. 1, a, Turris neglecta, much magnified ; 1, b, body seen from above, showing 
the muscular bands alternating with the vessels; 1, c, proboscis and lips ; \,cl, one of the 
lips magnified ; \, e, a. group of tentacles ; 1, g, a tentacle with its bulb, showing the ocellus 
and otolitic mass ; 1, h, portion of a tentacle highly magnified ; 1, ?', ova ; 1, A., muscular 
tissue. 



SAPHENIA DINEMA. 25 



Genus III. Saphenia, Eschscholtz (1839). 

Umbrella sub-cylindrical, campanulate, or hemispheric; ovaries four, double, 
convoluted, lining the upper part of the cavity of a highly extensile peduncle ; 
vessels simple, four ; margin of the umbrella with two large, and numerous very 
small tentacula ; mouth of the peduncle four-lipped. , 

I have no doubt whatsoever that the Medusa I am about to describe, is one of those 
regarded by Eschscholtz as characteristic of his genus Saphenia, although his definition, 
" Peduncidus apice simplex," does not apply. He founded his genus on figures and imperfect 
descriptions ; but, as it seems to be a good group wrongly defined, I have adopted it, and 
revised the character. 

1. Saphenia dinema (sp.), Peron (1809). 

Plate II, Fig. 4. 

Synonyms. Geryonia dinema. Peron, Ann. de Mus., p. 346 (1809). 
Dianaa dinema, Lamarck, First Ed., p. 505 (1817). 
Campanella dinema. De Blainville, Man. d'Act., p. 286 (1834). 
Saphenia dinema. Eschscholtz, Syst., p. 93 (1839). 

I have met with this species in our seas during two seasons, once near Hillswick, on the 
western coast of Zetland (1845), and thrice on the south coast of England, in Cornwall and at 
Portland, in 1846. It is small, being scarcely more than one fourth of an inch in length, but 
conspicuous, owing to the colour of its ovaries and tentacles. The umbrella is cylindrical, 
somewhat mitrate, obtuse (acute in young specimens), and smooth. It is quite transparent and 
colourless. From the margin spring two large orange purplish tentacles, with bulbous and 
rather large bases ; in the interspaces, and covered by a short veil, are a number of minute 
and very short colourless tentacles, about twenty-four in all. These appear to increase with 
age, as young specimens had fewer. The sub-umbrella is about two thirds the length of the 
body. From its centre depends the highly contractile peduncle, which is sometimes drawn 
out into a long tube, sometimes contracted into a short bell. In its uppermost part are lodged 
the four reproductive glands, short, convoluted, double, distinct, of a bright purple colour 
when adult, and when immature, tawny. The part of the peduncle below the ovaries is most 
contractile, and is of a pale fawn colour, witii four dark purple lines in the course of the 
ovaries proceeding to the four slightly fimbriated lips bordering the orifice, and which, when 
drawn together, give the club-like appearance that appears to have misled Eschscholtz and 
other observers. 

Plate II, fig. 4, a and b, represent this species (enlarged), as seen from the side and from 
above ; 4, c, is the appearance of the peduncle when contracted. 



26 OCEANIA. 

Genus IV. Oceania (nom. Peron), (1809). 

Tiara and Oceania (pars). Lesson. 

Umbrella conical, mitrale, or rarely globular; ovaries four, double, convoluted, 
lining the cavity of the peduncle; vessels simple, four; margin of the umbrella with 
few, or many, similar tentacula ; no conspicuous muscular bands; mouth of the 
peduncle four-lipped. 

The natural wish to preserve a name in a manner consecrated by long usage, has induced 
me to retain that of Oceania, even at the risk of employing it in a somewhat different generic 
signification from any of the many meanings hitherto given to it, and thereby perhaps almost 
increasing confusion. In such a case it becomes a toss-up whether the conservative or the 
destructive process be best ; but as even the most virulent of reformers gladly seize on an 
excuse for upsetting usurpers, whilst good-natured people are inclined to stand by ancient 
dynasties, however bad, rather than risk the chances of change, I tliink it safest (though 
prepared to do either) to adopt Oceania as a genus for the present, and, by defining it more 
strictly, endeavour to make it more useful. 

The term Oceania has been so often and generally applied to the Medusa pileata of 
Forskal. and similar forms, that I think it best to restrict it to that group. Peron, who first 
founded the genus, included them within it, though it is doubtful whether he would have 
regarded the Forskalian species as the type. Lesson gives the generic name of Tiara to 
M. pileata ; and those who are extremely anxious to use a new name may adopt it. Eschscholtz 
included its allies in his view of the genus, but also some very distinct forms. 

Using it in the sense here taken, the genus consists of those mostly mitre-shaped 
Medusae, which have an ample dependent peduncle and convoluted ovaries within, a conspicuous 
character indicating a close relationship with Turris. Their muscular tissue, however, is not 
nearly so highly organized as in the last-named genus, whilst the uniform character of the 
tentacles separates them from Saphcnia. 

The species of Oceania range throughout the European seas, extending into the 
Mediterranean. They will probably be found much more numerous than at present recorded. 
Sars has described and figured in his ' Beskrivelser,' &c., 1835, a species with twenty-four 
tentacula, and resembling in form our 0. episcopalis, which may probably inhabit our seas. 
He names it O. ampullacea, and describes it as ovato-campanulate, terminating above in an 
oblong, conical appendage ; the mouth furnished with very short fimbriations ; the length of 
the body about an inch. Forskal's name of '^ Medusa pileata" has been applied to more than 
one species of the genus. He described it as " ovato-campanulate, terminated above by a 
hyaline sphere ; within containing an oblong, red nucleus ; the margin fringed with numerous 
tentacula, having yellow bases." His figure does not show clearly the number of tentacula, 
but they appear to have been sixteen and upwards. The animal figured by Ehrenberg as 
Medusa pileata from the Norwegian seas, has a yellow nucleus, and appears to be the 
Oceania ampidlacea. The colour of the nucleus, the general form of the umbrella, and the 
number of the tentacula, are evidently the most important sources of character in this genus, 
and should be carefully noted. 



OCEANIA EPISCOPALIS. 27 

1. Oceania octona (sp.), Fleming. 

Plate II, Fig. 3. 

Synonym. Geryonia octona. Fleming, Edin. Phil. Journ., vol. viii, p. 299 (1822). 

British Animals, p. 501 (1828). 

Umbrella smooth, transparent, mitrate, somewhat constricted above the centre, and 
produced into a conical acuminated apex. Margin with eight elongated yellow tentacuhi, 
springing from thick bulbous bases. On each bulb is a minute red ocellus, and an otolitic 
cavity beneath, with an included vibrating mass. Between each pair of tentacles are three 
yellow minute tubercles springing from a narrow, yellow, marginal ring. On each tubercle is 
a minute red ocellus. The central one is largest. Down the sides of the sub-umbrella, which 
occupies about two thirds of the body, run four wide vessels to join a wide marginal vessel. 
The upper part of the sub-umbrella has often a lobed appearance. From its centre depends 
an ample, yellow, vasiform peduncle, including in its upper part four convoluted bright yellow 
ovaries. Its orifice is wide, and bordered by four fimbriated yellow lips. I have taken this 
species off the mouth of the Frith of Forth, where it has also been observed by my friend 
Mr. Henry Goodsir, and in the seas near the east coast of Zetland. 

It was first noticed and described by Dr. Fleming, who observed it in the sea of the east 
coast of Scotland in 1821. In his account of an excursion made that year, published under 
the title of Gleanings of Natural History, in the eighth volume of the ' Edinburgh Philo- 
sophical Journal,' he describes it as follows : " Having returned from the Bell rock to the 
vessel, I devoted some time to the examination of the molluscous cargo which I had brought 
on board. While observing the motions of some of the animals in a glass of sea-water, a 
Medusa presented itself belonging to the genus Geryonia of Peron and Lesueur. The body 
was diaphanous, round at the margin, sub-conical at the summit, and slightly acuminated. 
The central mouth was trumpet-shaped, and shortly pedunculated. The circumference of the 
body was furnished with eight similar tentacula, equal to its diameter. As it differs from 
Geryonia dinema and Geryonia proboscidalis, the only known species, I have named it 
G. octona.'' (Loc. cit. p. 298.) 

2. Oceania episcopalis, Forbes. 

Plate II, Fig. 1. 

The great fishing-banks which stretch along the coasts of the Zetland Isles, whether 
eastern or western, are among the most interesting of stations in the British seas for marine 
researches. The beautiful Medusa which I have now to describe was taken in the neighbour- 
hood of the western line of bank, forty miles from the mainland of Zetland, in the autumn of 
1845. When lying to there, as much in open ocean as if we had been in the very middle of the 
Atlantic, — the sea calm, though the hea\'y swell tossed our little vessel to and fro with a motion 
that would make land-loving naturalists wish themselves on shore, and vow never more to meddle 
with the wonders of the deep, but keep steadily at their studies among cockchafers and 
tom-tits, — we were delighted by the sight of shoals of swimming gelatinous animals, with 
brilliant purple nuclei, passing in succession near the surface of the water, and having all the 



28 OCEANIA TURRITA. 

aspect of Scilpce. On securing some, however, by means of the hand-net, they proved not 
to be molhisks but Medusse, alHed to, but very distinct from, the Geryonia octona of Dr. 
Fleming. Mr. Patterson appears to have met with a species very similar, if not identical, at 
Bangor, County Down. 

The largest of the individuals taken measured an inch and a half long in the body : its 
marginal tentacula far exceeding that dimension. The umbrella is very transparent, smooth, 
and mitre-shaped, its upper part more or less tumid, and at times assuming a sub-globular 
form, as if the body was crowned by a mobile glass ball. Round the margin are twelve 
(2X4+4) highly contractile tentacula, with thick bases. These tentacles are of a pale 
yellowish hue ; between each pair of them are three minute yellowish tubercles, the central 
one slightly the largest. All these tubercles and all the bulbous bases of the tentacles bear 
very minute red ocelli (12x3-|-12:^48). Round the orifice of the umbrella is a veil, borne 
upwards and inwards. The sub-umbrella is conico-cylindrical, and occupies about two thirds 
of the length of the body. From its centre is suspended an ample urn-shaped peduncle, 
including in its upper and most tumid part the eight convoluted ovaries, dense, and of a rich 
maroon purple colour. Four very wide gastric vessels run in the course of the reproductive 
glands, and proceed down the sub-umbrella to join the wide marginal vessel. The peduncle 
terminates in a campanulate proboscis, with a wide orifice, surrounded by four fimbriated lips 
of a pale purplish-tawny hue. 

It is a most active and graceful, but very delicate creature. 

Plate II, fig. 1, a, represents an individual slightly enlarged ; \,h, shows the disposition 
of the ocellated tubercles between the tentacula. 

3. Oceania turrita, Forbes. 
Plate II, Fig. 2. 

Umbrella campanulate, smooth, transparent, produced above into a long, conical, acute, 
and mobile process, turned more or less to one side. Margin with four long yellow tentacles, 
their bases much swollen ; on the bulb of each is a very minute crimson ocellus, composed of 
a well-defined group of pigment-cells, and in the substance of the bulb below it is a cavity 
containing a vibrating mass of crystalline particles (calcareous ?), mixed with brown pigment- 
cells. This, no doubt, is the otolitic body. Between each pair of tentacles are three yellowish 
marginal tubercles, with rudimentary ocelli. The sub-umbrella occupies about half the length 
of the body. Down its sides run four broad vessels to join a wide marginal vessel. From 
its centre depends a rather short, but wide yellow peduncle, in the upper part of which are 
four convoluted bright yellow reproductive glands. The orifice of the peduncle is cam- 
panulate, and bordered by four slightly fimbriated lips. The body measures about half an 
inch in length. 

When I first caught this singular and active little Medusa, I fancied we had secured the 
" Piliscelotus mtreiis" of Templeton. That anomalous animal, however, I now believe to 
have been merely a Sarsia tuhulosa turned inside out. The Oceafiia turrita was taken in 
the Zetland seas, in 1845. 

Plate II, a, represents this animal much enlarged ; 2, b, the body as seen from above ; 
2, c, the bulb of a tentacle with the red ocellus and the otolitic cell ; 2, d, the peduncle and 
included ovaries ; and 2, c, the arrangement of the tentacles and tubercles. 



OCEANIA GLOBULOSA. 29 

4. Oceania globulosa, Forbes. 
Plate III, Fig. 3. 

This is a minute species, — not larger than a pea, — but so very distinct, that it cannot 
possibly be confounded with any of its congeners as yet described. The umbrella is glo- 
bular, iniiated, and very wide, extending much beyond the orifice on every side, quite smooth 
and transparent. The sub-umbrella is also globular, and occupies rather more than half of 
the interior. Down it run four broad radiating vessels, to join an ample marginal vessel. 
Round the margin are forty close-set purple bulbs, the bases of as many filiform white ten- 
tacula. When the bulbs are magnified, their colour is seen to depend upon rich purple 
ocelli. From the centre of the sub-umbrella hangs a broad and rather short campanulate 
peduncle, of a fawn colour, including, in its upper part, four double, convoluted, orange brown 
reproductive glands. The orifice of the peduncle has four pale, tawny, fimbriated lips. 

I procured two specimens of this singular Oceania in the Sound of Bressay, in 1835. 
It differs so much from the other British species of the genus, that I had some hesitation in 
placing it among the Oceania. In some respects it approaches more closely Turris; but the 
inconspicuous character of the muscular tissue of the sub-umbrella prevents our assigning it 
to that highly-developed group. Plate III, fig. 3, a, represents it, much magnified, as seen 
from the side ; and 3, h, as seen from above ; 3, c, is the peduncle with the included ovaries ; 
3, d, the bulb of a tentacle with its ocellus ; and 3, e, the annulated appearance presented by 
a tentacle when in its most extended state. 



^QUOREADiE. 

Genus V. Stomobrachium, Brandt (1838). 

Umbrella depressed or convex; ovaries 8-12, linear, radiating on the surface of 
the sub-umbrella in the lines of the vessels. Margin with very numerous tentacula. 
Peduncle short, with lobed and fimbriated lips. 

Stomobrachium octocostatum (sp.), Sars. 
Plate IV, Fig. 1. 

Synonyms. Oceania octocostata. Sars, Besk. aj. Jagt., p. 24, pi. 4, f. 9 (1835). 

MeUcertumcampanulatum. Ehrenberg, Berlin Trans., 1835, pi. 8, f. 5-7. 
vEquorea octocostata. Lesson, Acal., p. 312 (1843). 
Thaumantlas Milleri. Landsborough, Arran, p. 265 (1847). 

Of all our British naked-eyed Meduste, I know least about the family to which the 
curious and elegant creature before us belongs. As yet we have only two members of it to 
record — this and a beautiful jelly fish discovered by Mr. Alder. The latter I have never seen 
myself; the former I have not met with since my first season's study of Medusse in 1839, 
when, though I made careful drawings of it, I did not examine its minute structure, trusting 
to meet vnth it again, as it seemed to be one of the most abundant of its tribe. Too often do 
we thus put aside unexamined what seems common and always at hand ; too often do we 
regi-et our inattention when the opportunity is gone ; and this with more serious subjects (some 
could add w-ith objects even more beautiful) than Medusae. 

The genus Stomobrachium was constituted by Brandt for the reception of a Medusa 
presenting characters intermediate between Mesonema and JEquorea^ and connecting the 
family of which the latter is the type with the Oceanida. The only species known to the 
Russian naturalist had been discovered and delineated by Mertens in the South Atlantic 
Ocean, near the Falkland Isles, in the month of January, 1827. 

In 1835, Sars described and figured a little Medusa of the Norwegian seas under the 
name of Oceania octocostata, with the diagnosis, " Disco campanulato, ore plicato brachiis 
nullis ; intus canalibus 8 clavatis, cirris marginalibus longissimis." He accompanied it by a 
fuller account in the Norwegian language. 

The same species had been excellently figured by Ehrenberg in 1835, under the name of 
Melicertum camjmnulatum (which had been apphed in 1829 by Eschscholtz to a very distinct 
Medusa from the Pacific), apparently under the impression that the animal in question was the 
Medusa campanula of Otho Fabricius {Campanella Fabricii of Lesson). 



STOMOBRACHIUM OCTOCOSTATUM. 31 

During the month of July 1839, when cruising with Mr. Smith in the Kyles of Bute, we 
met with numbers of a little Medusa identical with those figured by Sars and Ehrenberg, and 
afterwards found it equally common in the bays of the north-west coast of Ireland. 

The umbrella is very convex and campanulate, transparent, and smooth. The sub- 
umbrella is large, and truncated at the summit. Round the margin of the umbrella is a thick 
fringe of fine, very e.xtensile, tentacula (40 — 60, according to Sars ; if the former number, the 
formula might probably be 8x4+8). Between each of these Ehrenberg represents a very 
small tentacle, which is borne upwards or obliquely when the larger ones are extended, and he 
also figures a vesicle (otolitic ?) at the origin of each of the small tentacula. Down the 
sub-umbrella run eight simple vessels, and in the course of each is a conspicuous linear 
furbelowed ovary or reproductive gland. The ovary and the tentacles are of a golden yellow 
colour. The peduncle is also yellow, short, but broad, and is suspended from the centre of the 
sub-umbrella ; it is occupied by the stomach, and terminates in four fimbriated lobes or lips, 
which change their forms so as sometimes to appear as if eight in number. 

This account of a beautiful and probably not rare species is evidently insufficient, though 
enough for its recognition, and I hope on some future opportunity to observe it more completely. 
Of its aspect and habits in confinement, a graphic account has been given by my friend the 
Rev. David Landsborough, in his delightful volume on the Island of An-an, not the less 
graphic for proceeding in part from the pen of Hugh Miller, who could not fail to present a 
vivid picture of this animated and beautiful bubble, after having endowed the fragmentary and 
motionless remains of fossil fishes with such vitality, that no reader of his admirable ' Old Red 
Sandstone' rises from its perusal, without the fond impression of having seen in his mind's 
eye the Pterichthys and Coccosteus, and their strange companions of the deep, not as shapeless 
lumps of rock, ranged in orderly rows under sheet-glass, but as living monsters sporting in a 
primaeval sea. 

" There was a Medusa," writes Mr. Landsborough, " discovered by my son David, which 
was quite new to us, and, from its minuteness, probably known to few. We took it home, and 
put it in a tumbler of sea-water, that we might better observe its structure and graceful 
evolutions. I would have attempted to describe it, but glad was I, soon after we had seen it, 
to find this done to my hand by one who is acknowledged in the scientific world as a graphic 
describer of nature — Mr. Hugh Miller ; best known among men of science as the author of 
the truly interesting work on the ' Old Red Sandstone,' but better known to our countrymen 
in general as the talented Editor of the ' Witness.' Nothing escapes his scientific eye ; and 
from his ' Summer Rambles,' I learned that he had about the same time discovered it, when 
aboard the Betsy, off the Island of Eigg. He speaks of two — one scarcely larger than a 
shilling, ' another still more minute,' (ours, I think, about the breadth of a sixpence), ' and 
which presenting in the water the appearance of a small hazel nut of a brown-yellowish hue, 
I was disposed,' says he, ' to set down as a species of Beroe. On getting one caught, how- 
ever, and transferred to a bowl, I found that the brown-coloured, melon-shaped mass, though 
ribbed like a Beroe, did not present the true outline of the animal ; it formed merely the 
centre of a gelatinous bulb, which, though scarcely visible even in the bowl, proved a most 
effective instrument of motion. Such were its contractile powers, that its sides nearly closed 
at every stroke behind the opaque centre, like the legs of a vigorous swimmer ; and the 
animal— unlike its more bulky congeners, that, despite of their slow persevering flappings, 



32 POLYXENIA ALDERI. 

seemed greatly at the mercy of the tide, and progressed all one way — shot, as it rolled back- 
ward, forward, or athwart.' The transparent tumbler gave me this advantage in observing 
it, that I could use a magnifying lens when it approached the side. Notwithstanding this 
advantage, it was some time before I observed the true form of the animal, as Mr. Miller's 
excellent description had not then been published. The transparent ball that rose above its 
body was so very pellucid, that it was a good while before I observed it all. It rose to a 
considerable height above the buff-coloured body of the animal ; and it was elegantly shaped, 
like the fine crystalline shades often placed over stuffed birds, or artificial flowers, or miniature 
figures formed of pure alabaster. The finest crystal vase was clumsiness itself when compared 
with it. It was as fine as the transparent soap-bubble blown out of a pipe ; and we doubt 
not that, like this bubble, it would have been iridescent had it been so placed as that the sun 
could have shone on it. Delicate as its fabric was, the vigour of the little creature was very 
remarkable, and has been well compared to the efforts of a strong swimmer, as it alternately 
contracted and expanded its pellucid organization. The margin of its mouth had a close 
fringe of brownish tentacula. By aid of the lens, I could observe that they were drawn in 
when the body was contracted, but that at every stroke they were protruded like forked 
lightning, or like feathered serpents, darting and flashing forth, till they were longer than the 
body of the animal." {Landsborough, Arran, chapter xvi, p. 260.) 

When I met with this Stomohrachimn in 1 839, I fortunately made a drawing of it in 
two states, the one (Plate IV, fig. 1, a and b) having much wider reproductive glands than 
the other (Plate IV, fig. 1, c), in which they are linear and paler, and reach quite to the 
margin. The former state is identical with that observed by Ehrenberg, who found ova in 
the folds of the ovaries ; the latter is not improbably the male animal. 



Genus VI. Polyxenia, Eschscholtz (1839). 

Umbrella spreadinj^, more or less depressed. Ovaries numerous, linear, in the 
centres of triangular spaces which reach nearly to the circumference. Marginal 
tentacles between each pair of ovaries. Peduncle very short, terminating in an oral 
orifice, surrounded by four lanceolate arms. 

Polyxenia Alderi, Forbes. 
Plate IV, Fig. 2 (under the name of F. ci/anostilis). 

I have ventured entirely to redefine the genus Polyxenia, and to regard it as represented 
by the beautiful Medusa now figured from a drawing of my accomplished and' distinguished 
friend Mr. Alder, who observed it on the coast of Devon, in 1846. The umbrella is expanded, 
and rather depressed, smooth, and transparent. Its margin is deeply notched by sixteen 
indentations, out of which spring as many rosy tentacula, alternately longer and shorter. The 
sub-iuubrella is divided into sixteen somewhat triangular spaces, which commence at about 
two thirds of its height, and terminate near the margin. In the centre of each of these spaces 



POLYXENIA ALDERI. 33 

is a linear ovary. Each space terminates marginally in two truncated lobes, one on each side 
of the ovary, and always alternately longer or shorter. Each lobe, especially the shorter one, 
is slightly lobulated. From the centre of the sub-umbrella springs a short peduncle, very 
soon terminating in four linear-lanceolate, rather long, arm-like lips, white, tipped with rose 
colour. They project beneath and lower than the umbrella. The breadth of the disk is 
about two inches. 

I had referred this graceful creature to the Polyxenia cyanostyla of Eschscholtz, and it 
is so named in the plate. Anxious, if possible, to retain and illustrate an old species, little 
known, rather than add a new name, against my better judgment I persisted in this opinion. 
On more recent consideration, however, I feel bound to abandon it, and do so the more 
willingly, since it enables me to retain the name of its discoverer as patron-saint of at least 
one of the new species here described. 

It may perhaps be disputed whether the genus Polyxenia is its proper place, and 
whether I do right in changing the characters of the genus as above. An earnest conside- 
ration of the probability of the nature of the animal, very imperfectly described by the founder 
of the genus, induces me to do so. His figure of Polyxenia cyanostilis appears to me to 
represent a mutilated animal, deprived of its lips ; and could we restore it, I beheve the repre- 
sentation now given (Plate IV, 2, a, natural size ; 2, b, much enlarged) would closely approach 
the figure of the creature delineated in the ' System der Acalephen.' 



CmCEADiE. 

Genus VII. Circe, Mertens (1838\ 

Umbrella coinco-campanulate ; ovaries eight, placed on the sub-umbrella at a 
little distance around its summit ; vessels eight, simple, passing through the ovaries, 
and opening into a marginal vessel ; tentacula very numerous, and placed in a 
single series around the margin ; peduncle cylindrical, contracting near its extremity 
to form the small campanulate stomach, the orifice of which is furnished with four 
lanceolate lips. 

Circe rosea, Forbes (1846). 
Plate I, Fig. 2. 

The genus Circe was constituted by Mertens for a remarkable little Medusa found by 
him on the coast of Kamtschatka. Brandt adopted this group in his ' Prodromus,' and 
afterwards more fully in the ' Petersburgh Transactions,' where he described the species 
discovered by Mertens under the name of Circe camtscJuitica, and engraved it from the 
drawings of its discoverer. Lesson, in his ' History of the Acalephse,' added two more 
species to the genus under the names of C. elongata, and C. anais, both discovered by Rang 
in the African (west ?) seas, and figured from his drawings. 

The Medusa which I now figure and describe under the name of Circe rosea, is the 
first of the genus noticed in the European seas. Though very small, — the largest specimens 
taken not being more than half an inch in height, — its aspect is very beautiful and striking. 
The umbrella is oblong, and somewhat mitre-shaped, with an apiculate summit. The outer 
surface is quite smooth. Round the margin is a close and single series of small tentacula, 
which, however, seem capable of considerable elongation, though usually retracted. Their 
number varies slightly. They are all similar, and in the examples taken, were 56 in number. 
Their formula might stand as 6x8-|-8. They are all external to a veil, which guards the 
orifice of the sub-umbrella, and is marked by sixteen denticulations. I could detect no ocelli at 
the bases of the tentacula. From the centre of the sub-umbrella depends a long cylindrical 
peduncle, which reaches nearly to a line with the margin, and is contracted a little above the 
orifice, so as to form a kind of proboscis terminated by four simple lanceolate lips. The 
stomach is very short, and terminates almost at the upper point of contraction, where we see 
eight gastro-vascular canals commence, run up the side of the peduncle, distinct from each 
other, turn at its base, and descend the sub-umbrella, till they reach and unite with the marginal 
vessel. In the uppermost part of their course along the sub-umbrella, they pass through as 
many small ovate, simple generative glands, which, when the animal is seen from above. 



CIRCE ROSEA. 35 

appear like so many clubs or spokes round a central axle. The body is of various shades of 
pink, and the ovaries yellow ; consequently, although the creature is so transparent, and 
unfurnished with brilliantly coloured ocelli to mark its position in the water, it is not uncon- 
spicuous nor diiEcult to see. 

The first specimen observed of this species was taken by Mr. M'Andrew and myself in 
the Zetland seas, in August, 1845, off Vella, seven miles from land. We afterwards met with 
several in Brassay Sound, on the opposite coast of the mainland. 

Compared with Brandt's species, it differs in the form of the ovaries, in general aspect 
and colour, and in the absence of conspicuous ocelli, eight of which, apparently colourless, 
are represented in the beautiful drawing by Mertens (Petersburgh Memoirs, Sixth Series, 
Sc. Nat., t. ii, 1838, pi. 1, f. 1-5). The simple series of tentacula distinguish it from Circe 
anais, also rose-coloured. (Lesson, loc. cit. pi. 5, f. 1.) It is more nearly allied to 
C. elongata, but differs in form. (Lesson, loc. cit. pi. 5, f. 2.) 

The genus Circe is evidently closely allied to Thaiimantias, Dianma, and Geryonia, 
differing mainly in the number of ovaries and of gastro-vascular canals. The generative 
glands may be said to be those of Thaumantius, the peduncle to be that of Dianceu. Brandt 
placed it among the OceanidcB, and Lesson strangely introduces it between his genus Turris, 
and Brandt's genus Conis, genera belonging to a very different group, and, as I have shown, 
scarcely deserving of separation, both being close allies of Oceania. With the Oceanidce, 
however, Circe has evidently no affinity. 

Plate I, f. 2, Circe rosea, natural size ; 2, a, greatly enlarged ; 2, b, as seen from 
above, showing the relative arrangement of the vessels and ovaries ; 2, c, tentaculated 
margin and denticulated veil ; 2, d, an ovary ; 2, e, portion of the margin, between two of 
the radiating vessels ; 2, f, peduncle, and small stomach at its extremity, with the oral 
orifice ; 2, g, lips of the oral orifice. 



GERYONIADiE. 



Genus VIII. Geryonia, Peron (1809). 

Umbrella hemispheric, margin furnished with tentacula in variable number, 
usually few ; radiating vessels five ; ovaries four, phylliform, on the surface of the 
sub-umbrella in the course of the vessels ; peduncle rather long, conico-cylindrical, 
terminating in a small campanulate stomach, opening by four lips. 

Geryonia appendiculata, Forbes. 

Plate V, Fig. 2. 

One of the most phosphorescent Medusa of the British seas is the graceful little animal 
I have now to describe. The umbrella is smooth, transparent, and of a hemispherical shape, 
sometimes nearly sub-globose, slightly elongating during the act of ascending, becoming 
more spherical when descending. The margin is fringed by eight distant tentacula, four of 
which are of a different structure from the other four ; the former being cylindrical, and 
composed of similar tissue (fibrous cells) throughout, whilst the latter have an outer and inner 
side of different composition, the fibrous cells being collected in bundles or masses at regular 
intervals on the inner surface. Both orders of tentacles appear to be tubular. Those of the 
first kind are highly contractile, and capable of great extension, trailing after the body to 
twice or three times its length when the animal rises, becoming short and thick when it 
sinks. The intermediate tuberculated tentacles have not an equal power of extension. At 
the origin, and by the side of each of the more extensile tentacles, is a globular body within 
a cavity (otolitic ?) produced linearly upwards to terminate in a vesicular body, also with 
contents (a rudimentary ocellus ?). A similar, though smaller and less defined, structure is 
seen at the bases of the intermediate tentacles. Opposite to each of the longer tentacles is a 
simple gastric vessel joining a marginal one, and proceeding down the inner surface of the 
rather conical and truncated sub-umbrella. Towards the summit of the sub-umbrella in the 
course of the vessels are the bases of four leaf-shaped, cordate, reproductive glands (in the 
instances examined, ovaries). Round the opening of the sub-umbrella is a rather broad, 
shelf-like veil. From its centre depends a long conical peduncle, extending for half its length 
below the orifice of the disk. Its base is thick, and it tapers rather gradually nearly to the 
extremity, a little above which there is a constriction, marking the commencement of the 
broad campanulate stomach. The gastric vessels commence at this point, and run up the 
peduncle. The oral orifice is four-lobed, but these lobes or lips change form considerably, 
sometimes appearing acute, sometimes obtuse, and occasionally as if six-lobed. They are 
bordered by a highly contractile tissue composed of fibrous cells. Indistinct bands of similar 



TIMA BAIRDII. 37 

tissue are present in the substance of the peduncle, and give it its power of motion, so as to 
cause it at times to remind us of the proboscis of some pacliydcrm. With the exception of a 
tinge of green on tlie ovaries, and of pink on tlie tentacles, this animal is colourless. The 
specimens taken measure, body and trunk, from one inch to one inch and a half. 

Two specimens were taken by Mr. M'Andrew and myself in the Reach of Dartmouth, 
Auo-ust 31st, 1846. We afterwards took some at a distance of fifteen miles from land, off the 
coast of Devon and in the west bay of Portland. The peculiar distinctive characters of the 
tentacula distinguish this beautiful Genjonia from any recorded species. 

I have adopted the genus Genjonia in the limited sense accorded to it by Lesson, 
restricting it to those pedunculated forms of the group as first named by Peron and Lesueur, in 
which we find four phylliform ovaries. It thus constitutes a natural and probably extensive 
assemblage of species. Of the GeryotiicE, as restricted by Eschscholtz, four out of his six 
species, viz. G. tetrciphyUa, G. bicolor, G. rosacea, and G. exigua, will fall under the section 
so defined. Lamarck had united them with very different animals, such as Oceania and 
Pelagia, in his very unnatural assemblage, Dianaea. De Blainville places it as a section of 
his extended genus Genjonia, distinguished from his Dianmi by the presence of phylliform 
ovaries. The few species recorded of tins genus are widely distributed : three are from the 
seas of the tropics, both Indian and Pacific ; one is from Gibraltar ; and one from the Adriatic 
{G. planata), briefly but excellently described and figured by Dr. Will. I have found a 
species in the iEgean. 

Genjonia ajrpendicidata, as represented in Plate V, f. 2, is the animal seen in profile 
during the act of descending and contraction, (2, a, indicates the natural size,) and 2, c, is the 
same when ascending and expanding ; 2, h, represents the position of parts as seen from 
above ; 2, cl, is an ovary magnified ; 2, e, represents the two sorts of tentacles contracted, 
with the vesicles at their bases ; these are represented still more highly magnified at 2, /, 
and 2, g. The proboscis, with the stomach and its lips, and the commencement of the gastric 
vessels, is shown in 2, h. 



Genus X. Tima, Eschscholtz (1839). 

Umbrella hemispheric ; margin with (rather distant) tentacula ; radiating 
vessels four, simple ; ovaries four, linear in the course of the vessels ; peduncle 
cylindrical, terminating in a campanulate stomach with four fimbriated lips. 

Tima Bairdii (sp.), Johnston (1833). 

Synonym. Biancea Bairdii. Johnston, in Loudon's Magazine of Natural History, 

vol. vi, p. 320, fig. 41 (1833). 

One of the most earnest and enthusiastic of British naturalists, the author of the excellent 
' History of British Zoophytes,' discovered the elegant Medusa now to be described, and 
dedicated it to his friend Dr. Baird, a gentleman whose skill in some of the minutest and 
least investigated departments of zoology has gained him a high reputation, and who, devoting 



38 TIMA BAIRDII. 

his time and sacrificing his professional pui'suits to his favorite science, is now one of the 
unselfish band of worthy assistants in the National Museum. 

Dr. Johnston met with the Tima Bairdii in Berwick Bay, on the 27th of September, 
1832, floating on the surface of the water; and published an account of it in Loudon's 
Magazine for the following year, illustrating his description by an expressive sketch from the 
pencil of his accomplished and esteemed lady, to whom, as to her eminent husband, our 
science owes many debts of gratitude. He describes it as " a semi-globular mass of a perfectly 
translucent and almost colourless jelly, divided by four opaque, milk-white, narrow ligaments, 
or bands, into four equal compartments. These bands arise at the angles of the mouth, and 
are at first very fine, but become broader and somewhat curled in their course towards the 
upper surface. The very delicate membrane investing the body is folded at the margin, 
which is furnished with a circle of rather distant, tapered, white tentacula. In our specimen there 
were thirteen of these. The under side is produced in the middle, so as to form a kind of stalk, 
at the apex of which is the mouth, of a square form, and encircled by four plumose branchial 
appendages. When magnified, these are shown to be formed of a thin membrane, beautifully 
but irregidarly folded, like a frill, and edged with a neat thickened border. BiancBu Bairdii 
seems to be invested with two membranes, of great tenuity. The outer one covers all, like as 
it were a glass inverted over a smaller globe, the intermediate space being occupied with a 
consistent but colourless jelly, in which neither vessels nor membranes can be distinguished. 
This coat forms two loose folds around the circumference, from the innermost of which the 
tentacula arise, and the inner coat is probably a continuation of the outer reflected upwards ; 
but it is not a simple membrane, since several laminae can be perceived to cross the body. 
The white conical bands adhere to this inner envelope ; they are not muscular, but very 
probably Ijelong to the generative system. I consider the plumose processes at the oral 
aperture as subservient to respiration, partly because of their position, and partly because of 
their folded structure ; intended, as it appears to be, to expose the largest possible surface to 
the action of the water. This Medusa is a luminous species. It gives out a copious light of a 
whitish colour, when the water in which it swims is agitated, or when it comes in contact with 
foreign bodies." 

Dr. Johnston compares it with a minute phosphorescent Medusa taken by Dr. Baird in 
the Straits of Malacca, and considers the two identical. In all probability, however, they are 
quite distinct, though congeneric. 

During the winter of 1839, I met with this handsome Medusa in two localities : first, in 
the harbour at Burtisland, in the Frith of Forth, and afterwards at St. Andrews, where a 
number of individuals were cast ashore on the sands, along with Cydippe pileus, and 
C. Flemingii, and Aurelia aurita, after a very stormy night. 

The umbrella, in the specimens I examined and drew, was hemispheric, or rather cam- 
panulate, with a slight constriction, giving it an elegant undulation about the upper third. 
It varies, however, much in form. It is smooth, transparent, and colourless. The margin 
bears sixteen tentacula, usually contracted, and tinged with pale pink. Between the tentacles 
there is the appearance as of a scalloped veil. The sub-umbrella is truncated, and marked by 
four radiating vessels, opening into a circular marginal one. In the course of these four 
vessels appear to be four very narrow, linear, whitish reproductive glands. From the centre 
of the sub-umbrella depends an ample cylindrical peduncle, extending for about a third of its 



GERYONOPSIS DELICATULA. 39 

length below the opening. Down it run the gastric vessels to the most constricted point, 
where they join a short campanulate stomach set on as if obliquely, and opening by four 
ample, triangular, beautiful crimped lips. The breadth of the umbrella was two inches. I 
saw no ocelli, though very possibly they are present. I placed one of these animals in a 
tumbler of sea-water with some specimens of Cijdippe equally alive and active. Before lono- 
the Medusa seized one of them with its lips, and made such vigorous exertions to swallow the 
ciliograde, that I had great difficulty in rescuing its Adctim. 

The peculiar form of the reproductive glands, which, indeed, are difficult to detect, 
prevent us placing this animal in Geryonia, to which genus it has close affinity, or in BiancBa 
(regarding Diancsa iwohosciadalis as the type of the latter). The genus Tima of Eschscholtz 
seems to be its proper place. ["Discus facie infera in conum productus. Ventriculus 
plicatus in apice coni situs. Canali quaterni cum circulo marginali conjuncti. Cirrhi 
marginales numerosi."] The figure given by that author of his Tima flavilabris, from the 
neighbourhood of the Azores, bears considerable resemblance to that of the species before us, 
but the number of tentacula (80) is much greater. 

Plate V, f. 1, Tima Bairdii during contraction; 1, a, during expansion; 1, 5, as seen 
from above. 



Genus X. Geryonopsis, Forbes, 

Umbrella hemispherical ; ovaries four, clavate, conspicuous on the sub-umbrella 
in the course of the four simple radiating vessels ; margin of umbrella with numerous 
short tentacula ; stomach at the extremity of a short peduncle, terminating in four 
large fimbriated lips. 

Geryonopsis delicatula, Forbes. 
Plate IX, Fig. 1 (under the name of Tliaumantias cymbaloides). 

Anxious to retain, if possible, the names of recorded species in our British lists, I 
referred to the Medusa cymbaloides of the older acalephologists, a very graceful and tender 
jelly-fish, which is not uncommon on the south coast of England. With much reluctance I 
abandon the reference, for though the character of the stomach, " voluminous and much 
exceeding the border," mentioned by Peron, applies only to the animal before us among all 
the immediate allies of 'llianmantias, one of which, i]\e Medusa cymbaloides of Slabber, must 
be in the British seas, the other character of " tentaculis sedecim, basi bulbosis," adopted in 
all diagnoses of that species, cannot by any stretch be made to apply, and is too positive to 
be founded on imperfect obsen'ation, since the bulbs of the tentacula are distinctly stated to 
be brilliantly ocellated. I trust further research will, before long, make better known to us 
the original species of Slabber and Modeer.* 

The umbrella of my Medusa is hemispheric and rather depressed, smooth, transparent, 
colourless, and of a singularly delicate texture. The margin is encircled by a row of very 

* Plate IX was unfortunately altered and printed off before the name could be corrected. 



40 GEKYONOPSIS DELICATULA. 

short tentacula, numerous (l6x4-|-4), but not placed in contact, and irregularly developed. 
They are of a slightly milky hue, and have slightly bulbous bases, but no coloured ocelli. 
There appears to be an otolitic mass in the cavity of each bulb, but I could observe no motion 
The sub-umbrella is hemispherical and depressed ; down it run four simple vessels to join the 
marginal vessels, and in their course are four greenish conspicuous, linear-lanceolate, or 
rather claviform, reproductive glands, with wavy margins. Round the inner margin of the 
umbrella is a horizontal, broad, membranous veil or shelf. From the summit of the sub- 
umbrella depends a conical peduncle, which projects beyond the margin, and, after contracting, 
suddenly expands into a wide campanulate stomach, with four large, lanceolate, fimbriated 
lips, whose edges are bordered by a thick layer of fibrous cells, endowing them with a highly 
motor power. Into the summit of the cavity of the stomach the four gastric vessels arc plainly 
seen opening. The lips are tinged with green. The disk measures one inch and a half across. 
Several specimens occurred on the coasts of Dorset and Devon during August 1836, 
especially in the Reach of Dartmouth. It was usually in company with the Geryonia 
appen die id a ta . 

In 1839, Professor Goodsir and I took a Medusa at Scalloway, in Zetland, presenting 
many of the characters of the species we have described, but differing in having much longer 
and more numerous tentacula, more clavate and purple ovaries, and pale fimbriated lips. We 
announced it at the meeting of the British Association, at Birmingham, as a new Oceania, and 
it has since been quoted as such by Lesson. For the present, however, it is better to abstain 
from naming it, though in all probability distinct, since the drawing and memoranda made at 
the time are insufficient ; the animal, owing to unavoidable circumstances, not having been 
submitted to microscopic observation. I may here remark, once for all, that under no circum- 
stances can any of the naked-eyed Medusne (above all, those belonging to the tribe we are now 
treating of) be identified without aid of the microscope, and the accident of that instrument 
not having been employed for the examination of Acalephse by the greater number of 
zoologists who have v^Titten upon them systematically, has rendered published diagnoses and 
determinations so imperfect, as, in the majority of instances, to be little better than utterly 
worthless. The quoting of authorities, synonyms, and localities respecting the subjects of 
this monograph is, in fact, an act of courtesy to those who have gone before us, rather than 
of justice to science, which would have thriven better if half the bad figures and worse 
descriptions of the smaller Medusae had never seen the light. Too truly in this case might it 
be said of such describers — 

" They have perplexed, 
With a dark comment, Beautj^'s clearest text ; 
Tliey have not told lier face's story true. 
But brought false copies to our jealous view." Carew. 

Of course, there are some honorable exceptions to this censure, especially Sars, Wagner, Milne 
Edwards, and Will. 

The higher Medusne, probably because larger, have been more fortunate. The 
Geryonia peUucida of Will (Horse Tergestinae, p. 70, pi. 2, f. 8) is a Geryonopsis not far 
removed from the species before us. It difi'ers in having more clavate ovaries, simpler lips, 
and finer marginal tentacula (64). Dr. Will relates of his species that it devoured eggs of 
Beroe rufescens, of which some living specimens kept in the same vessel had laid many eggs, 



THAUMANTIAS. 41 

and states that his friend Dr. Koch observed it eat mutilated fragments of the ciUograde 
Eucharis multicornls. The coarser indigestible parts of its food were ejected by the mouth 
in doing which the stomach shortened considerably, and everted itself partially. 

Plate IX, f. \, a, represents a large specimen of Geryonopsis delicatula of the natural 
size; 1, h, shows the structure of the lips, and the origins of the gastric vessels ; 1, c, is a 
reproductive gland, the vessel passing through its centre ; 1, r/, a tentacle with its bulb ; and 
1, ^, a partially developed tentacle. 



Genus XI. Thaumantias, Eschscholtz (1829). 

Umbrella hemispherical, in some species almost globular, in others much 
depressed ; ovaries four, varying in form from ovate to linear, conspicuous on the 
sub-umbrella in the course of four simple radiating vessels ; margin of umbrella with 
tentacula in variable number (from 4 to 200) according to the species, their bulbs 
alvvaj's ocellated ; stomach sessile, dependent from, and almost always included 
within the sub-umbrella ; mouth vvith four lips, rarely fimbriated. 

This excellent genus was instituted by Eschscholtz for the reception of the Medusa 
cymhaloidea of Slabber, and the Medusa hemisphcerica of Gronovius. The latter is so much 
better known than the first, that it may be regarded as the type. They are probably, 
however, identical. When Lesson published his ' History of the Acalephae,' in 1843, he 
enumerated nine species of Tlmumantias, two of them Norwegian, discovered by Sars, and 
four British, described by myself in the ' Annals of Natural History.' The ninth was the 
Medusa lucida of Macartney, another name for T. hemispheerica. 

Of all the naked-eyed Medusae, those belonging to this genus are most common in our 
seas, swarming in countless myriads in many of our bays and harbours. They are among 
the most usual causes of phosphorescence. It might be expected that animals so 
abundant, when carefully sifted, although so similar, would be found to include several 
distinct kinds. I have now to describe no fewer than seventeen British species of Thaumantias, 
of which the greater number are all so very distinct from each other, that they cannot 
be confounded. I believe many more equally distinct will be before long discovered in the 
European seas. 

The characters in common presented by many of these kinds are such as to enable us 
conveniently to group them in sectional assemblages, dividing them, in the first instance, 
under two sub-generic heads: — 



42 THAUMANTIAS PILOSELLA. 

A. Marginal tentacles of two kinds (Cosmetira). 

1. Tliaumantias pilosella. 

B. Marginal tentacles of one order only (Thaumantias). 

* Marginal tentacles four. 

2. T. quaclrata. 

3. T. cBronautica. 

** Marginal tentacles eight. 

4. T. octona. 

*** Marginal tentacles sixteen and upwards. 

f Umbrella very convex or globose. 

5. T. maculata. 

6. T. melanops. 

7. T. glohosa. 

8. T. convexa. 

9. T. gibhosa. 

tt Umbrella depressed. 

10. T. lineata. 

11. T. pileatu. 

12. T. Sarnica. 

13. T. Tliompsoni. 

14. T. hemisphmrica. 
] 5. T. inconspicua. 

16. T. punctata. 

17. T. lucifera. 



A. Marginal tentacles of two kinds. 



'^ 



[Cosmetira.] 



1. Thaumantias pilosella, Forbes. 
Plate VIII, Fig. 1. 

By far the most beautiful, and among the largest, of the British kinds of Tlmumantias is 
that I have first to describe. The umbrella, which sometimes measures nearly two inches in 
diameter, but more usually one, or one and a quarter, is hemispheric, and shaped like a watch- 
glass, but much more convex. It is transparent and smooth, except on the sides towards the 
margin, where it is as if woolly, being invested with minute epidermic hairs composed of fibrous 
cells. These, though sufficiently conspicuous, may escape the observer who is not aware of their 
presence, in consequence of their transparency. The margin is fringed by very numerous 



THAUMANTIAS QUADRATA. 43 

(24x4+4), extensile (but usually borne rather short), pale pinkish tentacula, with bulbous 
bases. The bulbs are ocellated, with dense crescentic masses of purple pigment-cells. 
When the margin is much magnified, it is seen to be bordered by a narrow band or thread of 
fibrous cells, from which the tentacles spring, and between each pair there are six or seven 
short, fine, secondary tentacles, without ocelli at their bases. The coloured bulbs of the 
larger tentacles are very conspicuous, and appear in the water as a circle of brilliant purple 
gems. The inner margin of the umbrella is bordered by a shelf-like veil. The sub-umbrella 
is depressed, and on its surface run the four radiating vessels with a long, linear, somewhat 
clavate ovary, of a bright pink colour, commencing very near the centre, and terminating close 
to the margin in the course of each. The stomach is very short, but wide, of a rose colour, 
and has four lanceolate, fimbriated lips, bordered by a compact edging of fibrous cells. 

This beautiful Medusa is very abundant in the bays and harbours of Zetland, especially 
in the Sound of Brassay, where it is the most common species of its genus. Mr. Alder met 
with it on the south coast of England, at Falmouth, in 1847. 

Plate VIII, fig. 1, a, represents its usual appearance, twice the natural size ; 1, 6, as seen 
from above, of the size of nature ; \,c, an ovary ; I, <:/, a lip ; \,e,a. pair of the larger ten- 
tacula with their ocelli, and the smaller tentacula between them. 

B. Marginal tentacles of one order. 

[Thaumantias.] 

* Marginal tentacles, four. 

2. Thaumantias quadrata, Forbes. 

Plate IX, Fig. 2. 

Umbrella campanulate, rather elongated, becoming globose during contraction, smooth, 
pellucid, colourless ; margin with four equidistant tentacula springing opposite to the gastric 
vessels from large and conspicuous bulbs. The tentacula are rather stout, dusky purple, and 
conspicuously granulated on the surface. The bulbs are bright yellow, with ocelli composed 
of vermilion pigment-cells loosely grouped together. A similar ocellated bulb is in the centre 
of the marginal space between each pair of tentacles, but is not so large as those at their 
bases, and exhibits no rudiment of a tentacle springing from it. In the interspaces again, 
between each intermediate bulb and the tentacular one, is a very small yellowish tubercle, 
without an ocellus. I have no reason to think that tentacula are developed from any of the 
bulbs, except the four largest. The four ovaries are ovate, pale yellow, and placed in the 
course of the radiating vessels in the lower half of the sub-umbrella, towards the margin. The 
inner margin is surrounded by a veil. The proboscis is rather short, but slender and narrow, 
quadrangular, yellow, lineated with red. It terminates in four simple, triangular, acute lips. 

This little species was observed very abundantly in the harbour of Tarbet, Loch Fyne, 
in the autumn of 1845. The umbrella is scarcely more than two tenths of an inch in length. 

Plate IX, f. 2, a, represents it of the natural size ; 2, b, magnified, and seen from the side; 
2, c, as seen from above ; 2, d, an ovary ; 2, e, tentacle and its bulb, with the intermediate 
bulbs. 



44 THAUMANTIAS OCTONA. 

3. Thaumantias teronautica, Forbes. 
Plate IX, Fig. 3. 

The umbrella of this minute species, which is still smaller than T. quadraia, is oblong, 
and becomes elongated during conti'action. It is smooth, transparent, and colourless. The 
margin bears four fine, colourless, filiform tentacula, which are often extended to a prodigious 
length, as compared with the dimensions of the body, or with the small size, both of length 
and diameter, into which they can contract themselves. When magnified, they exhibit a 
moniliform appearance. They spring from four large bulbs of a pale yellow colour, with traces 
of an interior vesicle, but no brightly coloured ocelli. In the interspaces of the margin are 
yellow tubercles, three between each pair of tentacles, the central one largest, but all very 
small as compared with the tentacle-bulbs. Down the sub-umbrella, which is of an oblong 
form, and sometimes, when vigorously contracting, of a pear-shape, run the four gastric vessels, 
traversing at exactly half its height as many very small ovate reproductive glands, of a pale 
yellow colour. The stomach is sessile ; when contracted, shortly campanulate ; when ex- 
tended, long and quadrangular, reaching nearly to a level with the ovaries. The mouth is 
surrounded by four simple lanceolate lips. 

Small as this pretty creature is, it is evidently adult, and presents its permanent characters. 
When confined in a glass tube, filled with salt water, it resembles a miniature balloon moored 
by five silken cables. I met with it in localities far apart, viz. off Brassay, and in Hamna Voe, 
in Papa, both in the Zetland seas, in July 1 845 ; and during the following month in the Sound 
of Skye, among the Hebrides. All the specimens were alike. I judged it to be adult, from 
the defined and firm condition of the reproductive glands, and the microscopic structure of 
the tentacles and their bulbs. 

Plate X, f. 3, a, represents it of the size of nature ; 3, b, magnified, with the tentacles 
extended ; 3, c, seen from above, with the tentacles contracted ; 3, d, the stomach extended ; 
and 3, e, contracted ; 3,/", one of the reproductive glands. 

** Marginal tentacles, eight. 

4. Thaumantias octona, Forbes. 
Plate VIII, Fig. 4. 

The umbrella of this small and peculiar species, is only about two tenths of an inch in 
height, globose, or sub-orbicular, smooth, and transparent. The margin presents the peculiar 
feature of bearing eight exactly similar pinkish tentacles, springing from conspicuous, bright 
yellow, bulbous bases, each bearing a defined red ocellus. In the marginal spaces, between 
each pair of tentacles, are two colourless tubercles placed close together. These do not 
appear ever to give rise to tentacula. The sub-umbrella is short compared with the umbrella, 
and is hemispherical. In the course of the four vessels, which run to join the marginal 
vessel opposite to the origins of four of the tentacles, are four small yellow or tawny ovate 
ovaries, placed on the lower half of the sub-umbrella. The stomach is very small in proportion 
to the size of the animal, rather elongated, and quadrangular in shape, of a yellowish or fawn 
colour, with four minute black dots at its base. The four lips are short, acute, and triangular. 



THAUMANTIAS MELANOPS. 45 

Many specimens of all ages, all preserving the peculiar characters mentioned, occurred 
at Tarbet, in Loch Fyne, in company with Thaumantias quadrata, during the autumn of 1845. 
We had previously taken it by the tow-net at Oban. 

Plate VIII, fig. 4, «and i, represent this species magnified, seen in profile and from above ; 
4, c, is an ovary ; 4, d, the stomach ; 4, e, part of the margin and tentacula ; 4,y, the bulb 
of one of the tentacles. 

*** Marginal tentacles sixteen and upwards, 
t Umbrella very convex or globose. 

5. Thaumantias maculata, Forbes. 

Plate IX, Fig. 4. 

The umbrella of this very distinct species is globular, smooth, pellucid, and colourless. 
The margin of its rather contracted opening is ornamented with sixteen jet-black ocelli, 
(3 X 4+4), which are alternately larger and smaller, and all very conspicuous, and large in 
proportion to the body. Between each of these is a small colourless tubercle. From each of 
the ocelli springs a colourless tentacle. All the marginal tentacles are similar, and very 
nearly of a size. The sub-umbrella is hemispherical, and divided into four equal segments 
by the gastric vessels. On its lower half, in the course of the vessels, are four ovate repro- 
ductive glands, pale, with yellow or tawny centres. From the centre of the sub-umbrella 
hangs a short but wide campanulate stomach, with four broad, slightly fimbriated lips. 
The four lips correspond in position to the four ovaries, and on the sides near the base of 
the stomach, alternating with the lips, are four patches of black pigment-cells, giving the 
centre of the animal, when seen from above, the appearance of being marked by four 
conspicuous black spots. The black bulbs of the tentacles, when compressed and highly 
magnified, are seen to be coloured by a crescentic series of black pigment-cells, forming the 
ocellus, bounding a tawiiy space in which there is seen an otolitic capsule. The tentacula 
themselves have a highly annulated appearance. The body measures about a quarter of an 
inch in height. I have met with this curious Thaumantias in the Zetland Islands only. It 
occurred several times in the Sound of Brassay, but was never plentiful. The jet-black eyes 
and stomachal spots render it a veiy striking object in the water. 

Plate IX, f. 4, a, represents it rather more than twice the natural size ; 4, b, as seen from 
above ; 4, c, the ocellated bulbs ; 4, d, the base and portion of a tentacle highly magnified, 
showing the distinction between the ocellus and otolitic capsule in the bulb ; 4, e, the stomach 
with its lips, spots, and the origins of the gastric vessels, seen from above ; 4, f, one of the 
reproductive glands. 

6. Thaumantias melanops, Forbes. 

Plate X, Fig. 3. 

Another black-eyed beauty, though unarmed with such jetty piercing orbs as the sister 
species last described. Instead of a few conspicuous oceUi, we have here an almost countless 
number, all, however, of extreme minuteness. Argus, the hundred-eyed, must yield to our 
Thaumantias, for it has twice as many. 

The umbrella of the Tlmumantias melanops is sub-orbicular, inflated, very tender, 



4G THAUMANTIAS GLOBOSA. 

transparent, and smooth. Around its margin are ranged in close order more than 200 
fine colourless tentacula, ringed and granulated, when highly magnified. At the base of each 
is a very small but very black and well-defined ocellus. Round the inner margin of the 
umbrella is a rather wide veil, which, instead of being borne horizontally in all the specimens 
I met, was lax and dependent. The sub-umbrella is hemispherical, and di\aded into four equal 
portions by the four radiating vessels, which traverse through the greater part of their courses 
four long, clavate, yellow, rather narrow, reproductive glands. I have represented an 
appearance presented by one of these glands, as if of a much contorted tube within it. This 
is probably of the male sex. From the centre of the sub-umbrella hangs the short and 
very broad stomach, opening by four large, triangular, fimbriated lips. It is usually pale, 
sometimes slightly tinged with yellow. 

The umbrella of this species was often more than half an inch in breadth, and of the 
same height. It has hitherto occurred only in the Zetland seas, and is not very common there. 

Plate X, f, 3, a, represents the entire animal magnified ; 3, h, the stomach and lips ; 
3, c, one of the reproductive glands, and some of the marginal tentacula (contracted), with 
their ocelli ; 3, d, one of the tentacula magnified when in extension. 

7. Thaumantias globosa, Forbes. 
Plate X, Fig. 4. 

Umbrella globular, smooth, transparent, colourless, delicate. Margin with a rather close- 
set fringe of tentacula. These are tinted with purplish-yellow, and when magnified, present 
a ringed and granulated aspect. They are highly contractile, and very slender. They spring 
from reniform tubercles of a pale yellow colour, with a crescentic ocellus formed of tawny 
pigment-cells, inclosing a cavity in which a vibrating mass of otolites is plainly seen. The 
tentacular bulbs are very large in proportion to the diameter of the tentacles. The sub- 
umbrella is small as compared with the body ; it is intersected by the four radiating vessels, 
which traverse in that part of their course nearest the margin four lax, more or less reniform, 
reproductive glands, firmer and more defined in form, being ovate, in the females. They are 
pale yellow in the males, tinged with tawTiy in the females. The gastric vessels present a 
knob-like enlargement at the point of their union with the marginal vessels. Here and there 
among the tentacles are little colourless tubercles studding the margin. The number of the 
former in a large specimen was 7x4-f-4 ; in a small one 3x4+4. They evidently increase 
with age. The stomach is very short, of a pale yellow colour, and bordered by four lanceolate 
furbelowedlips. The umbrella, in well-grown specimens, measures about half an inch across. 

This delicate species is very abundant in the harbours of both sides of the Zetland Isles, 
usually in company with Tliauviantias hemisphcerica and T. pilosella. It has a remarkable 
habit of crumpling up, as it were, its tentacula into a confused mass. When very young, as 
represented at fig. 4, c, of plate X, it often extends its tentacles to a great length, and the 
reproductive glands appear of disproportionate size. Plate X, fig. 4, a and b, represent its adult 
state, magnified ; 4, d, is the stomach and lips ; 4, e, the reproductive gland of a male ; 4,_/', 
an ovary full of eggs ; 4, g, a quarter of the margin, with the tentacular bulbs ; 4, h (marked 
e by mistake in the plate), a tentacle-bulb greatly magnified, showing the ocellus, the otolitic 
capsule, and the structure of the tentacle. 



THAUMANTIAS PILEATA. 47 

8. Thaumantias convexa, Forbes. 
Plate XI, Fig. 6. 

Umbrella very convex, but not globose, smooth, transparent, colourless. Margin fringed 
with twenty (4x4-}-4) ringed and granulated colourless tentacula, springing from as many 
tubercles, each pinkish, with a small red ocellus. Between each pair of tentacular bulbs a 
smaller coloured tubercle, without any tentacle, is placed. The sub-umbrella is large, and very 
convex. The radiating vessels which divide it, run in the course of four ovate, or rather 
paddle-shaped, yellow ovaries, placed very near the margin. The stomach is short and 
narrow, yellow, and terminated by four lanceolate yellow lips, the margins of which are 
slightly fimbriated. The height of the umbrella is about one fourth of an inch. This 
TJiaumantias is a very common species in the Zetland seas, and among the Hebrides. We 
have taken it at Oban. It is an active little animal, and very tenacious of life. 

Plate XI, fig. 6, a and b, represent it as seen in profile, and from above ; 6, c, is an 
ovary and base of a tentacle ; 6, d, part of the margin with tentacula. 

9. Thaumantias gibbosa, Forbes. 
Plate XI, Fig. 3. 



o' 



This curious and very distinct species, which I have hitherto only met with once, has an 
oblong conical umbrella, not contracted below, smooth, and colourless. The margin bears 
twenty-eight (6x4-|-4) pink tentacula, which, in the specimen taken, were habitually borne 
coiled up. They spring from red ocellated tubercles. The sub-umbrella is very convex ; 
in the course of the four radiating vessels which intersect it, are as many long, linear, pale 
rose-coloured ovaries, reaching almost to the margin. From the summit of the sub-umbrella 
hangs a quadrangular, narrow, rather long stomach, opening by four triangular simple lips. 
The body is about a quarter of an inch in length. 

Thaumantias gibbosa was taken in the Hebrides. Circumstances at the time prevented 
such a minute examination of it as so curious a form deserved. It cannot, however, be 
mistaken for any other species described in this work or elsewhere, and I hope some future 
observer will seek for and re-examine it. 

Plate XI, f. 3, a, represents the body in profile magnified ; 3, b, as seen from above ; 
3, c, two of the tentacula ; 3, cl, the stomach and mouth. 

ft Umbrella much depressed. 
10. Thaumantias pileata, Forbes (1841 ). 

Plate XI, Big. 2. 
E. Forbes, in Annals of Natural History, April, 1841, p. 84, pi. 1, fig. 3. 

Umbrella smooth, transparent, pellucid, shaped like a Chinese hat, being prominent and 
conical in the centre, depressed and extended at the circumference. Margin bordered by 
twenty (4 X4-f-4) filiform, colourless tentacles, springing from ocellated bulbs, coloured black 



48 THAUMANTIAS SARNICA. 

and yellow. Stomach short, with proportionally large, triangular, sharp lips. Ovaries 
oblong, yellow, placed rather more than half way down the pileated sub-umbrella. Breadth 
nearly an inch. 

This pretty species was taken by Mr. Smith and myself, at Port Rush, on the north coast 
of Ireland, in June, 1839. 

Plate XI, fig. 2, a, represents it of the usual size ; 2, h, magnified, as seen in profile ; ' 
2, c, as seen from above ; 2, e, an ovary ; and 2, d, the stomach and lips. 

11. Thaumantias lineata^ Forbes. 
Plate XI, Fig. 1. 

This is rather a large species of its division, measuring nearly an inch across the umbrella, 
which is much depressed, sub-hemispherical, smooth, colourless, and tender. The margin 
bears thirty-six (8 X4-|-4) filiform, transparent tentacles, springing from as many very small 
yellowish tubercles, with minute red ocelli. The sub-umbrella is elevated, as compared with 
the umbrella. It is bounded below by a rather broad veil, and divided by four radiating 
vessels ; in the second third of the length of these are seen the linear, yellow, reproductive 
glands. The stomach is quadrangular, small, and very short, campanulate, with four fim- 
briated lips. 

Taken in the Zetland seas in 1846, but not found common. 

Plate XI, f. \, a, represents it as seen in profile of the natural size; 1, b, magnified, and 
viewed from above; 1 c, an ovary; 1, d, Xm'o of the tentacles at their origins, with their 
bulbs. 

12. Thaumantias Sarnica, Forbes (1841). 

Plate XI, Fig. 4. 
E. Forbes, in Annals of Natural History, April, 1841, p. 84, pi. 1, fig. 6. 

Umbrella hemispherical, regidarly convex, smooth, transparent, and colourless ; margin 
with twenty (4x4-|-4) transparent tentacles, with colourless bases, which do not present 
conspicuous eyes. Sub-umbrella rather convex, divided by the four radiating vessels. In 
the lower half of the course of the latter, occupying nearly one half their length, are the linear 
or slightly clavate, bluish, reproductive glands. The stomach is of the same hue, and is small, 
quadrangular, with four, rather large, simple, triangular lips. Breadth about half an inch, 
or rather more. 

I took this apparently distinct species between Guernsey and the Island of Herm, 
in the autumn of 1839. It is very closely allied to Thaumantias pileata — perhaps too 
closely. 

Plate XI, fig. 4, a and b, represents it as seen from one side, and from above ; 4, c, is 
the stomach and lips ; 4, d, an ovary. 



THAUMANTIAS HEMISPH^RICA. 49 

13. Thaumantias T/iompsoni, Forbes (1841). 
Plate XI, Fig. 5. 
E. Forbes, in Annals of Natural History, April, 1841, p. 88, pi. 1, f. 6. 

Among the most phosphorescent of the smaller Medusae on the coast of Cornwall, and 
elsewhere in the south, is a little Thaumantias, which several years ago (in 1840) was met 
•witli, for the first time, by Mr. W. Thompson, Mr. R. Ball, and myself, in Chfden and 
Roundstone Bays, on the coast of Connemara, in Ireland. When irritated, it gives out a 
brilliant light from the bulbs of its tentacula. 

The umbrella of Thaumantias Thompsoni is hemispherical and rather depressed, smooth 
and colourless; It measures about a quarter of an inch across. Round its margin are sixteen 
(3x4+4) white tentacula, springing from rather large yellow tubercles, each bearing a very 
small, dark red or nearly black ocellus. These tentacles, when highly magnified, appear 
annulated, and their bulbs distinctly exhibit an ocellus separate from an otolitic mass. The 
sub-umbrella is rather more depressed than the umbrella. On its sides, rather more than 
half way down, are the four reproductive glands, ovate, and of a clear yellow colour. In the 
centre is the small, rather slender, quadrangular stomach, also yellow ; it terminates in four 
triangular lips, margined with a thick band of fibrous cells. 

Plate XI, f. 5, a and b, represents this species magnified ; 5, c, one of the ovaries full of 
egg, and the bulb of a tentacle, greatly magnified ; 5, d, part of the margin with three ten- 
tacula ; 5, e, a lip, with its border of fibrous cells. 

14. Thaumantias hemisphcerica (sp.), Miiller (1776), 

Plate VIII, Fig. 2. 

Synonyms. Medusa Iwrnispheerica. Gronovius, Acta Helv., 438, t. iv, f. 37 (1760). 

O. F. MiUler, Prod. Faun. Dan., No. 2822 (1776). 

O. F. ;Muller, Zool. Dan., pi. 7, f. 1-4 (1788), 

(fig. Cop. in Enc. Meth., pi. 93, f. 8-10.) 
Lin., Syst. Nat. Cur. Gmelin, p. 1098 (1 789). 

Modeer, Kong. Vet. Ac. Nya Kand. vol, xii, 

p. 251 (1791.) 
Oceania hemispheerica. Peron, Ann. de Mus., vol. xiv, p. 347 (1809). 
Medusa hemispheric a, var. lucida. Macartney, Phil. Trans. (1810), 

p. 264, pi. 14, f. 3. 
Geryonia hemispharica. Fleming, Brit. Animals, p. 500 (1828). 
Thaumantias hemisphcerica. Eschscholtz, Syst. der Acal., p. 103 (1829). 

Blainville, Man. d'Act., p. 285 (1834). 

(Lamarck), An. sans Vert., 2d Ed., 

p. 162 (1840). 
Medusa [Geryonia) hemisphcerica. Thompson, Annals of Nat. Hist. 

vol. v, p. 248 (1840). 
Thaumantias hemisphcerica. Lesson, Acal., p. 335 (1843) 

7 



50 THAUMANTIAS HEMISPH.^RICA. 

It is very probable that, under the name of " Medusa hemisph(erica" the older and most 
of the more recent writers on Acalephce confounded many, or at least more than one, species 
of Thaumantias ; and, as very few of their notices extend to more than characters obvious at 
first glance, and common to a majority of members of the genus, it is difficult or impossible to 
ascertain what form or forms were meant when the name in question was cited. The figures 
given by Midler and GronoA'ius were evidently at such times borne in mind ; but, as 
the importance of ascertaining the number and structure of the bulbous bases of the 
tentacula was not understood by any of the naturalists from whom I have given citations, 
unless Peron and Lesson be excepted, the reference to those figures cannot be received 
without suspicion. 

I feel sure that the animal I am about to describe and figure as Thaumantias 
hemisphcerica, is identical with that which was delineated by Miiller in the Zoologia Danica, 
and clearly characterised by Peron, whose description, which I here cite, appHes to no other, and 
was probably drawn up after a study of MiUler's account and figure, for the French naturalist 
gives no other locality than " des cotes de Danemarck." He characterises the species thus : 
" un ombrelle hemispherique, deprime a son centre ; ovaires pedicelles et claviformes; rebord 
entier, garni de trente-deux tentacules tres-courts et de trente-deux petites glandes ; ombrelle 
gris-bleuatre, parseme de petits points plus gris ; ovaires jaunatres, glandes marginales rouge ; 
1 centimetre." The short diagnosis of Eschscholtz, "canalibus versus marginem disci 
clavatis," founded on his misapprehension of the nature of the reproductive organs, would 
apply equally well to half the allied species here described, and his account of the number of 
marginal tentacles, " ihr Rand ist mit 16 bis 24 kurzen Fangfaden besetzt,'' leaves us in no 
more certain position. The tentacles, however, as I have found by examination of very 
numerous specimens, do vary greatly in the several stages of the animal's growth, though I 
have never seen them fewer than twenty. When full grown, the number is, as Peron has 
stated, thirty-six. This variation of number of tentacles with age is seen in several species of 
Thaumantias, especially in those with depressed umbrellas. Many species, however, present 
exactly the same number in their earliest and their oldest stages. Fortunately, the large 
conspicuous red or orange ocelli, and brightly coloured claviform reproductive glands, afford 
features combined with form, which will always enable us to recognise the true lliaumantias 
hemisphesrica without much difficidty. 

The individuals which I have examined in Zetland, where they abound in the bays and 
harbours, have a hemispheric, slightly depressed, transparent, smooth umbrella, sometimes 
measuring nearly three fourths of an inch across. The margin in adult specimens usually 
bears thirty-two (sometimes more) tentacles, springing from as many large tubercle-like ocelli, 
which are vividly coloured with orange and red, and when magnified, are seen to present a 
small black dot. The formula for the oceUi and tentacles in the adult is probably 7x4+4. 
The tentacles are composed of granular tissue ; they are often carried short, not by contrac- 
tion, but by coiling up in a spiral. The sub-umbrella is moderately convex, and divided 
by the four gastric vessels, which pass in the lower half of their courses through four 
linear, claviform, purplish or yellowish ovaries, marked with purple or orange lines. The 
stomach is short, rather broad, purplish, or tinged with pink, terminating in four lanceolate 
fimbriated lips. 

The abundance of individuals of this species in all stages of growth enabled me, when 



THAUMANTIAS HEMISPH^RICA. 51 

in the Zetland Isles, to observe the mode of formation of their tentacula, a point of considerable 
- physiological importance. The results may be summed up as follows : — 

1st stage. The first indication of the tentacle is in the form of a minute lobe at the 
margin of the body, in the immediate neighbourhood, and in contact with the motor band and 
marginal vessel. 

2d. This lobe becomes a closed cell, which then rapidly enlarges in every direction, but 
chiefly superiorly and inferiorly. 

3d. It next presents a contraction, which is the commencement of its division into two 
cells, the superior being largest. 

4th. The inferior cell grows rapidly, and divides itself into two, the lowermost of which 
does the same, and this process is continued until the tentacle has elongated considerably in 
one direction. In the mean time the superior vesicle of all enlarges, but does not divide. All 
present the aspect of nucleated cells. 

5th. The partitions between the cells begin to be lined with minute, translucent granules, 
which give the tentacula a raoniliform aspect ; and from this time those organs become con- 
tractile. 

Gth. The minute translucent granules (which are themselves cells of a secondary order) 
multiply until they fill up the primary cells, and give the tentacula the aspect of homogeneous 
bodies, composed of highly contractile granular tissue. 

7th. When the tentacle has far advanced towards its perfect state, the bulb-like vesicle 
at its base also begins to be filled with granular tissue, many of the minute cells of which 
become pigment-cells. 

8th. In the middle of this mass of cells, one cell appears larger than the last, and which, 
increasing with great rapidit}', divides in the end the mass of minute secondary cells into two 
pad-like bodies, constituting together the conspicuous ocellus. 

9tli. When this division has taken place, there appears in the central cell a new and free 
mass of minute cells, in which are secreted crystals (of carbonate of lime). These are the 
otolites. The tentacle is now complete. 

The Thaumanlias hemisphccrica is an active little animal, gregarious. It is exceedingly 
tenacious of life, as the tortures to which it was submitted by Dr. Macartney, quoted in my 
mtroductory remarks,, show. It is very abundant in the Zetland seas, and occurs also in the 
east and west sides of the mainland of Scotland. It is also, according to Mr. W. Thompson, 
found on the north coast of Ireland. " In October, 1838, 1 obtained one of these Medusae in 
Belfast Bay, and a day or two afterwards many specimens were brought to me by Mr. 
Hyndman from the same locality ; in size they rather exceed Miiller's, measuring five lines in 
diameter in their most depressed state. Mr. R. Patterson informs me that he obtained the 
M. hemisphcBrica at Larne, in the summer of 1835." 

Plate VIII, f. 2, «, represents this species seen in profile, magnified ; 2, h, seen from 
aljove, a very large specimen (too many tentacles and tubercles are introduced in the figure, 
by mistake) ; 2, c, is one of the reproductive glands ; 2, d, those of the tentacles and their 
bulbs. 



52 THAUMANTIAS LUCIFEEA. 

1 5. Thaumantias inconspicua, Forbes. 
Plate VIII, Fig. 3. 

A very delicate, and rather large species, measuring usually about three fourths of an 
inch across. Its umbrella is hemispherical, smooth, and colourless. The margin bears from 
sixteen to twenty colourless tentacles springing from pale yellow, inconspicuous bulbs, each 
tinged with a faint tawny spot. Between each pair there is a rudimentary marginal tubercle. 
The sub-umbrella is much more depressed than the umbrella ; its opening is bordered by a 
rather broad veil. The ovaries are long and linear, and of a faint lilac or greenish hue, 
with a central fulvous line. They occupy more than half the course of each radiating vessel. 
The stomach is narrow, quadrangular, and of a yellow colour. It terminates in four lan- 
ceolate lips. The inconspicuous ocelli and pale reproductive glands easily distinguish this 
species from its congeners. It is common in the Hebrides. 

Plate VIII, fig. 3, a and b, represent it magnified as seen from the side, and from above; 
3, c, is an ovary ; 3, d, a portion of the margin, showing the tentacular bulbs, and the inter- 
mediate tubercles. 

16. Thaumantias lucifera, Forbes. 

Plate X, Fig. 2 (under the name of T. lucida). 

This minute and curious Medusa is one of the most phosphorescent of all the naked-eyed 
species. Small as it is, I have not seen it more than one fifth of an inch across ; it presents 
such very distinct characters, that I do not hesitate to describe and name it as a separate 
species, although specimens presenting truly adult characters have not as yet occurred. 

The umbrella is very much depressed, smooth, and transparent. The margin is fringed 
with a close-set series of tentacula, of which I reckoned no fewer than eighty-four (20x4+4) 
in several examples, but very small specimens had rather fewer of these organs. Their bases 
present a very minute ocellus, and there is a club-shaped, transparent organ, projecting, as it 
■were, into the umbrella above it. This is probably the auditory vesicle in an early stage of 
development. The structure of the tentacles also indicates an immature condition of the 
tissues. The sub-umbrella is rather high in proportion to the umbrella. On its lower half 
are the four reproductive glands, sub-orbicular, lax, and yellow. The stomach is rather large, 
quadrangular, yellow, with imperfectly developed lips, whose margins are not fimbriated. 

I have met with this species in Zetland and the Hebrides, and in vast abundance off the 
Lizard Point, on the coast of Cornwall ; also at Dartmouth, and in the west bay of Portland. 
When swimming, it carries its tentacula stiffly, and at nearly a right angle with the body. 
The Thaumantias plana of Sars (Beskrivelser og Jagtagelser, p. 28, pi. 5, fig. 13) bears a 
close resemblance to it, and is of the same size, but the tentacula are described as being more 
than one hundred in number. It is also evidently an immature species. The only other 
described form of Thaumantias with a depressed umbrella, and very numerous tentacles, is 
the T. multicirrata of Sars (Op. cit. p. 26, pi. 5, fig. 12), but that has linear or clavate 
ovaries, more than 200 tentacles, and very few and distinct ocelli in proportion to the number 



SLABBERIA HALTERATA. 53 

of tentacula. Both these species of Sars are from the coast of Norway, and will very pro- 
bably be found hereafter in Zetland and on the coast of Scotland. 

Plate X, fig. 2, a, represents the Thaumantius lucifera of the natural size ; 2, h, as seen 
from above ; and 2, c, in profile, much magnified ; 2, e, the stomach ; and 2,y, its mouth ; 
2, g, one of the tentacles with its bulb and vesicle ; 2, d, an ovary. 

The name Thaumantias liickla has been adopted by Lesson for Macartney's animal, 
under the impression that the latter was intended to be distinct from T. hemisphcBrica. 

17. Thaumantias punctata, Forbes (1841). 

Plate X, Fig. 1. 

E. Forbes, in Annals of Natural History, vol. vii, p. 85, pi. 1, fig. 5 (1841). 

The last of the species of TJiaumantias which I have to describe, is one of which I gave 
an account some years ago in the 'Annals of Natural History,' under the name of Thaumantias 
punctata. 

The umbrella is hemispherical, and rather depressed, smooth, and colourless. Its margin 
is bordered by thirty-two filiform, colourless tentacula, each with a rather swollen bulbous 
base, marked by a dark, almost black, ocellus. The sub-umbrella is convex, and divided into 
four parts by the radiating vessels which traverse, on its lower half, four short, linear, purplish 
ovaries. The stomach is very small, and purplish. It terminates in four short lips. This is 
a large species, measuring nearly an inch in diameter. It occurred abundantly off the Isle of 
Man, where it was taken in the month of June, 1839, by Professor Goodsir, Mr. Henry 
Goodsir, and myself. I regret not having encountered it since, as it requires to be submitted 
to more minute observation than time then permitted. 

Plate X, f. 1, a and b, represents this species magnified, in profile, and seen from above ; 
c, is the stomach ; and d, a tentacle with its bulb. 



Genus XII. Slabberia, Forbes (1846). 

Umbrella campanulate ; ovaries four, linear, in the course of the four simple, 
gastro-vascular canals ; peduncle proboscidiform, highly extensile, oral orifice 
circular ; a marginal tentacle springing from an ocellated bulb, and terminating 
in a coloured globular body, placed opposite each of the gastro-vascular canals. 

Slabberia halterata, Forbes. 

Plate VI, Fig. 1. 

Umbrella deeply campanulate, smooth, colourless ; sub-umbrella large, divided into equal 
parts by four simple vessels, which open into a circular marginal vessel. On the upper third 
of the sub-umbrella are seen in the course of the vessels four small linear ovaries or reproductive 
glands, pointed at each end. The border of the general cavity is provided with a shelf-like 
veil. The tentacula are strong, four in number, and colourless, except at their bases and lips. 



54 SLABBERIA HALTERATA. 

The bulbs at tlieir bases are more or less triangular, coloured above with bright verdigris 
green, and across the centre with a band of deep orange, below which, on the root as it were 
of the tentacle, is seen a conspicuous and rather large jet-black ocellus. The extremity of 
each tentacle is likewise swollen into a bulb, which is of a rich orange hue. From the centre 
of the sub-umbrella hangs a long and highly extensile peduncle or stomach, capable of being 
contracted entirely within the general cavity, but more usually elongated beyond the length of 
the tentacula. It is of a denser tissue than the other parts, and terminates in a circular 
orifice. The termination is not unfrequently swollen into the shape of a bell. The summit 
projects slightly above the surface of the sub-umbrella. The diameter of the disk does not 
exceed one eighth of an inch. 

This curious Medusa was observed in August, 1836, in Mount's Bay, Cornwall, where 
great numbers of them were taken. It swims with its four tentacles either stretched out 
straight, or at a slight angle, and as if quite stiff, so that they seem with their loaded extremities 
to serve as poisers. The brilliant terminal bulbs, following at equal distances the conspicuous 
ocellated bulbs of their bases, give the creature a very striking aspect, and at first glance 
seem as if they belonged to two Medusae, one of which had partially enveloped the other. 
The Slabheria halterata is a very active, and apparently hardy Uttle animal. This remarkable 
combination of characters, and the features quite peculiar to itself, render it well worthy of 
o-eneric distinction. The position and form of its ovaries indicate a relationship with Thau- 
vHintias, whilst the pedimcle is that of a Sarsia. It thus links together genera, which, were 
it not for such a connecting form, would seem to be far apart. I have dedicated the genus to 
Martin Slabber, an ingenious Dutchman, who amused himself with the microscope, and 
pubhshed an account of his observations at Haarlem, in 1778. He was one of the first to 
direct attention to the minuter forms of Medu,Sce inhabiting the German Ocean, and therefore 
has good claim to preside over a sound generic group. 

Plate VI, f. 1, a, represents Shihheria halterata of the natural size ; 1, ^, magnified; 
1, c, disposition of vessels and ovaries, as seen from above; 1, (J, ovary; \,e, a tentacle 
with its bulbs ; 1,/, proboscis when most dilated. 



SARSIAD.E. 

Genus XIII. Sarsia, Lesson (1843). 

Umbrella hemispherical ; radiating vessels four, simple; no conspicuous ovaries , 
four marginal tentacles opposite the point at which the radiating vessel joins the 
marginal one ; ocelli four, more or less conspicuous ; stomach in a very extensile, 
cylindrical, proboscidiform peduncle, with a simple orifice. 

This genus was instituted by Lesson for a very remarkable Medusa discovered by the 
eminent naturalist of Norway, whose name it bears — a philosopher who, pursuing his re- 
searches far away from the world, buried among the grand solitudes of his magnificent 
country, where the pursuit of science is his recreation, and the holy offices of religion his 
sacred duty, has nevertheless gained name and fame wherever the study of nature is followed. 
The unpretending writings of this parish priest have become models for the essays of learned 
professors in foreign lands, and his discoveries the texts of long commentaries by experienced 
physiologists. The French naturalist, in seeking to be the first to honour the name of Sars, 
has done himself honour by his recognition of high and modest merit. With peculiar 
pleasure, therefore, do I offer a testimony to the value and permanency of the genus before 
us, by announcing three very distinct species additional to the only recorded type, con- 
firmatory of the excellence of its constitution. 

1. Sarsia tubulosa (sp.), Sars (1835). 

Plate VI, Fig. 2. 

Synonyms. Oceania tubulosa. Sars, Besk. og. Jagt., p. 25, pi. 5, fig. 11 (1835). 

Thompson, Ann. N. H., vol. v, p. 249 (1840). 

Dujardin, in 2d Ed., Lamarck, An. s. Vert., t. iii, 

p. 165 (1840). 
Sarsia tubulosa. Lesson, Acaleph., p. 333 (1843). 

This is the type of the genus. The umbrella, which is sometimes nearly an inch in 
length, is smooth, colourless, and of an oblong hemispherical shape. Its margin bears four 
equidistant tubercles, of a bluish or purplish colour, each marked with a well-defined dark 
ocellus. From each tubercle arises a tentacle, lilac, or greenish, or blue, e-\tremely extensile, 
often elongated to three times the length of the body. The structure when magnified is seen 
to be moniliform and granular. The sub-umbrella is hemispheric and very convex, divided 
into four equal parts by as many radiating simple vessels, which join the marginal vessel 



5G SARSIA TUBULOSA. 

opposite the bases of the tentacula. The margin of the sub-umbrella is bounded by a rather 
broad membranous veil. From its centre hangs the long, cylindrical, fleshy, proboscidiform 
peduncle, of a blue or lilac, sometimes greenish colour, and fleshy substance. It is highly 
contractile, being often extended, like a long tube, so as to be twice the length of the body ; 
sometimes contracted within the bell, and then assuming a bottle shape ; the central part 
being inflated, and suspended by a slender portion formed by its base. Its fixed point rises 
conically for a little way above the summit of the sub-umbrella. The free extremity is more 
or less claviform, and terminates in a round or indistinctly four-lobed orifice, the lips of which 
are sometimes everted. The vessels do not appear to run down the peduncle, the interior of 
which is occupied throughout by the stomach. 

In a note, communicated by my friend Mr. Patterson, he describes the tentacular bulbs, 
in some specimens met by him at Larne, as exhibiting brilliant crimson ocelli, and the central 
peduncle as of a dirty pinkish yellow, with a bright crimson spot at its junction with the body. 
" The peduncle," he writes, " was ever changing its form ; sometimes flung out beyond the 
body, and then becoming proportionally thin." The animals were from a quarter to three eighths 
of an inch in length, and other specimens afterwards taken measured two, five, and seven lines 
in the body. The tentacula extended so much, that on one occasion they measured two inches 
and a half, by a rule applied to the sides of the glass jar in which they were confined. 

In his " Additions to the Fauna of Ireland," published in the ' Annals of Natural History' 
for 1840, Mr. W. Thompson has the following note respecting this species, in which it was 
first announced as a British animal, though, as I have remarked before, when describing 
Oceania turrita, it is probable that the PeUscelotus vitreus of Templeton, figured in the 
ninth volume of Loudon's ' Magazine of Natural History,' was our Sarsia turned inside out. 

"April 11, 1840. I had the satisfaction to-day of identifying with the Oceania {?) tuhulosa 
of Sars a Medusa of which several individuals were brought to me by Mr. Hyndman, just 
after their capture in Belfast Bay. On calling the attention of Mr. R. Patterson to them, a 
reference to his notes on Medusse showed that he had procured the same species at Larne, 
county Antrim, in May 1835, and June 1838; and again at Bangor, county Down, in July 
1839. As my friend could not find the species described — Sars's work he had not, for 
reference — he drew up a detailed and interesting account of the animal, accompanied by several 
characteristic sketches of it in various positions. Having remarked that one of my specimens, 
which was in a phial containing one ounce and a half of sea-water, appeared as lively after 
four days' captivity as at first, although the fluid had not been changed, nor any nutriment 
added, I, before leaving home for some days, handed it over to Mr. Patterson, that 
the period the animal would live, under such circumstances, might be noted. From him I 
learn that this individual lived thus for twelve days (from the 18th to the 30th of April) and 
that for the first ten, it retained its ordinary vivacity." (Thompson, loc. cit., p. 249.) 

The first time I had the pleasure of seeing this elegant little Medusa, was at Scalloway, 
in Zetland, where it was taken by Professor Goodsir and myself in 1839. During the month 
of June in the same year, I met with it again abundantly when dredging with Mr. Smith of 
Jordanhill, in the Kyles of Bute. On visiting Zetland in 1845, in company with Mr. 
Mac Andrew, we found it very abundant in the bays and harbours on both east and west coasts. 
Some we took at Hillswick were exceedingly lively and active, swimming obliquely through the 
M'atcr with great rapidity. Being kept in a jar of salt water with small Crustacea, they 



SARSIA GEMMIFERA. 57 

devoured these animals, so much more highly organized than themselves, voraciously ; appa- 
rently enjoying the destruction of the unfortunate members of the upper classes with a truly 
democratic relish. One of them even attacked and commenced the swallowing of a Lizxia 
octopunctatu, quite as good a Medusa as itself. An animal which can pout out its mouth 
twice the length of its body, and stretch its stomach to corresponding dimensions, must indeed 
be " a triton among the minnows," and a very terrific one too. Yet is this ferocious creature 
one of the most delicate and graceful of the inhabitants of the ocean — a very model of ten- 
derness and elegance. 

Plate VI, fig. 2, «, represents the Sarsia tubulosa of natural size ; 2, b, magnified ; 
2, c, a tentacle, bulb, and ocellus ; 2, d, the peduncle retracted and inflated. 

2. Sarsia pulchella^ Forbes. 

Plate VI, Fig. 3. 

A much smaller species than the last ; none of the specimens which I have met exceeding 
a quarter of an inch in length of body. The umbrella is sub-orbicular, and very convex, 
transparent, colourless, and smooth. The margin is quadrate, each angle bearing a large 
ocellated tubercle, from which a rather thick tentacle springs. The tentacle is of a pale pink 
colour ; the tubercle at its base transparent, with a mass of pink or orange pigment-cells at 
its upper part, from which depends a brilliant green pedicle, with a jet-black ocellus at its 
extremity. The sub-umbrella is prominent, and rather conic. Down its sides run the four 
gastric vessels, coloured pale pink. Round its opening is a four-lobed veil. From its centre 
hangs the proboscidiform peduncle, which is rarely protruded beyond the umbrella, and more 
frequently contracted into various flask-like shapes. Its point of afiixment rises as a short 
pink cone above the sub-umbrella ; its orifice is round. It is of a brownish-red colour, with 
a green-tinged oral extremity. 

It is a very active animal, and very tenacious of life. It never extends its tentacula so 
far as the preceding species, and often carries them coiled up spirally. I found several 
specimens in Brassay Sound, Zetland, in 1845. 

Plate VI, fig. 3, a, represents this species of the natural size ; 3, b, magnified ; 3, c, the 
bulb of a tentacle ; 3, d, a tentacle coiled up. 

3. Sarsia gemmifera, Forbes. 

Plate VII, Fig. 2. 

The very remarkable animal which I have now to describe, was discovered in the Zetland 
seas by Mr. M'Andrew and myself in 184.5 ; several specimens were taken. It was the first 
Sarsia which we found exhibiting a distinct mode of reproduction, and that by gemmation 
from the walls of the peduncle. 

The Sarsia gemmifera is a very small species, scarcely a quarter of an inch in length of 
body. Its umbrella is pyriform, smooth, and colourless. The aperture of it is rather con- 
tracted and quadrangular. At each angle there is a conspicuous ocellated tubercle of a pear 
shape, its upper part pale tawny, its middle dark orange, and its base colourless, with a well- 
defined, jet-black ocellus. Round the orifice of the sub-umbrella, which is pyriform, is a four- 

8 



58 SARSIA GEMMIFERA. 

lobed veil, festooning, as it were, the spaces between the ocellated tubercles. From each of 
the latter arises a tentacle, rather short, thick, cylindrical, moniliformly granulated, and of an 
orange colour. Each tentacle is placed opposite the point at which one of the four gastric 
vessels joins the marginal one. From the centre of the sub-umbrella is suspended the 
peduncle, perforating its summit, and terminating there in a small conical process of granular 
tissue. The peduncle is shorter than the umbrella, but may be extended slightly beyond it. 
It is slender, cyhndrical, and tubular, but is capable of changing form greatly, and often 
swells out into a club- or bottle-shaped extremity, ending in the mouth, which is round. The 
peduncle is of an orange colour, like the tentacles, but much paler. 

The peduncle presented the appearance of being ramified, or rather pinnated, variously- 
shaped processes projecting from the sides. When several individuals were compared, it was 
found that these pinnations did not correspond, and a closer inquiry made it evident that they 
were in reality young individuals, in various stages of development, budding from the pedun- 
cular tissue. They are not distributed over its surface in any regular order according to their 
decrree of advancement, but intermingled, as may be seen in the much magnified representation 
of the peduncle in an individual, different from that which is drawn entire. (See 2, e, and 
compare it with 2, b) At the same time there is an indistinct spiral arrangement to be 
observed, and the peduncle has a tendency to assume angular bendings at the points from 
which the buds spring. The earliest stages of one of these buds is that which I have 
represented at 2, /, where the tissue of the surface of the peduncle simply bulges out as a 
small wart, whilst there is a corresponding indentation in the tubular cavity beneath. This 
indentation increases with the growth of the wart, which takes upon itself a club shape, and, 
at the same time, there is a notch-like appearance towards the base at the upper part of the 
club (2, g). In a stage more advanced, the club begins to assume a globular form, and the 
excavation, if anything, decreases, not projecting beyond the notch, but a new cavity has 
appeared independently within the club, and has possibly been formed by the division of the 
old cavity into two parts, and the isolation of its upper part (2, //). In the stage represented 
at 2, i, the development has advanced rapidly ; the club begins to present a four-lobed aspect, 
and the internal cavity has greatly enlarged. An assemblage of small dots, indicative of an 
ocellus, appears opposite each lobe. After this it would appear, that correspondent with the 
increase of the lobes, is the opening of the cavity and the formation of a peduncle ; for, in the 
highest condition of these gemmse which I have had an opportunity of examining, the bud 
presents the appearance of a little bell (3, Ic), open, and having a rudimentary proboscis 
within it ; the lobes are much more elongated, and at their bases are seen not only the little 
ocelli present in the last stage described, but others below them of a jet-black colour. It is 
evident that the former are the orange portions of the tentacular bulbs, and the latter, the 
black ocelli beneath them, whilst the lobes, which have already become of a conspicuous 
orange colour (as well as the peduncle), are the tentacles in course of development. The 
four gastric vessels are also now manifest, so that we have a young Sarsia nearly ready to 
drop from its parent, and shift for itself. 

Plate VII, fig. 2, a, represents this interesting animal of the natural size ; 2, h, the same, 
magnified ; 2, c, the summit of the peduncle ; 2, rf, a tentacular bulb with its ocelli ; 2, e, the 
peduncle of another individual, with its bulbs ; and 2,/ to 2, k, the various stages of develop- 
ment of the buds as just described. 



SARSIA PROLIFERA. 59 

4. Sarsia prob'fera, Forbes. 
Plate VII, Fig. 3. 

Equally remarkable with the Medusa I have just described is that now before us — indeed 
more so, for the last was an instance of gemmation from the peduncle, a phenomenon, as we 
shall hereafter see, pi-eviously discovered by Sars in another genus (Lixzia) ; but in Sarsia 
prolifefu we have gemmation from the tentacular bulbs, an entirely new and most remarkable 
mode of reproduction. 

On the 21st of August, 1846, we found great numbers of minute Mcdusse in Penzance 
Bay. Among them was one which, whilst it presented the simple cylindrical proboscis and 
four tentacula of a Sarsia, differed from all the members of that genus hitherto seen, in having 
at the base of each tentacle a supplementary bulb, or a bunch of little tubercles suspended 
like a bunch of grapes. Fortunately, individuals of this curious little creature — it is even less 
than the last species — were plentiful, so that we were soon enabled to unriddle the anomaly 
by a careful e.xamination of numerous specimens under the microscope. The supplementaiy 
bulbs and grape-like tubercles proved to be young Sarsia, sprouting by gemmation from the 
bases of the tentacula. 

The Sarsia prolifera is a very delicate little animal, so faintly coloured as to be incon- 
spicuous in the water, in that respect differing from the other members of its genus. 
Its umbrella is campanulate, and somewhat inclined to a sub-globose form, smooth, and 
colourless. 

The opening of it is quadrangular, the spaces between the angles curtained by a veil, the 
angles themselves bearing each a pale yellow tentacular bulb, marked with a minute black 
ocellus. From the four bulbs spring as many pale yellow, moniliformly-granulated, slender, 
coiling tentacula. The sub-umbrella occupies about two thirds of the length of the umbrella, 
and to about a third of its length is suspended the pale yellow tubular peduncle, which is very 
changeable in form, sometimes inflating itself into a bottle-shape, but apparently never 
protruded beyond the umbrella. Its summit projects slightly above the sub-umbrella ; its 
orifice is round, and bounded by a highly contractile rim of fibrous cells. 

In every specimen we found a different arrangement or degree of development of the 
little buds at the bases of the tentacula ; and not only did each individual differ from the 
other, but rarely were the arrangements of the germs at the bases of the four tentacles alike 
in the same example. I have accordingly figured the four bunches of a single specimen to 
show how they differ, and what the nature of the curious bud-like bodies is. 

In fig. 3, e, all the buds are in a low state of advancement ; the bulging above the 
coloured ocellus indicates the lowest and most rudimentary stage, corresponding to the wart- 
like condition described in the account of the buds in the peduncle of Sarsia gemmifera. 
To the left of the ocellus, a bud more advanced, and exhibiting traces of an interior cavity, is 
seen, and dependent fi'om the bulb at its right side is a gemmule already presenting traces of 
lobation. In fig. 3, g, these two stages are repeated, with the advance in one that four 
dark masses of pigment-cells indicate the bases of the rudimentary tentacula, and the formation 
of ocelli. In fig. 3, h, whilst two of the gemmae are rudimentary, a third shows not only 
ocelli, but the tentacula distinctly in course of formation, as yet, however, folded in. In 3,/, 



60 SARSIA PROLIFERA. 

one of the gemmae has attained a stage of development far beyond that of the other children 
of its parent, for it has assumed a distinct campanulate form ; its sub-umbrella is lineated by 
the gastro-vascular canals ; its tentacular bulbs are defined, and separated from the margin, 
and present large and conspicuous ocelli, and its tentacles have assumed a definite form and 
ample dimensions. The whole hangs by a very short and slender peduncle to the tentacular 
bulb, which exhibits besides three other buds in early stages of development, ready to advance 
when the firstborn of the parent finger has loosened its ties, and embarked on a free voyage 
of its own. A youthful Sarsia, which has just cast oft its leading-strings, is represented at 
3, i. The funiculus still remains projecting from its summit, and its stomach is so rudi- 
mentary, that, for its own sake, we must wish it a rapid accumulation of new tissue, since at 
present it can scarcely hope to live very long, unless it provides itself with a more efficient 
receptacle for nourishment. Perhaps it was undutiful, and left its mother too soon ; all the 
worse for it, and the better for us, since we learn from its examination, that the little process at 
the summit of the sub-umbrella is the remains of the funiculus, and that the umbrella does 
not attain its true shape and dimensions till after the sub-umbrella has been formed, and the 
tentacula and organs of sense comparatively advanced. The great size of the ocelli in these 
young animals, as compared with the entire body, is very striking, especially in the species 
before us, where the ocellus is eventually very small and inconspicuous, though well defined. 
The order of formation of tissues and organs in the Sarsice seems to be as follows : — 1st, the 
motor tissue begins ; 2d, the ca\aty of the disk is outlined, though closed ; 3d, the pigment 
cells of the ocelli commence to appear ; 4tb, the cavity opens, and the tentacula grow as lobes ; 
5th, the vessels are formed, and the distinction between the tentacle-bulbs and tentacles 
appears ; at this time the veil between the tentacle-bulbs is indicated by lobes, and the 
peduncle appears ; and 6th, the mass of the cellular tissue of the umbrella is formed, and the 
peduncle completed after the bud has become free. 

What strange and wondrous changes ! Fancy an elephant with a number of little 
elephants sprouting from his shoulders and thighs, bunches of tusked monsters hanging 
epaulette-fashion from his flanks in every stage of advancement ! Here a young pachyderm 
almost amorphous, there one more advanced, but all ears and eyes ; on the right shoulder a 
youthful Chuny, with head, trunk, toes, no legs, and a shapeless body ; on the left an infant, 
better grown, and struggling to get away, but his tail not sufficiently organized as yet to 
permit of liberty and free action ! The comparison seems grotesque and absurd, but it really 
expresses what we have been describing as actually occurring among our naked-eyed Medusae. 
It is true that the latter are minute, but wonders are not the less wonderful for being packed 
into small compass. The multitude, being muddle-headed, love magnitude, but the philo- 
sopher does not estimate a whale above a minnow for his mere bigness. " Nosci digna base 
animalcula, non quia Deus maximus in minimis est, seque enim magnus in omnibus, at ob 
eximiam membrorum exilitatem, miram organorum diversitatem, varia Creatoris eundem finem 
obtinenda media et pulchritudinem et proportionem quam nihil excellit." So wrote Otho 
Frederic Midler — filled, by his studies of minute life, with a deep spirit of reverence and 
admiration of his monoculi ; so might we write of our Medusae. But when to all the wonders 
of their structure are added such surprising physiological facts as those which we have just 
been naiTating concerning their reproduction, the spirit of reverent astonishment fills us 
fuller and fuller. " La force qui developpe, rintelligence qui specific et co-ordonne, I'amour 



BOUGAINVILLEA. 61 

qui unit et vivifie"* — the triune powers manifested in each and every being, in each single 
and all combined, are revealed as clearly in our little Sarsia, as in the mightiest monster of 
the ocean, beneath whose shadow it may swim invisible to the unarmed eye. And when we 
behold how its perpetuity in that ocean is secured, we are tempted to explain with Spenser — 

" Wonder it is to see 

How diversly Love doth his pageaunts play. 

And shewes his powre in variable kinds."t 

Plate VII, fig. 3, a, represents Sarsia prolifera of the natural size ; 3, b, magnified ; 
3, c, its peduncle ; 3, d, a tentacle, with the ocellated bulb at its base, and a gemmule beside 
it ; 3, e, to 3, i, gemmae in various stages of development. 



Genus XIV. Bougainvillea, Lesson (1829). 
HiPPOCRENE, Mertens (1835). 

Umbrella spherical ; ovaries in the form of four equal lobes, on the sides of the 
short peduncle; margin of the umbrella, with four fasciculi of tentacular bulbs, 
sending forth one or many tentacles, each fasciculus opposite one of the four single 
radiating vessels ; stomach shorter than the sub-umbrella ; mouth with four rami- 
fying tentaculated lips. 

This very remarkable group was first strictly defined by Brandt from the drawings and 
notes of Mertens, who had recognised its generic value, and assigned a name to it in his 
manuscripts. Lesson, who appears to have been the first naturalist who observed any species 
belonging to it, had equally perceived its importance, and claims priority for the name which 
he gave, dedicating it to the honour of the distinguished French voyager. Admiral 
Bougainville. Both Lesson and Brandt appear to have met with the same species, the 
former, in the bays of the island of Soledad ; the latter, in Behring's Straits. I had the good 
fortune, in 1839, to add a second and representative form, inhabiting the North Atlantic. 
Since then a third has occurred, and it is not improbable that before long many species may 
be discovered of this beautiful genus, seeing that its characters are peculiarly susceptible of 
specific modifications. Dr. Gould mentions one as occurring on the coast of the United 
States. 

The peculiar structure of the lips in the species of this and the following genus, appears 
to bear some relation to the grouping of the marginal tentacula in fascicles, and to constitute 
a character of sufficient value to cause in the end the establishment of a family distinct from 
Sarsiadce, for the reception of Bougainvillea, Lizzia, and a genus as yet undefined, of which 
the Medusa described by Rathke, under the name of Oceania Blumenbachii, is the type. 

* Lamennais, Esquisse d'uue Philosophic, B. v, ch. i. 
t Faerie Queen, canto v. 



62 BOUGAINVILLEA BRITANNICA. 



1, Bougainvillea Bntannica, Forbes (1841). 

Synonyms. Hippocrene Britannica. E. Forbes, in Annals of Nat. Hist., vol. vii, 

p. 84, pi. 1, fig. 2 (1841). 
Bougainvillea Bi'itannica. Lesson, Acalephes, p. 291 (1843). 
Medusa duoclecilia. Dalyell, Animals of Scotland, p. 70, pi. 11, 

figs. 11, 12(?) (1847). 

This beautiful little animated bubble is nearly globular, and usually not much larger than 
a marrowfat pea. Its umbrella is transparent, colourless, and quite smooth, therein differing 
essentially from the Hippocrene Bougainvillii of Brandt, which has pilose sides resembling, 
in this respect, Thaumantias pilosella. (See Brandt, in Petersburg Memoirs, Sixth Ser., Sc. 
Nat., vol. ii, pi. 20, figs. 2, 3, 4, and 6.) The opening of the umbrella is contracted and 
quadrangular. At each angle is an oblong group of tentacle-bulbs, closely packed together, 
six to eight in each group. Each bulb is particoloured, orange below, and white above, with 
a red eye-dot on the white portion. The bulbs seem all united into one mass or pad at their 
lower part, so that the tentacles are more close together at their origins than the ocelli. The 
tentacles are as many as the bulbs, not very long, yet slender, white towards their bases, 
orange towards their tips. The outer ones are usually borne curled upwards. The sub- 
umbrella is small as compared with the umbrella, less than half its size. It is divided into 
four sections by four simple gastric vessels, which join the marginal vessel opposite the groups 
of tentacular bulbs. From its centre hangs the massy peduncle, consisting in its upper part 
of four equal, compressed, quadrate lobes, of a bright orange colour, contracting below into a 
short, tubular, orange stomach. The latter terminates in a mouth surrounded by four very 
curious lips, for each is prolonged into a white filiform tentacle, which twice dichotomously 
divides ; each division terminates in a bulbiform extremity of an orange colour, with dark 
specks. The structure of these singular appendages to the mouth reminds us of the root- 
like cotyledonary tentacles of Cephea among the higher Medusce, and serves to bear out 
the view that those bodies are not substitutes for stomachs, absorbent roots, as it were, as 
formerly supposed, but only a modified form of fimbriated lips. The gland-like appearance 
of their extremities in Bougainvillea, seems to depend on terminal accumulations of fibrous and 
pigment- cells. 

The Bougainvillea Britannica is a very active little animal, and very tenacious of life. 
Its tentacula are continually in motion, and sometimes so contracted, that none appears to be 
present. It is abundant, but probably not gregarious, in various localities in the north. I 
have taken it in the Kyles of Bute, whence it was first described, at the entrance of the Frith 
of Forth, in Zetland, and in Ballycastle Bay, on the north coast of Ireland. It has also been 
taken on the east coast of Scotland by Mr. Henry Goodsir, and Mr. Patterson has.communicated 
a memorandum of a little Medusa, evidently this species, procured by hira at Portaferry, 
Strangford Loch, on the 7th of August, 1 838, so that he had met with and observed it before 
I had the same good fortune. 

Plate XII, fig. 1, a, represents this Bougainvillea of the natural size ; 1, b, magnified, 
and seen in profile; 1, c, as seen from above; \,d, the lobes of the peduncle. Between 



BOUGAINVILLEA NIGRITELLA, 63 

these lobes will probably hereafter be found young Bougainvillecs produced by gemmation, 
in the manner described as occurring in the following genus. The appearance of eio-ht 
peduncular lobes represented in Mertens's drawing of Hippocrene Bougainvillu, {Bougainvillia 
Macloviana, Lesson,) is probably due to this cause ; the four intermediate ones being gemmae, 
symmetrically developed, in this respect differing essentially from the unsymmetrical develop- 
ment so strangely exhibited by Lizzia. Those who have opportunities of hereafter examining 
our British Bougainville a, should let no specimen pass until the mode of reproduction be 
discovered. When found, I feel quite confident it will prove to be of the order now indicated. 
1, e, represents one of the ramified lips, with its gland-like extremities ; 1,/, is one of the 
four fasciculi of tentacles and tentacular bulbs. 

2. Bougainvillea nigritella, Forbes. 

A second British species of BougahiviUea — one, too, remarkably distinct from the first — 
was discovered by Mr. M'Andrew and myself in the Sound of Brassay, Zetland, durino- the 
autumn of 1845. 

It is very minute, not more than half the size of its congener. The umbrella is globose, 
smooth, transparent, and colourless. It is contracted at its opening, which is quadrano-ular, 
each angle bearing a compact, oblong, or almost kidney-shaped mass of tentacular bulbs, 
apparently four in number, closely united together, so that, but for indications of lobation at 
the lower part of the pad, the number of these bodies would be indeterminable. The upper 
half of the pad is yellow, the lower jet-black, the two colours separated in a very defined 
manner. On one side of each pad arises a very short, thick, yellow tentacle, and one only. 
The sub-umbrella, which occupies about two thirds of the body, is divided into four parts, the 
simple gastric vessels, each of which unites with the marginal vessels opposite the centre of 
one of the ocellated pads. The peduncle is short, and divided above into four rather thick 
oblong lobes, of a yellow colour ; below it is produced into a short, campanulate, yellow 
stomach, terminating in four tentacle-shaped, white lips. Each lip becomes suddenly filiform, 
proceeds for some distance simple, and then divides into two, again bifurcating before 
it terminates. The end of each division is a conical gland-like body, white speckled with 
black. 

Plate XII, fig. 2, a, represents Bougainvillea nigritella of the natural size ; 2, h, mag- 
nified in profile ; 2, c, seen from one side ; 2, cl, the peduncle and lips ; 2, e, one of the lips 
'with its sucker-like terminations ; 2, f, one of the masses of tentacle-glands, and the single 
tentacle. 



64 LIZZIA OCTOPUNCTATA. 



Genus XV. Lizzia, Forbes (1846). 

Umbrella spherical or campanulate ; ovaries in the form of four lobes, on the 
sides of the short peduncle; margin of the umbrella with eight unequal, compound, 
tentacular bulbs, all tentaculiferous, the four larger opposite the four radiating, 
simple, gastric vessels ; stomach shorter than the sub-umbrella ; mouth with four, 
simple, or ramifying tentaculated lips. 

I founded this genus for the reception of the remarkable Medusa described and figured 
by Sars under the name of Cytais {?) octoptmctata. I had previously referred it to Hippo- 
crene (i. e. Bougainvillea), and had been followed in such reference by Lesson, but the discovery 
of several true Bougahwillem with four fascicles of tentacles, and of more than one form with 
eight fascicles, indicated the propriety of separating the two tyjies, and of assigning each a 
generic value. Further observations have rendered it probable that the one genus produces 
its young by gemmation symmetrically, and the other unsyrametrically, which difiference, if 
constant, would of itself be sufficient to induce a generic separation of the two groups. 



1. Lizzia octopunctata (sp.), Sars (1835). 
Plate XII, Fig. 3. 

Synonyms. Cijt(sis{?) octojiunctata. Sars, Besk. og Jagt., p. 28, pi. 6, f. 14 

(1835), and Fauna Littoralis Norwegise, 
t. iv, figs. 7-13 (1846). 
Hippocrene octopunctata. Forbes, Annals of Nat. Hist., vol. vii, 

p. 84(1841). 
Bougainvillea octopunctata. Lesson, Acalephes, p. 292 (1843). 

Among the many important discoveries which have rewarded the patient observation of 
Sars, that of the power of Medusae to reproduce by gemmation is not the least significant. 
The animal now to be described was that in which the Norwegian naturalist met with the 
phenomenon. Hitherto it has been found only on the coasts of Scandinavia ; I have now 
the pleasure of making it known as a member of the British Fauna. To add a new form, 
even though of little interest, in a class with so few recorded native members as the Acalephae, 
is a pleasure ; much more so, to increase our lists with one of such curious physiological 
import as the Lizzia octopunctata. 

This little Medusa — it is scarcely a quarter of an inch in length — swarms in the bays of 
the eastern and western coasts of Zetland. I have not met with it elsewhere. Its umbrella is 
sub-globose or elongato-convex, smooth, transparent, and colourless. On the margin there 
are eight, jet-black, triangular ocelli, four of which are larger than the other four. All are 
compound, being composed of the united bulbs of several tentacula. Three of those organs 



LIZZIA OCTOPUNCTATA. 65 

spring from each of the larger bulbs, and either two or three from the smaller, the number 
varying in different specimens. These tentacles when contracted are rather short in pro- 
portion to the body, colourless, and not usually very extensile, though Sars has observed them 
extended to a great length. The animal, when swimming, often turns them up and curls them. 
The sub-umbrella occupies not quite two thirds of the body ; it is divided into four parts, by 
the four vessels running to join the marginal canal opposite the larger tentacular bulbs. Its 
upper part often appears as if truncated. The peduncle is short, thick, and four-lobed. It 
is marked with four patches of black pigment-cells. Between the lobes are seen, in the 
majority of specimens I have examined, four budding gemmules, one of which is invariably in 
a stage of advancement far beyond the others, and usually exhibits distinctly the black 
ocellated tentacular bulbs. The stomach occupies the lower part of the peduncle ; it is 
narrower than the upper, and more extensile. It is colourless, and terminates in four ten- 
taculiform lips, each one bifurcating. 

In St. Magnus Bay I took specimens similar in every respect to those just described, 
except in being a little larger, having slightly smaller ocelli, and no buds on the peduncle. 
These may possibly have been males. I have never seen the process of gemmation in the 
females advanced beyond the stage noticed above. Sars, however, traced all its stages, and 
as his account is of great interest, and contained in a work probably accessible to very few of 
my readers, I extract it entire : — 

" I considered the short cylindrical knots or appendages on the stomach (which hangs 
free in the cavity of the campanulate disk) of the Acalephse, described by me under the name 
of Cijtmis octopunctata, as very remarkable even at the moment of the discovery of the species. 
I could not at the time state their purport with certainty, but supposed that they had some 
connexion with the mode of procreation. 

" In the spring, 1836, I had an opportunity of observing a number of individuals of this 
species of Acalephse ; and I then discovered, to my astonishment, that the parts mentioned are 
nothing else than the young ones produced by gemmation, — a phenomenon hitherto unknown 
among the class of the Acalephse. I have briefly mentioned this interesting discovery in 
Wiegmann's Archives for 1837, Part V, p. 406. 

" I obsei-\red in some individuals, which I examined on the 5th of May, that these knots 
are all placed in a horizontal position (viewing the animal erect or with the mouth downwards), 
at the sides of the square-formed stomach. They are usually four in number, and are seated 
opposite one another. There are likewise frequently seen an additional two or four much 
smaller ones, placed beneath the former number. They are, moreover, usually of uneven 
dimensions, the two seated opposite one another being larger than the other two, and one of 
the larger pair is larger than the other. In one of these individuals a knot was developed 
into a perfect young animal, with a bell-shaped, colourless, transparent disk, in the cavity of 
which the oblong, pear-shaped, brownish-gray stomach was quite distinct. At the margin of 
the disk there were eight brownish-black, marginal granules, and the marginal fibres that 
spring forth from them, of which I counted sixteen, as long as the disk. The marginal fibres 
moved and bent slowly, and the entire disk was contracted occasionally. The young one was 
attached by means of a very short and rather thick peduncle (which issued forth from the 
back or from the convex surface of the disk) to the stomach of the mother, whilst it otherwise 
projected with its entire body independently. The young one seated opposite it had probably 

9 



66 LIZZIA OCTOPUNCTATA. 

already dropped off, for traces of the peduncle were perceived at the place where it had 
grown. Of the two other buds issuing forth fi'om this individual, the one was round at the 
independent extremity, and had only four brownish-black marginal granules, without traces 
of marginal fibres ; whilst the other exhibited very short, thick, prominent, marginal fibres, 
and traces of stomach internally. 

" In another individual the two opposite knots were small, rounded, and as transparent 
as water, without traces of stomach, marginal granules, or marginal fibres ; of the two 
other larger ones, the one was simple, without marginal fibres, but furnished with four 
marginal granules ; the other and largest of all also did not exhibit any marginal fibres, but 
had eight marginal granules, of which four were much larger than the rest, the latter being 
evidently those that had latest budded forth, owing to which circumstance they were seated 
alternately with and between the larger ones. 

" The number of these knots I found unequal in different individuals, depending upon 
the circumstance whether several or few young ones had already severed themselves, namely, 
from one to three, independent of the small knots seated farther below on the stomach, 
which become smaller the nearer they are to the mouth. The form of the latter, however, is 
quite the same as that of the others, excepting that they are as transparent as water, and 
without visible organs. 

" Among the larger young ones, which possess eight distinct, equally large, marginal 
granules, marginal fibres are likewise always found to grow forth, being equally long or even 
longer than the young one itself ; they are, however, generally found to be lying together in 
a bent position, and only become visible when they are unfolded by the aid of a needle, or 
when the young one severs itself forcibly from the body of the mother, in which latter case 
they begin to unfold and to move spontaneously. Their number is usually twelve, (viz. three 
grow from each of the marginal granules that first show themselves). Sixteen, however, are 
found with the largest young ones, or those that are about to sever themselves from the 
mother, (viz. one marginal fibre grows from each of the four subsequent marginal granules). 
In the last mentioned young ones, the stomach as well as the short mouth-tentacles are 
distinctly developed. 

" On the same day I remarked, in one of the largest of the individuals of this Acalephe, 
a young one, which was from five to six times smaller in diameter that the mother ; it appeared 
recently to have been severed, and still was slightly glued to the stomach of the mother, but 
separated from it immediately as soon as I touched it mth a needle, swam about in the water, 
and exhibited the same phenomena of life as its parent. It had eight marginal granules and 
sixteen marginal fibres. 

"On the 1 0th of May, I found in such a large individual a perfectly-developed young 
one, of the size of the one that I have just mentioned, and which was still glued to the body 
of the mother. I observed it with great attention, v^-ith a view, if possible, to see the process 
of separation. The disk, the stomach, the mouth-tentacles, marginal granules, and the four 
radiating canals, running from the stomach towards the margin of the disk, each of which corre- 
sponded exactly with the same parts in the mother, were evident. It was moreover as colourless 
as water, excepting the bro^iiish-gray stomach, and the brownish-black marginal granules. 
Occasionally it would violently contract, and again expand (just as the mother when swimming), 
a systole and diastole by which it strove to sever itself ; its contractions were quite independent 



LIZZIA BLONDINA. 67 

from those of the mother, and indeed evinced a distinct individual Hfe. The marginal fibres, 
the number of which were sixteen, viz. three and one alternately issuing from the marginal 
granules, were of the length of the disk or a little larger, and moved themselves, worm-like, 
in every direction. 

" I placed this individual by itself in a vessel filled with sea-water. On the very evening 
of the same day I found that the young one had severed itself from the mother, swimming 
about rapidly in the water. The bell-formed disk (from five to six times smaller in diameter 
than that of the parent) was more rounded at the upper part, and not so high as in the 
mother ; every trace of the locality of adfixture, which, as already mentioned, is at the back 
of the disk, had already disappeared. At its stomach I observed two small knots of an 
unequal size, being probably the first commencement of the issuing young ones of the second 
generation. In other young ones (that had not yet freed themselves from their progenitor), of 
about the same size as this, I have found four such unequally-sized little knots, — otherwise 
growing young ones — issuing forth from the stomach. 

" On the morning of the following day I had another rather smaller young one, which had 
been attached to the same mother, swimming about merrily with the above-mentioned one, 
the latter having grown to I — ^ of the size of the mother in diameter." 

The Li%zia octopunctata is a gregarious species. It is very conspicuous, owing to the 
jetty colour of its ocelli and ovaries. It moves much less gracefully than most of its allies, 
jerking itself through the water with sudden and ^^gorous leaps. It is lively, vigorous, and 
tenacious of life. 

Plate XII, fig. 3, a and b, represents it seen in profile, and from above, magnified ; 
3, c, is the peduncle with a budding young one far advanced, and the curious bifurcated lips 
of the stomach ; 3, d, is the same seen from above ; three of the buds in this instance are very 
rudimentary ; 3, e, represents two of the tentacular bulbs. The variety figured is that with 
only two tentacles springing from the intermediate bulbs ; that with three, as figured by Sars, 
is equally, if not more, abundant. I have no reason to suppose them to be specifically 
distinct. 

2. Lizzia blondina, Forbes. 
Plate XII, fig. 4. 

Rather smaller than the last species, not so common, and of solitary habit, is another 
Lhzia inhabiting the Zetland seas, and met with first in the Sound of Brassay, and afterwards 
off Fitful Head, during the autumn of 1845. 

The umbrella of the Lis,%ia blondina is sub-conical, smooth, inflated, and colourless. 
Its margin is ornamented with eight oblong, yellow, compound, tentacular bulbs, alternately 
small and large, each of the latter giving origin to three yellow tentacula, the former to only 
one, equally yellow, however, and not difl'erent from the other tentacles in dimensions. Their 
substance, when highly magnified, appears minutely and granularly ringed. The sub-umbrella 
is rather small in proportion to the umbrella, conic, and truncated above. Four simple mar- 
ginal vessels run down its sides, opposite the four larger fascicles of tentacula. A broad veil 
borders its opening. From the centre hangs a short, four-lobed, yellow peduncle, producing 
gemmules unsymmetrically, exactly as the last species does, in the intervals of the lobes. 



68 LIZZIA BLONDINA. 

Stomach very short ; mouth bordered by four tentaculiform lips, which do not bifurcate, but 
terminate with simple, gland-like, black-spotted, conical extremities. The motions of this 
very distinct species are jerking, and its habits, except in being solitary, like those of its 
congener. 

Plate XII, fig. 4, a and b, represents the Lhzia blonclbia much magnified, seen in 
profile and from above ; 4, c and d, are similar views of the peduncle, tentacula, and 
gemmules ; 4, e, is one of the larger fascicles of tentacula, with the compound bulb from 
which they spring ; 4, f, the appearance of a portion of one of the tentacles, greatly 
magnified. 

I have met, both on our southern and northern coasts, with several little Medusae in an 
immature state, presenting the essential characters of Lizzia, and evidently distinct from 
either of the preceding, but have abstained from describing or figuring them as species until 
they shall have been observed in an adult condition. I do this the more willingly, as it would 
seem that the intermediate stage of several hydroid zoophytes appears to be similar to, if 
not identical with, this genus. Thus the Medusa state of the Eudendrium ramostim, figured 
by Van Beneden in the 'Memoirs of the Brussels Academy' (vol. xvii, pi. 4, figs. 10-13), is 
closely related to both Bougainvillea and Lizzia, nor is the Medusa produced by Tuhidaria 
far removed. Sir John Dalyell has recognised this affinity of forms, in his account of the 
probable Medusa of Tithulnria, under the name of Medusa ocilia, and his comparison of it 
with a Bougainvillea (apparently B. Britannica), denominated by him Medusa duodecilia. 
His description is so interesting, and bears so importantly on the history of the Lizzia and its 
allies, that I venture to extract it from his most valuable and beautiful work ' On Remarkable 
Animals of Scotland,' in the hopes of attracting the attentive observer to a subject evidently 
prolific in future discoveries : — 

" Medusa ocilia. Minute pyriform bodies, as above specified, are dispersed on the stalk 
of different parts of the Tubularia at considerable intervals ; sometimes three are together, 
sometimes two are opposite to each other, or only one terminates a twig, where it might be 
readily supposed a regenerating hydra. Each pyrulum is affixed by its own distinct pedicle, 
at first of some length, but gradually shortening as the remainder becomes more globular, or 
flattens. In a few days, the whole may be compared to the opening bud of a white rose. 

" Now the dilatation and collapse of the subject commence ; convulsive struggles ensue ; 
four pair of long, rough, muricate organs resembling tentacula, or ciliary processes, are 
gradually unfolded, and after what seems repeated, severe, and protracted exertions, a perfect 
animal of great transparence is liberated as a Medusa, suspended amidst the waters. 

" This Medusa resembles a large transverse section of an ovoid, not half a line in diameter, 
the sides of excessive tenuity, the tentacula, or ciliary organs, four or five times as long as the 
diameter of the ovoid, and quite flexible. I conjectured there might be an orifice in the upper 
surface, and that some stump or particular organization by which it was penetrated, and 
remaining behind, the animal, amidst its struggles, was kept in its place. It is the upper 
surface which is that in application or adhesion, as may be seen of others ; the lower portion 
whence the cilia originate, is meantime free. Here, as we know, the mouth or proboscis of 
the Medusarian race is situated. In the under surface, four cruciform organs were apparent. 
But the difficulty of ascertaining relative position is never to be forgot ; and, in fact, organs 
apparently below, in such creatures may be in the middle, or actually above. 



"MEDUSA DUODECILIA." 69 

" As in some other Medusae, the ciliary or tentacular organs resemble knotted cords. 
Each pair originate from a prominent knob or button on the margin of the cavity in the 
under part ; and a black speck, like an eye, is conspicuous at the root of each tentaculum : 
there are, therefore, eight in all. The transparent convex surface of the animal is crossed by 
two darker lines at right angles, as if dividing it into quarters, and terminating in the four 
buttons or knobs on the margin. 

" The opacity of this creature, during its earlier stages, is not such as entirely to intercept 
the light, though its transparence refines in proportion to the evolution of its organization. 
Thus, the cross lines running from what was conjectured an orifice are discovered to be four 
vessels, wherein a fluid carries a number of black particles down to the marginal knobs below. 
It rather appears also, though I could not satisfy myself of the fact, that the current may pass 
in another course around the margin. 

" We should be much deluded, as well as our fellows, in believing that sufScient oppor- 
tunities are afforded for similar observations by the Medusa free of the zoophyte, and swimming 
at large. On the contrary, they can be effected only while immaturity yet restrains the 
animal to its pristine site ; and where, dui'ing progressive evolution, the microscopical focus 
can be accurately adjusted for distinct vision. The motion of so restless a being, when 
liberated, renders correct observation incompatible with that condition. 

" One of the pyrula seemed to be united to a hydra, just at the orifice of the twig from 
which it issued. Circulation manifestly advanced in both. Black particles were carried up as 
well as down the neck of the former, the pyi'ulum, and during their descent, a current was 
obviously conveying black particles up the body of the hydra ; something similar seemed to 
be going on in an isolated pyrulum, where there was no hydra. 

******* 

"The preceding animals originated and disappeared without leaving any traces behind. 
Indeed the smaller Medusae never leave anything that I have seen. 

" Another species {Medusa duodecilia) has sometimes appeared and decayed mysteriously, 
without affording obvious indications of its origin. This animal occurs in April ; the former is 
developed from June until August. It is an eighth of an inch in diameter, with si.xteen long, 
slender, tentacular, or ciliary organs, disposed in four bundles, which issue from four marginal 
prominences. At the base of each set is a black speck. The transparence of the Medusa is 
such, that while suspended in equilibrium in the water it is scarcely perceptible. 

" The evolution of the Medusa ocilia is progressive, accelerated perhaps by external 
temperature. It has occurred to me under no other conditions than as above specified. A 
number of whitish corpuscula seem to be generated rather suddenly among the hydrae, either 
singly, in pairs, or in clusters of three, four, or five, as already stated. Each is afiixed by an 
independent pedicle, and generally about the orifice of a twig of the zoophyte. One, wherein 
no subordinate organic parts were visible, on first inspection, was observed to unfold in three 
hours. 

" There is reason to believe that this Medusa is confined in a diaphanous vesicle or 
involucrum, remaining in its place after the animal has escaped. If actually so, which requires 
confirmation, we cannot but recognise strict analogy to the vesicles of the Sertularia, some 
being hardly perceptible after discharging their contents, from extreme transparency. 

" A colony, computed at 130 individuals, of the Medusa ocilia was produced in four or 



70 MODEERIA FORMOSA. 

five days ; and there are grounds for assuming that successive colonies come from the same 
specimens of the zoophyte. 

" I cannot presume to aflBrm that any connexion, immediate or remote, connects the 
Medusa oc'il'ia and the Tabular ia ramosa ; far less to conclude that the former shall be 
metamoqjhosed in progress of time to the latter, with its numerous and beautiful appurte- 
nances. I have not seen both the beginning and the end ; nor does the fugitive existence of 
so delicate a creature seem well adapted for permanent observation. (Dalyell's Rare and 
Remarkable Animals of Scotland, vol. i, p. 66, pi. XI.) 



Genus XVI. Modeeria, Forbes, (1846). 

Umbrella globose ; radiating vessels four, simple ; four marginal tentacles 
opposite the four simple vessels ; ocelli conspicuous ; peduncle inflated, balloon- 
shaped, contracted below, and terminating in four lanceolate lips. 

I have dedicated this very distinct genus to Adolph Modeer, a naturalist of no mean 
eminence, whose name has hitherto remained without such association in zoology. No one 
has a juster right to preside over a good genus of Medusa than Modeer, since to him we owe 
the first attempt of any value towards systematising the knowledge of the tribe acquired up to 
his time. This is contained in the twelfth volume of the ' New Transactions of the Swedish 
Royal Academy,' published in 1791, and is a very elaborate and masterly treatise for its day. 
The author was a Swede. He was a man of considerable and varied acquirements, and 
brought a mind exercised in comprehensive and practical thought, to bear beneficially upon 
natural history. As secretary to the Patriotic Society of Stockholm, he worked advan- 
tageously for the benefit of his native country, especially directing his attention towards the 
improvement of its agriculture and industrial arts. He wrote a ' History of the Commerce of 
Sweden,' and was the author of a ' Bibliotheca Helminthologia,' pubUshed at Erlangen in 
1776. He died at the age of 60, in 1799. 

Modeeria formosa^ Forbes. 
Plate Vn, Fig. 1. 

The largest objects are not always the most beautiful. Little diamonds may sparkle 
brighter than the monster gems of a regal crown. There is not a Medusa in all the ocean 
which can match for beauty vnth the minute creature now before us, though its smallness is 
such, that a split pea would overtop it. Yet small though it be, it has shape, colour, and 
substance so disposed, that as yet no explorer of the sea has met with another like it. It is 
gorgeous enough to be the diadem of the smallest of sea-fairies, and sufficiently graceful to be 
the nightcap of the tiniest and prettiest of mermaidens. 

The umbrella is globular, or slightly elliptical, smooth, and transparent. Its inner margin 
is bordered by a membranous veil. On the outer edge are four conspicuous tubercles, each 



EUPHYSA AURATA. 71 

with two crimson lateral pads, and an ocellus of the same colour between them. From each 
springs a tentacle, long, translucent, slender, curling, pale, or colourless. Centrally on the 
margin, between each pair of tentacles, is a smaller tubercle ocellated with crimson, but 
bearing no tentacle. On each side of this are three colourless tubercles, all very small, but 
the central one slightly larger than the others. The marginal vessel is rather large and 
conspicuous. The sub-umbrella occupies about two thirds of the umbrella, and is slightly 
conic in form. From its centre hangs an ample balloon-shaped peduncle, obsoletely four-lobed, 
the lobes of a brilliant crimson hue, the interspaces yellowish-white. I think it probable 
that the young are produced by gemmation from the crimson spaces. The peduncle 
terminates in a contracted neck, widening into a campanulate mouth, with four, rather short, 
lanceolate hps. This orifice is white, with four crimson lines. The peduncle sometimes 
contracts into a distinctly four-lobed form, and the lips are then more distinct than usual. 

The Modeeriaformosa was taken by Mr. M'Andrew and myself off Mull, in the Hebrides, 
during the autumn of 1845. 

Plate VII, fig. 1, a, represents it of the natural size ; \,b, magnified ; 1, <*, seen from 
above; 1, d, the stomach or peduncle contracted; a slight inequality of the ovarian lobes 
maybe observed; 1, e, a portion of the crimson bands of the peduncle much magnified, 
showing the structure of the supposed germinal membrane \,f, two of the tentacles, and the 
tubercles of the interspace (by mistake the small tubercles are represented as too few by four) ; 
1, g, structure of a tentacle and its bulb. 



Genus XVII. Euphysa, Forbes. 

Umbrella globose, inflated ; ovaries in the base of a flask-shaped peduncle, with 
a simple orifice at the end of a proboscidiform stomach ; vessels four, simple, joining 
the gastric vessel opposite four conspicuous ocellated tubercles, from each of which 
arises a short, slender, recurved cirrus, and from one a supplementary large tentacle. 

Euphysa aurata, Forbes. 
Plate XIII, Fig. 3. 

I have constituted this genus for the reception of a very beautiful little Medusa taken 
in 1835, in Brassay Sound, Zetland. At first I included it in my genus Steenstrupia, and it 
is one of the three species of that group announced at the meeting of the British Association 
at Southampton. But although bearing a close aifinity with Steen.itrupia through the 
arrangement of its tentacles, and the form of the stomach, it presents characters, such as the 
construction of the umbrella, the presence of conspicuous ovaries, the absence of an apical 
appendage, and the freedom of its lesser tentacles, which must be regarded as generic. It is 
indeed evidently of higher rank in the scries than Steenstrupia, which, we shall presently see, 
has many of the features of a rudimentary animal, a stage in the metamorphoses of some 
creature in a different order. 



72 STEENSTRUPIA. 

The EnpJiysa aurata has a smooth, inflated, globose, transparent umbrella. The orifice 
is rather contracted, and square. At each of the four angles is a large diamond-shaped 
ocellus, the upper half of which is bright golden yellow, and the lower vivid scarlet or 
crimson. From each of the ocelli springs a short, reflexed, cylindrical, yellow tentacle, which I 
have never seen to extend itself. From one of the ocelli, below the short tentacle, arises a long 
and thick one, highly extensile, and of a golden colour, usually presenting a club-like shape. The 
smaller and larger tentacles are identical in structure. A marginal vessel and veil bounds the 
opening of the umbrella. The peduncle is flask-shaped, the neck of the flask being the 
stomach, and terminating in a contracted, simple, round opening, which I never saw protruded 
beyond the disk, though often moved about in various directions ; the walls of the peduncle 
are apparently banded with motor tissue. The orifice of the stomach is tinged with 
red. The remainder is yellow, with four slightly-marked taT\Tiy bands on its lower part. 
Within the centre, and at the base of the peduncle, is a p3Tamidal cluster of reproductive 
cells, constituting the ovary. The size of the body scarcely exceeds one sixth of an inch. It 
is an active and lively little animal. 

Plate XIII, fig. 3, a, represents it of the natural size ; 3, h, magnified ; 3, c, is one of 
the smaller tentacles and ocellus ; 3, d, the same seen in front ; 3, f, the extremity of the 
larger tentacle ; 3, e, the peduncle and ovaries as seen under compression. 



Genus XVIII. Steenstrupia, Forbes (1846). 

Umbrella conical, apiculate ; apex connected by a cord with the sub-umbrella ; 
four marginal elongated glands opposite the four simple radiating vessels ; a single 
tentacle developed from one of the glands only ; peduncle proboscidiform, with a 
simple round orifice. 

Whatever may be thought of the other genera of Sarsiacleee, there can be no question 
that this is intimately related to zoophytes of the hydroid order, and is, in all probability, an 
intermediate stage. Yet animals, so singular and significant as those which I have brought 
under Steenstrupia, cannot be left unnamed until their larva or final conditions, as the case 
may prove, be determined. Nor, I trust, will the illustrious naturalist of Denmark, in lionour 
of whose genius I have ventui'ed to designate the group, disdain it, even though it prove to 
be only provisional, since whatever may become its eventual rank, in all probability the 
creatures it includes will yield valuable illustrations of the ingenious and important theory 
which he first presented in definite shape. 

Indeed several of the figures which Steenstrup has given in the first plate of his ' Essay 
on the Alternation of Generations,' representing the hydroid polype Corijne fritillaria, and 
those in the first plate of Sars's ' Beskrivelser, &c.' delineating the gemmules of that 
exquisitely beautiful zoophyte, the Corymorpha nutans, bear so close a resemblance to the 
two forms of Medusa which I am about to describe, that I can scarcely doubt their close 
affinity. It is not impossible that my Steenstrupia rubra may eventually prove to be the full- 
grown mcdiisoid of the Corymorjiha, — a supposition rendered the more probable by the 



STEENSTRUPIA RUBRA. 73 

circumstances that both animals as yet have been found only in the Zetland and Orkney seas, 
■where that giant zoophyte was first enrolled among British species by Professor Goodsir and 
myself, in 1839. The specimens taken by us had, however, their buds too immature to 
permit of our perceiving the close resemblance and affinity of those bodies to true Medusae. 
The apical process, which is so striking a feature of the following animals, is almost certainly 
the remains of a funiculus by which the young animals were attached to its parent — but 
whether that parent was a " nurse," guised as a hydriform Coryne, or a Medusa, like the 
offspring, as we have seen to be the case in the gemmiparous Sarsia and Zizxice, is a point 
which fortunate future observations only can determine. 

1. Sleenstrupia rubra, Forbes. 
Plate XIII, Fig. 1. 

The umbrella of this strange little Meduste is conical, rather elongated, transparent, 
smooth, and colourless. Its summit bears a little tentacle-like, fleshy, red appendao-e. Its 
orifice is rather contracted and quadrangular. At each angle there is an elongated, slanting, 
tentacle-like ocellus, fixed throughout, and terminating above in a bulb. This body is 
entirely of a vermilion red. From the side and lowest part of one of them springs a very lono-, 
thick, fleshy, bright-red tentacle, which twists and coils like a worm, and under the microscope, 
presents a ringed and granulated structure. The sub-umbrella is oblong. It is comparted 
by four simple gastric vessels, running to the tentacular bulbs. Its orifice is surrounded by a 
veil. From its centre depends a thick, tubular, very contractile, fleshy, red peduncle, 
terminating in a round or imperfectly quadrate mouth. This peduncle is capable of great 
changes of form, and sometimes presents an appearance as if it had a nucleus denser than 
the substance composing its walls. Though highly extensile, it does not appear even to be 
voluntarily produced beyond the opening of the disk. Its base is connected with the httle 
finger-like process on the apex of the umbrella, by a rather tortuous colourless cord, presenting 
a tubular appearance. 

The length of the body is about a line and a half. Small as this Medusa is, it is very 
conspicuous in the water, owing to the brilliant colouring and fleshy substance of its tentacles 
and stomach. It is very active and tenacious of life ; before dying, assuming all manner of 
strange shapes, doubling itself up, and turning its organs inside out in a terrific manner, givino- 
up the ghost with convulsions as fearful as those of a popular actor in the death-scene of a 
tragedy. One of the least strange of these moribund attitudes is represented in Plate XIII, 
fig. 1, d, where the creature has constricted its body so as to assume the aspect of some twin 
Acaleph, such as Diplnjes. At such times, if we had not seen the animal previously in a 
healthy state, it was very diiEcult to perceive any resemblance between it and the other 
genera of its family. But when well and uninjured, it is an extremely active and regularly 
formed creature, though, owing to the weighty and unbalanced tail which it is doomed per- 
petually to drag as its train, it cannot advance through the water with the easy grace and 
rapidity for which its allies are remarkable, but struggles forward with frantic energy, con- 
tracting and expanding rapidly, and without ceasing, reminding us of an escaped felon impeded 
in his course by the dragging of his heavy fetters. When I first saw how the weight and 

10 



74 STEENSTRUPIA FLAVEOLA. 

one-sidedness of the simple tentacle arrested the motions of this Steenstrupia, I fancied that 
the animals before me had by accident been deprived of their corresponding marginal 
appendages. But among hundreds of specimens secured in the bays of both sides of Zetland, 
there was never the slightest sign of a symmetrical arrangement by the development of more 
tentacles than one. 

Plate XIII, fig. 1, a, represents Steenstrupia ruhra of the natural size ; \,h, magnified, 
as seen in profile ; 1, c, the arrangement of tentacular bulbs and tentacle, as seen from below ; 

1, d, the distorted specimen noticed above; 1, e, structure of the tentacle when very much 
elongated ; and \,f, its usual appearance when contracted. 

2. Steenstrupia flaveola^ Forbes. 
Plate XIII, Fig. 2. 

Umbrella conical, mitrate, transparent, colourless, not exceeding a line in length. Its 
summit is produced into a mucronatcd termination, which, though transparent and colourless, 
appears to be of a different tissue from the rest of the disk, which, besides, is quite smooth, 
whilst the apex is as if pilose. The margin is contracted as in the last species, and 
quadrangular, each angle bearing a similar elongated ocellus or tentacle-bulb, in this instance 
of a fawn yellow colour. One only of these bulbs gives origin to a tentacle, very slender, long, 
and moniliformly granulated. From the summit of the sub-umbrella, a cord or tube runs to 
the apex. Its sides are marked by four simple, radiating vessels. The peduncle is very 
changeable in form : sometimes contracting into a very short, thick, quadrate mass ; at others, 
assuming the shape of that of Sarsia, but not protruding beyond the disk. It is also of a 
pale yellowish, fawn colour. The mouth has no produced lips. 

This singular little animal was taken in Penzance Bay towards the close of August, 1846. 
Plate XIII, fig. 2, a, represents it of the natural size ; 2, b, magnified, its peduncle contracted; 

2, c, with the peduncle extended ; 2, d, the apex of the umbrella ; 2, e, one of the tentacle 
bulbs. 



75 



Having now enumerated and described those naked-eyed Medusae, which have come 
under my notice* in the British seas, I shall proceed to offer a few remarks on their systematic 
relations with other animals of their class, and on the affinities of the Pulmograde Acalcphae, 
with the members of other sections of Radiata. But before I do so, I think it best for the 
convenience of my readers to enumerate very briefly the higher or Steganopthalmatous 
Pulmograda, known to me as inhabiting our coasts, in the hope of directing attention to the 
study of the larger species, which afford fine materials for original researcli. I look forward 
at some future time to describe and figure them in a companion Monograph to this, but require 
many more observations and drawings before that can be done in a satisfactory manner. 

Being so much larger than the subjects of this volume, they are much more familiar to 
frequenters of the sea-side ; and as several of the species are gregarious and very generally 
distributed, they present good opportunities for the acquirement of a knowledge of the 
structure of Meduste in general. Every person who has been in a boat on a calm day, or 
looked over the side of one of our harbours when the tide was flowing in summer, must 
have seen large transparent gelatinous disks, with fringed margins, contracting and expand- 
ing, making their way beneath the surface of the water. All who have walked much 
along the wet sands when the tide is out, must have met with great pads of transparent jelly, 
marked in the centre with purple circles, or edged and rayed with bro\vn. The latter, when 
handled, sting severely ; the former are harmless. They are the two most common kinds of 
covered-eyed Medusae, members of the genera Attrelia and Cyanaa. The following brief 
notices will enable the reader to distinguish between our native species of Pulmograda 
Gymnoptlialmata . 



Genus Aurelia, Peron. Medusa, Eschscholtz. 

1. To this genus belongs the commonest of our native species, the Aurelia aurita, a 
hemispherical, translucent, bluish, gelatinous disk, margined with a close fringe of fine 
filiferm tentacula, interspersed at eight points by as many ocelli, each composed of an egg- 
shaped, pedunculated, black body, with a red speck above it. The sub-umbrella is marked 
by numerous radiating vessels, dichotomously dividing and most of them anastomosing in their 
course towards the margin. Sometimes these vessels present a deep purple hue, and then 
we have the spurious species Aurelia lineolata of Peron, A. radiolata of Lamarck, and 
Medusa jnirpurata of Pennant. Borlase first noticed this variety, and correctly considered it 
such, and not distinct from the ordinary form. Four long arms, with membranous and 
fringed edges, spring from the centre of the sub-umbrella. In the middle of them is the 
mouth. In particular states of the animal, the fringes and margins of the arms serve as 
marsupia for the eggs. Between each pair of arms is a raised cartilaginous tubercle, with an 

* Notices of doubtful or imperfectly described forms, will be found in the Bibliographical 
Appendi.x. 



76 PELAGIA. 

opening at its inner side entering the cavities of the body in which we find the ovaries. These 
are four in number, shaped hke horse-shoes or half-moons, of a bright purple colour. They 
are the four purple crescent-like marks which shine through the disk of the jelly fish as we 
see them swimming in the water. In some monstrous varieties they become united, and form 
a circle round the disk, or are multiplied, or, more rarely, aborted. The disk often measures 
nearly a foot across. It is very minutely granulated ; when more coarsely so than usual, we 
have the variety which has been called A. granulata. The specific names rosea, Surirea, 
Uneolata, radiolata, jmrjmrata, are all so many s)monyms of the Aurelia aurita ; and, 
judo-ing from the description, the Biblis Aquitania of Lesson was, in all probability, nothing 
more than this common Medusa cast high and dry on the sands ! It is everywhere abundant 
around our coasts, and sometimes occurs in vast numbers, impeding the course of boats 
through the water. Figures of it may be found in the Zoologia Danica, in the Berlin Trans- 
actions for 1837, and in the commemorative edition of the Regne Animal. 

2. A second species of Aurelia, for which I propose to retain the name Campanula, 
occurs abundantly in Southampton water, and some other confined localities. It appears 
to be the Medusa campanula of Otho Fabricius, and is a much more delicate animal than the 
Aurelia aurita, differing from it also in size, proportions, and ocelli. The umbrella is much 
more shallow, and never attains one fourth of the dimensions. The margin and arms are 
fringed with white tentacles, so that when the animal is seen in the water it appears as if 
conspicuously marked by a white cross. The ovaries are of a pale or tawny purple. The 
egg-shaped bodies of the ocelli are white, with a red spot above. This is probably the Medusa 
cruciata of some authors. There is no figure of it published. 



Genus Pelagia, Peron and Lesueur. 

3. Until the autumn of 1826, no example of Pelagia had occurred in the British 
seas ; in August of that year, several specimens of the Pelagia cyanella, one of the most 
beautiful and phosphorescent Medusse of the Atlantic, were taken by Mr. M'Andrew and 
myself, off the coast of Cornwall. The disk is sub-globose, and measures nearly three inches 
across. It is tinged with a rich rose colour, and is speckled over, especially at the sides, by 
small orange warts. Its margin is scalloped into sixteen lobes, from beneath eight of which 
spring as many highly contractile, purple, tubular tentacula, and in the notches of the other 
six, are eight red, protected, pedunculated ocelli. From the centre of the sub-umbrella hangs 
a thick peduncle, which soon divides into four lanceolate, winged and furbelowed, rose- 
coloured, orange-spotted arms, nearly four inches in length. Around the bases of the arms 
are, the openings above the four purple ovaries. A full description and figure of this beautiful 
species, will be found in the Annals of Natural History, vol. xix, p. 390, pi. 9; fig. 5. 

When the Pelagia phosphoresces, it seems like a great globe of fire in the water, an 
appearance familiar to those who have sailed on the coast of Italy, where animals of this 



CYAN.EA. 77 



Genus Chrysaora, Peron. 

4. The Mediisafusca and M. tuherculata of Pennant, described by him after Borlase, 
are varieties of the animal called Medusa hysoscella by Linnjeus, and belong to the genus 
Chrysaora of Peron. The umbrella is hemispherical and expanded ; its margin is festooned, 
and furnished with more than eight tentacula. The arms beneath are long, simple, and all 
separate ; the mouth is centrally in the midst of their bases. Four openings conduct to the ovaries. 
Our British species has a yellowish or reddish disk, marked with more or less distinct pale 
rays, and often spotted with brown at the margin. It attains considerable dimensions. It 
varies much ; Peron constituted seven spurious species out of its varieties. A good figure 
after an original drawing by Milne Edwards, may be found in the commemorative edition of 
Cuvier's Regne Animal. 



Genus Rhizostoma, Cuvier. 

5. The Rhizostoma pidmo {R. Aldrovandi and Cuvieri of Peron) is one of the largest 
of European Medusse, its disk growing to two feet, or even more, across. The umbrella is 
hemispherical, thick, and of a bluish hue, scalloped at the margin. Beneath is a thick peduncle, 
di\-iding into eight, long, tapering arms. At their bases are great, fringed, ovarian lobes, 
tinted with yellow and puq^le. It is usually stated to be plentiful on the English shores, but 
I have seldom met with it. 



Genus Cassiopea, Peron. 

6. The Medusa lunulata of Pennant, Urtica marina octopedalis of Borlase, is a species 
of Cassiopea. It is the Cassiopea Borlasea of Peron, and C. rhixosiomoidea of Tilesius. 
The disk is large, wide, depressed, and campanulate ; the margin scalloped, but not ten- 
taculated. There are eight ovaries below, opening by as many orifices, and eight pinnated 
arms with furbelowed appendages at their bases. A good figure of this species is given by 
Tilesius, in the Act. Acad. Nat. Cur. vol. xv, pi. 71. It is named on the plate C. aiiglica. 



Genus CvANiEA, Peron. 

7. Next to the Aurelia aurita, the commonest Medusa of our seas, is the large Cyanaa 
capillata, formidable on account of its stinging power. The disk is wide, and nearly flat, of 
a pale yellow or tawny colour, deeply scalloped at the margin into sixteen quadrate lobes, 
between each pair of which, in a deep notch, is a conspicuous pedunculated ocellus. Eight 
brownish rays proceed towards these ocelli, from a circle of reticulated, quadrate, brown mark- 
ings, giving the whole disk a beautifully stellate appearance, depending on the arrangement 



78 ' CYANEA. 

of the origans on the sub-umbrella, which is furnished with long, plicated, and furbelowed 
membranous arms, and fasciculi of extremely extensile, stinging, filamentary tentacles. The 
varieties of this Cyanea have been made into numerous spurious species by Peron. 

8. CyancBU LamarcMi, Peron. Not quite so common as the last, though very frequent 
in the Irish sea. It is easily distinguished by its more convex disk, of a deep ferruginous hue 
in the centre, and divided at the margin into eight four-lobed triangular lobes, the eye notches 
at their apices. The ovaries are of a rose colour. It stings equally severely with C. capillata. 
Better figures of both these Medusa are wanted. For one of the former, the memoir of 
Gaede, in the ' Bonn Transactions,' may be consulted ; the latter is represented in the 
92d plate of the ' Encyclopedic Mcthodique, Vers' (copied from Dicquemare). 



All the genera and species of higher Pulmograda are distinguished from the lower forms, 
to the description of which this treatise is devoted, by a much greater complexity of structure, 
especially in the vascular system and organs of sense, and also in the arrangements of the 
reproductive system. The vessels branch and anastomose ; the ocelli are protected by 
complicated coverings, and are themselves of more perfect organization ; the generative glands 
are more highly developed in the Sleganoiitlialmata than in the Gymnopthalmata. We have 
no instance of the Medusa or perfect state of the former propagating itself by gemmation, 
whilst such a mode of procreation occurs in several cases, as we have seen, among the 
latter. Taking one character with another, then, we cannot doubt that the Pulmograda 
Steganoiithalmata are higher in the series than the Pulmograda Gymnopthalmata. This is 
borne out by an examination of the phases of development and metamorphosis of the larva in 
the latter. The observations of Sars, Dalyell, Reid, and Steenstrup indicate that the early 
stages of the Aurelim and CyanecB, correspond closely structurally with the perfect condition 
of the naked-eyed Pulmogi-ades. 

Hitherto the genera of these two very distinct, yet proximate, tribes have been grouped 
together, without much respect to their natural affinities or serial order. Among the more 
important attempts at arrangement of the Acalephte, are the systems of Peron and Lesueur, 
of Lamarck, Eschscholtz, Cuvier, Blainville, Brandt, and Lesson. In the first of these, all 
the Pulmograda are grouped under two tribes, termed agastric and gastric, the latter being 
subdivided into monostomous and polystomous. Such a division is founded on a complete 
misapprehension — one which prevailed generally among zoologists, until Milne Edwards 
carefully examined the anatomy of Caryhdea marsupialis, which, however, in the arrange- 
ment under consideration, was placed in the gastric division, owing to the- naturalists who 
proposed it having mistaken the whole concavity of the sub-umbrella for a stomach ! Geryonia 
was, on the contrary, enumerated among agastric genera, the nature of tlie true stomach, 
situated at the extremity of the peduncle in that group not having been recognised. We 
find Oceania, which included a heterogeneous assemblage of naked-eyed species, placed side 
by side with Pelagia, in a division of the Monostomous Gastric Medusee. The greater number 



CLASSIFICATION OF MEDUSiE. 79 

of the Steganopthalmata are included in the Polystomous section, and the association as a 
natural assemblage indistinctly recognised. 

The arrangement of Lamarck was no better than that of Peron and Lesueur, and, in some 
of the details, inferior, as for instance, in the comprising of the species of Pelagia in his genus 
B'mnaa, associated with numerous forms of Geryonidce. All the Pulmogruda were grouped 
under two great sections, the one characterised by a single mouth, the other by the presence 
of several mouths; Lamarck, like Peron, having mistaken the ovarian orifices in certain 
Steganopthalmatous species for so many digestive openings. 

Eschscholtz, whilst he greatly improved, through his pei'sonal experience of the Medusae, 
the generic and specific arrangements, went astray as widely as his predecessors when he 
attempted their classification. For, as we have already seen, he mistook the ovaries in many 
genera for appendages of the digestive system, and regarded such forms as constituting a 
great cryptogamic section. Hence he divides all his Biscophoroi — a happily chosen term by 
which he designates the Pulmograda — under Discophone plianerocarpa and Discophora 
cryptocarpa>. But though this was a classification based on false notions of structure, so 
true was Eschscholtz's perception of the natural affinities of the genera, that the covered-eyed 
forms are all assembled under his first division, and the naked-eyed under the second. His 
minor groups are generally very excellent, though throughout all their characters the great 
mistake just mentioned prevails, and consequently nullifies them. 

Cuvier assembled all the Pulmograda in the section of Acalephm, which he termed 
" Meduses Propres," dividing it into five groups, of which the first, " les Equorces," is 
characterised by the presence of a simple short mouth, without tcntacula ; the second, " les 
Pelagies," by the mouth being prolonged into a peduncle, which becomes divided into arms ; 
the third, " les Cyanees," in which the mouth is central, and there are four lateral ovaries ; 
the fourth, "les Rhizostomes," in which there is no conspicuous mouth, nourishment being 
derived through the ramifications of the peduncle, the ovaries four or more ; and the fifth, 
" les Astomes," without central mouth, or ramified peduncle, or distinct cavities for the 
ovaries. His first and last tribes included the naked-eyed species. The whole arrangement 
is a mistake, founded on misapprehension of the value of characters in the order. The groups 
are neither natural nor of equal systematic value. The classifications of Peron and of 
Eschscholtz, though founded on mistakes as great, if not greater, are much superior to those 
of Cuvier and Lamarck, doubtless owing to the superior practical acquaintance of the former 
naturalists with the objects under arrangement. Cuvier's personal knowledge of the 
Discophorae seems to have been Umited to two or three of the higher species ; Lamarck had 
probably no experience in this tribe. Peron, Lesueur, and Eschscholtz had observed and 
studied numerous forms in the living state, and consequently, though of inferior order of mind 
to the former great naturalists, came nearer the truth in their systems, because their know- 
ledge was sound and practical, and not gained at second-hand. 

The arrangement of the Pulmograda proposed by De Blainville, is likewise the result of 
book-study, and not of sea-research, and is consequently objectionable. He divides them 



80 CLASSIFICATION OF MEDUSA. 

into simple, tentaculate, sub-proboscidean, proboscidean, and branched. In the last section 
are assembled the covered-eyed species. The juxtaposition of Mquorea and Ohelia in the 
second section, of Thaumantias and Conis in the third, and of Hippocrene and B'lanaa 
in the fourth, are instances of the unnatural way in which the genera are distributed in this 
otherwise ingenious system. 

Brandt, who appears to have founded his studies among Medusae chiefly upon the 
drawings and notes of Mertens, divided the DiscophorcB into Monostomous and Polystomous, 
subdividing the former into Oceanidm, in which we find Circe and Conis placed together, 
Mqtiorklce and Mediiskla:, the latter including the covered-eyed species very naturally 
assembled, except the Tilu%ostomkl(B, which, along with the Geryonidm (in which tribe he 
includes Hippocrene), constitute the polystomous section ; so that it is evident this eminent 
author did not clearly perceive the affinities of the several groups. 

Lesson, who, besides an extensive acquaintance with living forms, had the advantage of 
being last in the field, arranged the Biscophoree under four groups: \iX," Les Mediises 
non-proboscidees," in which we find several families, including genera, juxtaposed, having no 
immediate affinity; 2d, '^Les Oceanides ou Meduses vrais," including Mquorea and its allies, 
with a heterogeneous assemblage of forms in the genus Oceania ; 3d, " Medtises agaricines 
OH prohoscidees" — here are Sarsia, Biancea, Geryonia, Tima, Thaumantias, &c., the group 
being in the main equal to my Geryonida, though some genera, as Saphenia, are brought 
into it far away from their fellows ; and 4th, '■'^Meduses a pedoncule central ou Rhixostomees" 
consisting entirely of the covered-eyed species, much more naturally assembled together than 
in any of the preceding classifications,.except those of Peron and Eschscholtz, a personal 
familiarity with the objects he describes, having led Lesson, as it did them, to similar arrange- 
ments of the more conspicuous tribes. 

All the authors I have just cited, regard the Discophora, or Pidmograda, as a separate 
division of the Acalephw, or Arachnoder7nata, the whole class being distinct from the 
Zoophyta. Recent discoveries, however, would go far to show that such a separation is 
unnatural, and that the hydroid Zoophytes, at least, are very closely allied, if not belonging, 
to the same natural order with the Pulmograde Medusse. 

On the side of the Zoophyta, the facts bearing on this question have been chiefly derived 
from the families Corynidm, Tubidariada, and the genus Campanularia. More than a 
century ago, when the nature of Zoophytes, whether animal or vegetable, was under dis- 
cussion, Bernard de Jussieu, who pronounced rightly for their animal origin, described 
certain little round, red, pedunculated bodies, encircling the head of the Tuhularia. Nearly 
half a century after. Otto Frederic Muller observed similar bodies around the head of the 
Cory7te, and maintained that they were eggs, and for an equal period this view of their nature 
was generally received. In the year 1833, Professor Rudolph Wagner gave an account* of 
the production of medusiform bodies in a Zoophyte of the Adriatic, the Coryne aculeata, 
which bodies he regarded as the young of the animal, although they themselves contained 

* In Oken's Isis, for 1833. 



CLASSIFICATION OF MEDUSA. 81 

eggs. In 1835, Sars represented medusiform bodies similarly produced, from the bases 
of the tentacula of the Corymorpha fiufans ; in this case the Mcdusoids closely resembled 
Steenstrupia* In the same year. Dr. Loven observed the formation of Medusoids on the 
Syncoryne, the animals produced being very similar to Medusse of the genus Sarsia. About 
the same time, Sir John Graham Dalycll described the formation of Medusa-like buds on 
Tuhular'm. In 1 840, Professor Steenstrup, when in Iceland, found a polype, which he named 
Coryne J'riiillaria, from whose head bell-shaped bodies, closely resembling our Steenstrupia 
in form, hang, and were regarded by him as individual animals.f In 1843, M. F. Dujardin 
communicated to the ' Annales des Sciences Naturelles' a short but interesting paper, " On a 
New Genus of Medusaires, proceeding from the Metamorphosis of Syncoryne." He calls 
the parent animal Stauridia, and the medusoid, which he saw detach itself and swim away, 
Cladofiema. He remarks, that it is closely allied to the genera Oceania, Thaumantias, 
and Cytais. In 1844, Professor van Beneden, of Louvain, published his 'Researches on the 
Embryogeny of the Tubularise ;' and in his memoir described and figured medusiform-bodies 
produced from species of Tubiilaria, Endendrium, and Syncoryna. Those of the last-named 
genus resembled Sarsia ; those of the two former had close affinities with Lixzia, especially 
the medusoid oi Eudendrium. In 1846, Sars, in his ' Fauna Littoriilis Norwegiae,' figured the 
medusoids oi Syncoryna Sarsii, Podocoryna carnea, and Ferigonytniis irinscoides, all closely 
resembling Sarsics. That of the Podocoryna comes very near the Medusa papillata figured 
by Abildgaard in the ' Zoologia Danica.' 

Similar observations have been made, from time to time, on the CampamdaricB. Ellis 
appears to have been the first to notice the productions of Medusoids in that tribe, though 
he evidently did not understand what he saw. In 1834, Mr. Lister communicated his 
valuable microscopic observations on Zoophytes to the Royal Society, and in his paper 
describes and figures Medusa-like animals in course of production fi-om CampamdaricB. In 
1836, Sir John Graham Dalyell distinctly proved and made known the production from 
Campamdarim of free animals like true Medusae. He named the creature " Animalculum 
tintinnabulum.":j: In 1839, Nordman announced his observation of the free Medusa condition 
of young Campunularice .^ In 1843, Van Beneden, in his ' Memoirs on the Campanularise 
of the Coast of Ostend,' entered into full details on the subject. The figure which he 
gives of the medusoids in this tribe reminds us strikingly of Tima and Geryonia. He remarks 
that the Medusa marina of Slabber, the type of the genus Ohelia of Peron and Lesueur, 
is a young Campanuluria. During the same year, similar phenomena were observed by 
Kolliker. 

Various interpretations were offered of these phenomena. Many zoologists followed 
MuUer, and regarded them as eggs. Others held them to be gemmules. Ehrenberg put 
forth the strange theory that they were female polypes, a view supported by Loven and 
Krohn. Van Beneden considered them young budding polypes, a notion held previously by 

* Besk. og Jagt., pi. 1, fig. 3, g, 3,f. 

t Steenstrup, Alternation of Generations, Mr. Busk's Translation, p. 27, pi. 1, fig. 41-3. (Raj' 
Society.) 

X Ed. New. Phil. Journ., vol. xsi. 
§ Comptes Rendus, 1839. 

11 






82 CLASSIFICATION OF MEDUSA. 

Dalyell, and supported by Kolliker. Dr. Johnston maintained the same view of their nature. 
Steenstrup struck out a most original and distinct speculation, holding them to be alternate 
generations, produced by gemmation from a dissimilar parent, and producing eggs from which 
should spring dissimilar children. 

In all instances where a Medusa has been observed originating from a hydroid polype, 
the new animal bears the closest resemblance to a naked-eyed Medusa, — a resemblance not 
merely of external form, but also of internal structure. Indeed, in many cases it would be 
impossible to draw a line between the two. 

Is it desirable to draw such a line ? Are not the so-called Zoophytes and Medusae animals 
of the same section ? Discoveries exactly comparable, and still more wonderful, have shown 
that the higher Medusae themselves afford instances of parallel plicnomena. We now know for 
certain — all the stages of the history having been demonstrated — through the researches of 
Sars, Dalyell, Siebold, Steenstrup, Price, and Reid, that the ova of the covered-eyed Medusae, 
belonging to the genera Aurelia, Chrysaora, and Cyanaa, give rise to polypoid animals, which 
in their turn originate individual Medusae by fission — or, more properly, as Dr. Carpenter 
has well suggested, by a peculiar process of gemmation. The higher genera of Biscophora, 
therefore, are closely linked, anatomically and physiologically, with the Anthozoa hydroida 
among the polypes. Assuredly any separation of such nearly allied animals, especially the 
placing of them in different classes, is exceedingly unnatural. 

In what light are we to regard the relationship between the Medusa and the Polype ? The 
one is not the larva of the other, as often improperly said, because there is no metamorphosis 
of the one into the other. The first is the parent of the last, and the last of the first, but neither 
is a stage of an individual's existence destined to begin life as a Medusa and end it a Polype, 
and vice versa. The notion that the Medusoid of the Campanularia, or Coryne, or Tuhidaria, 
fixes itself, and changes into the typical forms of those zoophytic groups, is as inadmissible as 
the supposition that the hydroid product of the Aurelia is metamorphosed into a Medusa of 
that genus. Facts show that such is not the case. These facts may be summed, in the 
abstract, in the following formulae : — 

1st. The case of Tubidaria and Campanularia. 

a. The medusoid produces eggs. 

b. The eggs produce infusoria. 

c. The infusoria fix and become polypidoms. 

d. The polypes of these polypidoms produce medusoids. 

2d. The case of Aurelia, S)'c. 

a. The medusa produces eggs. 

b. The eggs produce infusoria. 

c. The infusoria fix and become hydroid polypes. 

d. The hydroid polypes produce medusae by gemmation. 



CLASSIFICATION OF MEDUSA. 83 

3cl. The case of Coryne, ^c. 

a. The zoophyte produces medusae by gemmation. 
h. The medusae produce eggs. 

c. The eggs produce infusoria. 

d. The infusoria fix and become zoophytes. 

4th. The case of Li%%ia and Sarsia. 

a. The medusa produces medusae by gemmation. 
(The remaining stages as yet unobserved, but probably) — 

b. The medusae produce eggs. 

c. The eggs produce infusoria. 

d. The infusoria fix as polypes, and produce medusae. 

With such facts — unquestioned facts — before us, it seems to me that we have no choice 
between theories, and that we must admit the idea of " Alternation of Generations" to be true. 
Steenstrup was assuredly the first naturalist who announced that idea as a general fact 
dependent on a law. " The special subject of this Essay" — I quote from the author's preface 
to the German version of his celebrated work, as translated by Mr. Busk — " is the funda- 
mental idea expressed by the words '■Alternation of Generations,' or the remarkable, and 
till now inexplicable, natural phenomenon of an animal producing an offspring, which at no 
time resembles its parent, but which, on the other hand, itself brings forth a progeny, which 
returns in its form and nature to the parent animal, so that the maternal animal does not meet 
with its resemblance in its own brood, but in its descendants of the second, third, or fourth 
degree of generation ; and this always takes place in the different animals which exhibit the 
phenomena in a determinate generation, or viith the intervention of a determinate number of 
generations. This remarkable precedence of one or more generations, whose function it is, as 
it were, to prepare the way for the later succeeding generation of animals destined to attain 
a higher degree of perfection, and which are developed into the form of the mother, and 
propagate the species by means of ova, can, I believe, be demonstrated in not a few instances 
in the animal kingdom." 

The main position thus stated appears to me sound and true : the assumption of a definite 
regularity in the alternations is a secondary and non-essential one, and true probably when 
disturbing conditions are not at work. But numerous observations, especially those of 
Dalyell,* Reid,t and Price,^ show that under peculiar circumstances, in what may be termed 
unnatural situations, the polype generations may go on continually producing polype genera- 
tions ; and those of Sars and myself, on the other hand, that a Medusa generation may go on 
producing Medusa generations ; although, under normal conditions in each instance, there is 
every reason to suppose that zoophytic and Medusoid forms would have regularly alternated. 

I am anxious to bear testimony to the value of the idea enunciated by Steenstrup, 
because I believe it has given a strong impulse in a right direction to Invertebrate Zoology. 

* Remarkable Animals of Scotland. 

t Annals of Nat. Hist., Second Series, vol. i. 

+ British Association Report, 1846, p. 86. 



84 CLASSIFICATION OP MEDUS.^. 

It is not a vague generalization founded merely on book-reading, but an induction interpreted 
by a naturalist combining the philosophic spirit with the requisite obsersdng power, — equally 
capable of, and practised in, minute specific research and speculative studies. The Ray 
Society did great service to British science when it sent out a translation of the remarkable 
essay alluded to under the able superintendence of Mr. Busk. This, I am sure, the most 
severe critic upon Steenstrup who has yet appeared — my distinguished and learned friend 
Dr. Carpenter — would be the first to admit, and it were greatly to be desired that some other 
critics on the publications of our Society had a tithe of his knowledge, reasoning power, and 
gentlemanly spirit. 

In a review, " on the Development and Metamorphoses of Zoophytes,"* devoted chiefly 
and most honorably to rendering justice to the untiring labours of Sir John Graham Dalyell, 
the worthy representative of Spallanzani among living naturalists. Dr. Carpenter has opposed 
in strong terms the views of Steenstrup, and, it seems to me, has not done justice — uninten- 
tionally without doubt — to the labours and theory of the Danish naturalist. 

Thus the omission in limine of the name of Steenstrupf in the list (" Sars, Siebold, 
Loven, and Van Beneden") of principal practical continental observers of the phenomena upon 
which the theory of that author, and the new interpretation proposed by Dr. Carpenter, are 
based, is not right, since it conveys the impression to the reader that the Danish zoologist 
theorized on this subject from the researches of others only ; whereas, in reality, some of the 
most valuable observations on the polypiform transformations, were those made by Steenstrup 
himself, detailed in the second chapter of his Essay — that on the " development of the claviform 
polypes." In fact, so far as the subject of this Monograph is concerned, the observations 
referred to, and those of Dujardin (whose name has also been inadvertentl}' omitted by the 
reviewer), are more important than any others in establishing the affinity of the naked-eyed 
Medusa with the Corynoid Polypes. Moreover, the discoveries and researches among the 
Entozoa, announced in the ' Essay on the Alternation of Generations,' are the fruits of its 
author's special obsen^ation, and among the strongest pillars of the edifice which he has built. 
Let not any one suppose, then, that Steenstrup ingeniously constructed a mere closet-theory. 
I doubt much whether any hypothesis or theory in natural history of any value in fostering the 
progress of the science — and may it not be said, too, of other sciences of observation ? — was 
ever eliminated, otherwise than as a dim dream, — dimmer to its author often, than even to other 
men, — by any one not a practical worker in the field where he would raise his specvilations ; 
not merely an occasional visitor, but a day-labourer in science. Goethe has been cited as an 
objection, but Goethe himself would have rejected with indignation the reputation of being a 
discoverer of laws in natural history, without having undergone a severe apprenticeship of 
jiractical study. The great poet who so clearly enunciated the morphology of the vegetable 
individual (unaware of the previous and clear, though premature as to time, announcement of 
the law by Linnfeus), and attempted to work out a like idea in the vertebrate skeleton, warmly 
contended that the doctrines he put forth were not sudden inspirations and lucky guesses, but 
the results of long continued and laborious study. The names which shine brightest in our 
science for their elucidation of its philosophy from the time of Aristotle to that of Linnaeus, 

* Britisli and Foreign Medico-Cliirurgical Eeview, No. I (1848). 
f Log. cit., p. 10. 



CLASSIFICATION OF MEDUSiE. 85 

and from that of Linnaeus to the epoch of Robert Brown and Cuvier, are those of practical 
naturahsts. In their names, and in the names of many eminent men, happily enrolled in the 
list of the Ray Society, I protest against the doctrine that naturahsts (properly so called) are 
only to " record exactly what they sec, and leave it to otlicrs to estimate the value of their 
facts, and to build upon them such inferences as they may think proper."* 

Dr. Carpenter remarks, on the theory of Steenstrup, as follows : " We regard this as a 
very premature, erroneous, and limited expression of the real facts ; and shall endeavour, in 
the course of our exposition, to show what is the real truth of the matter. The proposition, 
in the form enunciated by Steenstrup, is totally inapplicable to the vegetable kingdom ; and a 
strong suspicion of its incoiTCCtness is suggested by that simple circumstance, inasmuch as it 
is chiefly based upon the phenomena presented by those tribes of animals which have most 
in common with plants in their general structure and history." 

The reader of this passage might suppose, if he had not previously read Steenstrups 
Essay, that the Danish naturalist had not taken the phenomena of the vegetable kingdom into 
consideration when stating his proposed law ; nor do I find in the review, although the 
phenomena of vegetation are abundantly cited in favour of the reviewer's opinions, any 
reference made to the fact, that Steenstrup had not only cited them in illustration of his theory, 
but regarded them as presenting the strongest evidences in its favour. The last passage of 
his concluding chapter — that " on the real nature of the alternating generations," runs as 
follows : — 

" I conclude with the remark, that, inasmuch as in the system of ' nursing^ the whole 
advancement of the welfare of tlie young is effected only by a still and peaceful organic 
activity, is only a function of the vegetative life of the individual, so also, all those forms of 
animals in whose development the ' nursing'' system obtains, actually remind us of the 
propagation and vital cycle of plants. For it is peculiar to plants, and as it were their special 
characteristic, that the germ, the primordial individual in the vegetation or seed, is competent 
to produce individuals which are again capable of producing seeds or individuals of the 
primary form, or that to which the plant owed its origin, only by the intervention of a whole 
series of generations. It is certainly the great triumph of morphology, that it is able to show 
how the plant or tree (that colony of individuals arranged in accordance with a simple 
vegetative principle or fundamental law) unfolds itself through a frequently long succession 
of generations, into individuals becoming constantly more and more perfect, until, after the 
immediately precedent generation, it appears as calyx and corolla, with perfect male and 
female individuals ; stamens and pistils — and after, the fructification brings forth seed, which 
again goes through the same course. It is this great and significant resemblance to tlie 
vegetable kingdom, which, in my opinion, is presented by the entoxoa and all nurse genera- 
tions {amme), and to which I have alluded in the preceding Essay ; I might almost say, that 
the condition of continued dependence incidental to the animal life, is, to a certain extent, one 
of less perfection than that which is presented in the progressive elevation in development 
effected by the agency of the vegetative life."t 

This remarkable passage had surely escaped the notice of the reviewer ; for in it the 

* Loc. cit., p. 7. 

t Ray Society's Translation of tlie Alternation of Generations, p 114. 



8C CLASSIFICATION OP MEDUSA. 

argument drawn from analogy between the animal and vegetable kingdom, is as clearly and fully 
expressed as in the sentences I am about to cite from the review itself. Indeed it was very 
unlikely that a Lecturer on Botany* would allow such obvious analogies to escape him. 

The theory enunciated by Dr. Carpenter, and proposed to be substituted for that of 
Steenstrup, has special reference to the Medusae and Polypes, and is stated in the following 
passages : 

" The fertilized ovum of the medusa-pai'ent is like the seed of the plant ; and the polype 
that grows from it resembles tlie first leaf-bud into which the embryo expands. From this bud 
are at first produced others, by the process of continuous growth, which are repetitions of 
itself ; these in the plant usually remain connected with each other so as to form a compound 
structure, and so they do also in the ordinary zoophyte ; but in the common hydra, and in 
the hydraform medusa-larva, they become detached like the bulbels of the marchantia or lily. 
But under certain conditions, a new and different set of buds, containing a sexual apparatus, 
are produced ; these, too, become detached, and, by their inherent powers of movement, they 
convey the germs of a new generation to a distance from the parent stock. The whole of these 
phenomena appear to us to constitute but a single generation, instead of two, as represented 
by Steenstrup. We are not in the habit of speaking of the leaf-buds and the flower-buds of a 
l)lant as of two distinct generations ; nor, if our comparison be correct, have we any ground 
for giving such a designation to the polypoid larva, and the medusa-imago, which are con- 
tinuous developments from the same germ. Hence the whole doctrine of the ' alternation of 
generations,' falls to the ground, so far as this individual case is concerned ; the phenomena 
being simply those of metamorphosis or change of form, attending the evolution of successive 
products from the same original germ. The metamorphosis is not really so great as that 
which presents itself in the course of the development of any one of the higher organisms, 
tiie several parts of which depart more widely from each other, and from the early embryonic 
cell-cluster, than do the polype-buds and medusa-buds we have Ijcen describing. The chief 
difference lies in the capacity of the latter to maintain a separate and independent existence ; 
a capacity which is evidently connected with tlieir low type of organization." (Review 
cited, p. 23.) 

And again, in the recapitulation (at p. 29) — " The true Ilydra, which may be regarded as 
uniting the general form and structure of the polype with the locomotive powers and dis- 
positions of the medusa, propagates in both the modes characteristic of the vegetable 
kingdom ; namely, by gemmation, and by the production of ova. The huds are not destined 
to remain in continuity with the parent, but are thrown off like the bulbels of certain plants ; 
having previously acquired, however, the form of the parent. The ova also are developed 
into polypes resembling the parent. The usual mode of propagation is here by bulbels ; the 
ova being destined apparently to continue the race through the winter season, the cold of 
which might be fatal to the parents. 

" In other cases, however, we find a greater specialization of characters ; the locomotive 
and proper generative apparatus being especially developed in the Medusce ; whilst the true 
polypoid condition presents its most complete evolution in the plant-like Sertularida, yet 
these two groups are not to be dissociated from one another ; for each of them, in one of its 

* PrcfLSsijr Steenstrup was Lecturer on Botany and Mineralogy in the Academy of Sorije. 



CLASSIFICATION OF MEDUSA. 87 

stages of development, presents the characters of the other. The Medusa begins life as a 
polyjje ; as a polype it is attached ; as a polype it grasps and digests its food ; as a polype it 
reproduces parts that have been removed ; and as a polype it propagates by gemmation ; the 
buds being detached from the parent as soon as they have acquired the form of the latter, 
and are capable of maintaining an independent existence. But in this condition it forms no 
ova. A new and distinct series of buds (flower-buds) is produced for this purpose ; these 
buds are detached like the preceding ; they become developed into perfect Medusae, in which 
state alone they have been known until recently ; and from these Medusae are produced ova 
by a true sexual process, which are first evolved into the polypoid form, and go through the 
series of changes just enumerated." (p. 30.) 

In this theory, proposed by Dr. Carpenter for adoption instead of that of Steenstrup, I 
can see only verbal differences. The main facts upon what Steenstrup built his proposed 
law are not denied. It is admitted that a polype may produce a Medusa by gemmation, and 
that the egg of the Medusa may produce an animal altogether different from itself, Init like 
the polype which produced it. The reviewer admits an alternation of forms, but he denies 
that they are generations which alternate. Yet in the ordinary sense of the word generation, 
as here applied by Steenstrup, they must either be such or be the same individual. A father 
belongs to one generation, a son to a second, a grandson to a third — at least, this has been the 
case hitherto, as far as my knowledge goes. Surely the first polyjie represents one generation, 
the offspring of that polype a second, and the offspi'ing of that offspring a third. And if so, 
the middle term here being a Medusa and not a polyjie, and its offspring a polype again, 
which produces a Medusa, we are warranted to speak of an alternation of dissimilar genera- 
tions. It does not affect the question, if they be regarded as individuals, whether they are 
produced by gemmation or from ova ; nor whether we hold with Sars that we have not an 
alternation of animals of distinct classes, but of fixed and free animals of the same class. It 
is not the less an alternation of dissimilar generations. 

But when Dr. Carpenter says that the phenomena are simple metamorphoses not really 
so great as those which present themselves in the course of the development of any one of 
the higher organisms — " the several parts of which depart more widely from each other, and 
from the early embryonic cell-clusters, than do the polype-buds and Medusa buds we have 
been describing" — he may mean, that the Medusae produced by gemmation are not distinct 
individuals, but parts of some one capable of maintaining a separate and independent 
existence. If so, he has certainly enunciated a new theory altogether distinct from that of 
Steenstrup, but one so opposed to ordinary notions of individuality among the lower animals, 
that few, if any, naturalists will assent to it until more fully and satisfactorily stated. 

To argue that, because " we are not in the habit of speaking of the leaf-buds and the 
flower-buds of a plant as of two distinct generations," we are, therefore, not to regard the 
alternations of polypes and Medusae as such, is to bring the common, popular, unscientific, 
and untrue notion of the nature of a plant into a scientific discussion on the nature of animals. 
We are not in the habit of regarding a leaf as an individual — popularly, we look upon the 
whole plant as an individual. Yet every botanist knows that it is a combination of individuals 
and if so, each successive series of buds must certainly be strictly regarded as generations. 
The first generation of a Lupine, for instance, is the pair of individuals constituting the 
cotyledons of the embryo, dissimilar from the second generation, which consists of the several 



88 CLASSIFICATION OF MEDUSiE. 

phytons comprising the first bud or plumule. A series of similar buds may be produced 
until one of different aspect is developed, composed of a generation of altogether different 
individuals, through whose agency the foundation of a new series of generations is laid in the 
formation of the ovum or seed. Whether we style the members of one generation nurses, or 
call them all by the same name, does not matter so far as the fact and law of an alternation 
of generations is concerned. 

I see no reason therefore to dissent from the theory of Steenstrup ; it is the simplest and 
most intelligible, as well as most original expression hitherto offered of the astonishing facts 
which he was the first to generalize. Granting it, we can no longer adopt the usually 
accepted classification of Radiate animals, nor separate them into Echinodermata, Jcalephee, 
Zoophyta, and Sponges, as so many distinct and equal orders ; but must unite the Acalepha 
with the Zoojjhi/ta, excluding from the latter the Bryoxoa which are polypoid Tunicata. 
The Acalep)h(e or Aruchnodermata must undergo reconstruction, for the Polypes cannot even 
be regarded as forming a primary division when united with the usual members of this great 
section. They evidently form part of a sub-class with the D'lscojihorce, equal to the sub- 
classes, CiUograda, Cirrhigrada, and Phjsograda. The Bhcophorm must again undergo 
subdivision into orders. The Antlioxoa will stand first, next the Steganopthalmata, then the 
Gymnopthalmata, and lastly the Hijdroida. That the Anthoxoa are intimately related to the 
Medusae is evident to any unprejudiced naturalist who has studied the structure of Lucernaria, 
or of the Actineada, especially of any floating form of the last tribe, such as the Arachnactis 
of Sars. The close affinity of these tribes has been excellently treated of in an Essay by 
Drs. Frey and Leuckart, who, after comparing organ with organ in the Anthozoa, the several 
usually received orders of Aculephm and the Polypes, observe, in conclusion, that these various 
tribes ought no longer to be placed apart in a natural system. "They rather go towards 
constituting a larger section, having one common type of structure — a type chiefly charac- 
terised by the peculiar arrangements of the viscera and the stomachal cavity." They 
propose to designate such division by the name of Colenterata.* 

Even among the animals figured and described in this Monograph, we see abundant 
evidences of the close affinity of the Medusae, on the one hand, with hydroid poljqies ; on the 
other, with the Anthozoa. The Steenstrupice are in all probability Medusa-generations of some 
corynoid polype, yet, through Euphysa, they are intimately related with Sarsia, and through 
Sarsia with Slahberia, whence the affinities upwards are easily traced. The 'Purris digitalis, 
on the other hand, closely reminds us of an Actinea ; so nearly, that when I first found a 
specimen, I mistook it for an animal of that genus. 

Thus, in the end, we revert, curiously enough, to the views of the affinities of these 
animals proposed by Aristotle, who plainly included, under the designation of aKa\v,pi,, both 
ActinecB and Medusa ; not from any vague guess, or in compliance with the popular recognition 
of their resemblances, but from a careful study of their structure and habits, as the varied 
notices of them preserved to us in the first, fourth, fifth, eighth, and ninth Ijooks of the ' History 
of Animals,' prove beyond question. 

* Frev and Leuckart's Beitrage, p. 38. 



I shall conclude with a few remarks on the best methods of studying and preserving the 
Naked-eyed Medusae. 

They are to be sought for in summer and autumn, when the weather is warm and dry, and 
the sea calm and clear. They abound, within reach, mostly in the afternoon and towards night- 
fall — probably also during the night, though not then so near the surface of the water. A small 
bag of fine muslin, attached to a metal ring, is the best instrument by which to take them, 
and may be used either as a hand-net fixed to the end of a stick or pole, or as a tow-net 
suspended over the stern of a vessel, when at anchor, or making very gentle way through 
the water. My friend Professor Acland took great numbers at Oban, by attaching a tow-net 
to the buoy in the bay, and leaving it there during the night. They abound most in sheltered 
bays near strong tideways or headlands projecting into the Atlantic. The majority 
being oceanic, they are most numerous and varied on those parts of our shores which are 
touched by oceanic currents. Hitherto the Zetlands, Hebrides, and coasts of Cornwall have 
yielded the greater number of species. Many new forms may be expected to occur on the 
Atlantic coasts of Ireland. Indeed, I fully expect that the number of British species will be 
doubled within the next ten years, now that attention is directed to these beautiful little 
animals. 

When the tow-net is taken out of the sea it is to be carefully reversed, and its contents 
gently emptied into a basin or glass jar, filled with clear salt water. It is best to plunge the 
net beneath the surface when being emptied, as thus the Medusae are enabled to detach 
themselves from the threads, and swim away without injury. When the net is out of the water, 
they appear like little, adhering, shapeless masses of clear jelly, and exhibit no traces of their 
elegant form and ornaments. When in the jar or basin, they are often, on account of their 
extreme transparency, very difficult to distinguish, but by placing the vessel in the sun, or 
beside a strong artificial light, we see their shadows floating over the sides and bottom of the 
basin, like the shadows of flitting clouds on a landscape. These soon gviide us to the creatures 
themselves, and before long we distinguish their ocelli and coloured reproductive organs. 
The next step is to secure such as we wish to examine closely, and transfer them to watch- 
glasses or small glass tubes. To do this is often not an easy task, for when alanned they 
are extremely agile and alert ; so that if we attempt to capture them with a teaspoon, they 
usually escape us, or if taken, by their slippery nature, slide out of the spoon whilst we pour 
away the superabundant water. This difiiculty may be got over by using a small but deep 
glass spoon, with its handle set very obliquely. When we have placed some in a glass tube 
with a little water, or in a small compressed glass jar, which I find an excellent aid in exa- 
mining them, we can observe their profile, the changes their body undergoes when 
contracting and expanding, and the extent to which the creature can lengthen its tentacula. 
We then place them in a watch-glass and submit them to microscopic examination, 
carefully noting the number, colour, form, and structure of the ocelli and tentacula, the 
arrangement of the gastro-vascular canals and reproductive glands, and the form and 

13 



90 PRESERVATION OF MEDUSA. 

structure of the central peduncle. Convepng our prizes then to a dark place, we irritate 
them, and observe whether they phosphoresce or not, of what colour the light is, and how 
long it endures. 

In every case a drawing, as careful and detailed as possible, and always coloured, should 
be made at the time. This is the more necessary, since they are animals extremely difficult 
to preserve, shrivelling up into indistinguishable curd-like masses in spirits, and most 
preserving fluids. In fact, the only specimens which I have seen preserved in a distinguishable 
state, have been so by means of one of Mr. Goadby's fluids. When Mr. Goadby accompanied 
Mr. M'Andrew, in 1 837, on a cruise among the Hebrides and Zetlands, he made many 
experiments on the preservation of these delicate creatures, and succeeded so well that I have 
been able to distinguish among them even the several species of the critical genus TJiaumantias . 
I do not despair of seeing, before very long, a series of these creatures so preserved exhibited 
in the British Museum, and contributing to render more perfect the finest natural-history 
collection in the world. The indefatigable director of the zoological department in that truly 
national establishment, will yet, I trust, sanction such an addition, and, as he no longer 
remains a sceptic in bones or disbeliever in spirits, may consistently extend his faith to 
Goadby's fluid. 

Note. — 'Kindly communicated by Mr. Goadby. 

To preserve the Acalephm. — These animals contain so large a quantity of water, that they 
reqviire great care and attention to preserve them. 

The B fluid of itself is not enough for the purpose, the assistance of alum being imperatively 
necessary to give firmness and support to the several tissues. 

The plan that I adopted with great success was the following : i. e. 

Make a saturated solution of bay salt, and when cold, test it with a specific-gravity bubble 
prepared for that purpose. 

When required for use, dilute it (with water) to 1148, indicated by another bubble so marked. 
To this latter fluid add alum, at the rate of =ij to every quart of fluid, aud dilute the whole to ^ strength 
with water. Pour this into a dish, and empty the contents of the tow-net (containing the well-draiued 
specimens of Acalephse) into it, and let them macerate therein for twenty-four hours, by which time 
they will be found saturated with the fluid, and at the bottom of the dish. If the specimens be small, 
they shoidd now be moved, and placed in fluid consisting of dissolved bay salt, only to the strength of 
1148, as the alum destroys transparency. 

Large specimens of the Acalephae, as Aiirelia, &c., off'er exceptions to this rule ; they may be 
allowed to remain in aluminous fluid (to be changed daily) of the strength described, or somewhat 
increased by additions of stronger saline fluid, for a longer period (two or three weeks), but ultimately 
they, too, must be removed from the continued influence of alum, and kept in the bay salt fluid, of not 
less strength than before described, viz. 1148. 

The fluid should be tested with the bubble daily, aud its strength made up by additions of the 
saturated solution, until it obey the test for several consecutive days, when endosmose and exosmose being 
at an end, the process of preservation may be considered complete. 

For permanent preservation, corrosive sublimate should be added to the preserving fluid, in the 
proportion of grains ij per quart of fluid, but its use is unnecessary in the early stages Of preservation. 
I did not employ it until the collection of last summer was complete, and on shore. Neither is it 
essential to filter the fluid if time be pressing : at sea, of course, it cannot be done. Finally, marine 
animals require for their preservation saline fluid of the specific gravity of 1148. Fresh water and 
terrestrial animals are preserved at the diminished strength of 1100; fluids of less strength (respectively) 
are insufficient, and greater strength is injurious. 



BIBLIOGRAPHY. 



In order to facilitate the studies of those among my readers who may be inchned to 
pm'sue researches among the Pulmograda Gymnopthalmata, I have drawn up the following 
catalogue raisonne of authors, works, papers, and figures bearing upon the subject, not 
confining the notices to British species only, but extending them to all described or figured 
forms of Naked-eyed Medusae. 

1739. Janus Plancus. ' De Conctis minus notis Liber.' 4to, Venice. 

In this is contained tlie first figure of the Carybdea marsupialis, — and a very miserable 
representation it is, — in Plate iv, f. 5, /, under the description of " Urtica soluta 
Marsupium referens et motus vitaleis manifestissime edens Maris Ariminensis." The 
figure, plate xcii, fig. 9, of the ' Encyclopedie Methodique,' is sometimes quoted as if it 
had been taken from Plancus, and represented the same species (2d ed. of Lamarck, 
An. sans Vert., vol. iii, p. 131), but is really copied from Slabber, and represents in all 
probabiKty an Oceania. 

1746. Linnaeus. 'Fauna Suecica,' 1st Ed. 

In the first edition of this work three Medusae are enumerated, of which the third is a 
naked-eyed species. " 1288. Medusa orbicula cruce alba picta. Hsec omnium minima 
est, tota gelatinosa vitrei colons, discum pingit crux magna alba, ad margines usque 
extensa, margo integer est : caret appendicibus omnibus, sc. cavitatibus, pistUlis, 
staminibus, branchiis." p. 368. 

In the second edition the specific name Criiciata is added. 

1758. Linnaeus. ' Systema Naturae.' 10th Ed. 

Medusa cruciata, M. pilearis, and M. marsupialis, are the naked-eyed species enumerated. 

1758. The Rev. W. Borlase published his ' Natural History of Cornwall,' in which there are some of 
the earliest figm'es of British ISIedusie, but no naked-eyed species is represented by him. 

Pennant (British Zoology, vol. iv, 1787) and Turton (British Fauna, 1807) contented 
themselves with following Borlase, making no additions to his list. 

1760. L. T. Gronow (Gronovius). " Observationes de Animalculis aliquot Marinae Aquse Innatantibus 
atque in Littoribus Belgicis obviis," in the 'Acta Helvetica,' vol. iv, p. 38. 

In his paper is contained the first notice of Thaumantias /lemisp/iarica. The description is 
full and good for its time ; the figm-e bad, and scarcely recognisable. Cydippe pileus is 
described and figui'cd in the same paper. 

1775. P. Forskal. ' Descriptiones Animalium quae in Itinere Orientali observavit Petrus Forskal. 
Post mortem auctoi'is edidit Carsten Niebuhr.' Havnife, 1775. 
Forskal observed and described twelve pulmograde Medusae during his voyage. Of these, five 
were inhabitants of the Eed Sea, and the remainder of the MediteiTanean. Among the 



92 BIBLIOGRAPHY. 

latter are four naked-eyed species, viz. M. j'roboscidalis (i. e. Dianaa proboscidalis) 
M. mollicina (Genus?), M. pileata {Oceania pileata), M. aqiiorea (i. e. ^quorea 
Forskalina). His descriptions and drawings are very characteristic. The latter were 
pubhshed in a separate volume, of ' Icones.' 

1776. Otho Frederic Miiller. ' Zoologise Danicse Prodromus, sen Animalium Daniae et Norvegiae 
Indigenarum Characteres Nomina et Synonyma.' 8vo, Havniae. 
Eight species are enumerated under the genus Medusa in this work. One of them is the 
Medusa palliata of Bohadsch, which is an Actinea, being the Adamsia maculata of British 
naturalists. Three appear to be naked-eyed Medusre, viz. M. hemisphcerica, M. bimorpha, 
and M. diyitale. The first is the Tliaumantias hemispharica ; the second and third were 
communicated to Miiller by Otho Fabricius, the latter being our Turris digitalis. 

1778. Martin Slabber. ' Natuurkundige Ver-Gustigingen.' 4to, Haarlem, 1778. 

This work contains several figures of Medusae. Plate ii, figs. 1 and 2, are very bad figures 
of Saphenia dinema : six vessels are represented instead of four. Plate xii, figs. 1 and 2, 
is the Thamnantias cymbaloidea of authors, and in all probability a bad representation of 
T. hemisp/usrica. Plate xii, fig. 13, is either Turris neglecta, or an allied species. Plate 
xiv, fig. 1, appears to be an Oceania. The two latter are the Oceania tetranema, and 
O. sanguinolenta of Peron and Leseuer. 

1780. Otho Fabricius. ' Fauna Groenlandica,' 8vo, Hafniae et Lipsiae. 

Of the Medusie described in this excellent work, M. digitale is our Turris digitalis; 
M. bimorpha and M. campanula appear also to have been naked-eyed species, though the 
latter may possibly have been a young Cyancea. 

1788. Olof Swartz, on " Medusa unguicidata wad Actinea pusilla," in the 'New Transactions of the 
Royal Swedish Academy,' vol. ix. 
The Medusa here described is probably a naked-eyed species, though I confess I do not 
clearly understand the figure (not badly executed) given. It is the Linuche unguicutata 
of Eschscholtz. Actinea pusilla appears to be a floating animal of its tribe, and possibly 
a species of Arachnactis. Eschscholtz and Lesson make a Medusa of it under the name 
of Melicertum pusillum. 

1788. J. F. Gmelin. The 13th edition of ' Systema Naturae,' of Linnaeus. 

The Medusae are contained in vol. i, part 6 of this compilation. Such naked-eyed species 
as are given (viz. Medusa marsupialis, M. hemispliarica, M. dimorplia, M. campanula, 
M. digitale, M. proboscidalis, M. mollicina (?) and M. pileata), are taken from Plancus, 
Miiller, Otho Fabricius, and Forskal. 

1788-9. O. F. Miiller. ' Zoologia Danica.' 

A good figure of Thaumantias fiemispjharica is contained in this excellent and most useful 
work. 

1791. Adolph Modeer. " Om Slagtet Siokalf, Medusa," in the ' Nya Handlingar,' of the Royal 
Swedish Academy, vol. xii. 

A synopsis of the Medusae known up to that time, and a very valuable one for its date. 
The naked-eyed species described by Forskal, Otho Fabricius, &c., are enumerated and 
characterised with great acuteness. 

1791. The collection of figures of Medusae in the six plates (pi. xc-xcv) devoted to Acalephte, in the 
' Encyclopedie Methodique,' contains several naked- eyed species. 
Plate xcii, figs. 9 and 10, are copied from Slabber, and represent a Turris and an Oceania; 
fig. 11 is Oceania pileata, copied from Forskal; figs. 12-15 are copies of Slabber's figure 
of Obelia sphcerulina, and figs. 7-8 of his Medusa perla, both, however, evidently, as we 
have seen, the figs, of higher Medusae. In plate xciii, we have fig. 1 representing Diatuea 
proboscidalis, copied from Forskal; figs. 2-4 is Thaumantias cymbaloidea, from Slabber; and 



BIBLIOGRAPHY. 93 

8-11, Thaumantias hemuphmrica, from Miillcr; 5, 6, 7, are copies of the "Medusa 
cntciatu," of Porskal. In plate xciv, figs. 4, 5, are copies of the figures given by Bastcr, 
of tlie Medusa whicli has received the name of Callirhoe Busteriana. Plate xcv, figs. 
1 and 2, are the jEquorea moUicina ; fig. 4, Mesonema calum-pensile ; and fig. 3, JEquoreu 
Forskalina, all copied from Porskal. 

1809. Peron et Lesueur. " Tableau des Caracteres generiques et specifiques de toutes les especes de 
Meduses connues jusqu'k ce jour," in the ' Annales du Museum d'Histoire Naturelle,' 
vol. xiv. 

A standard paper. Unfortunately the plates and figures referred to in this valuable memoir 
have never been made public, so that it is beyond the power of the British naturalist to 
determine the species mentioned as inhabitants of the Channel, for the descriptions are 
too often insufficient. The following genera of naked-eyed Medusse, are characterised 
for the first time in this paper : Eudora, Berenix, Orythia, Favonia, Lymnorea, Geryonia, 
Curybdea, Phorcynia, Eulimenes, JEquorea, Foveolia, Pegasia (?), Callirhoe, Oceania, 
Aglauru, Me/icerta (?), Euryale. The names of the species will be found in the table 
which I have constructed from Lesson, further on. An issue of the original plates would 
be a great boon to science, as few naturalists have had such opportunities of observing the 
MedusEe in all parts of the world. 

1816-18. De Lamarck. ' Animaux sans Vertebres.' 

The iledusse are described at second-hand. The naked-eyed species are arranged under 
the genera Eudora, Phorcynia, Caryhdea, jEquorea (?), Callirhoe, Orythia, and Diancea. 
Peron and Lesueur are evidently the chief source of the descriptions. 

1821. A. de Chamisso, and C. G. Eysenhardt. " De Animalibus quibusdum e Classe Vermium 
Linneana in circumnavigatione Terrse, auspicante Comite N. Romanzoff, duce Othone 
de Kotzebue, annis 1815-18 peracta, observatis;" in the ' Acta Academise Naturte 
Curiosorum,' vol. x. 

Several Medusa; are represented in the plates to this paper. Of these, one, the Geryonia 
tetraphyUa, is a naked-eyed form, allied to our G. appendiculaia, and resembling it 
in having eight tentacles alternately differing in size. Their structure is not given. The 
extremity of the peduncle is represented as having a round orifice, which is a mistake, as 
in the description, the peduncle is said to be " bipollicaris, cylindricus, flexilis, apice 
(ore) truncato dUatato quadrivalvato membranaceo, maculis quatuor viridibus notato." 
It inhabits the Indian Ocean. (Loc. cit., t. xxvii, f. 2.) 

The " Medusa campanulata" of this paper (pi. xxx, f. 1) seems to me to be a mutilated 
animal, doubtfully of this division, and the M. niucilaginosa is possibly a mutilated 
Polyx'enia. Both are from the Pacific Ocean, and the imperfection of the drawings is due 
to the specimens, and not to the describers, as they expressly state their doubts respecting 
the generic afBnities of both forms, and suggest the necessity of fresh observations. 

1821. Quoy and Gaimard. 'Zoology of Voyage of the Urania and Physicienne' (under Freycinet). 
Plates Ixxsiv and Ixxxv are devoted to the Medusae. 

Of naked-eyed species there are figured jEqiwrea grisea (Admiralty Isles), JEquorea 
cyanogramma, from the same locality, yJSquorea punctata, from between the Philippines 
and Sandwich Isles, and ^quorea semirosea, from New Guinea; all species well marked 
by peculiarities of colour. Dianma balearica (a Geryonia ?), from the western Mediterranean, 
a two-tentaculated species, remarkable for its thick peduncle (?). Diantea endractensis, 
a six-tentaculated species of a reddish tinge, from New Holland. In neither the figures 
nor descriptions of these are the ovaries definitely stated. The introductory remarks show 
that the authors did not very clearlj' comprehend what they saw. 

1826. Risso. ' Histoire Naturelle de I'Europe Meridionale.' 

The Medusfe of the neighbourhood of Nice are enumerated in the fourth volume, including 
several known naked-eyed species. The author's knowledge appears to have been very slight. 



94 BIBLIOGRAPHY. 

1827. MM. Quoy et Gaimai-d. ' Observations Zoologiques faites a bord de I'Astrolabe, en Mai 1826, 
dans le detroit de Gibraltar.' 

In this valuable paper a number of radiate animals, chiefly pelagic, are described, and among 
them several naked-eyed Medusae. The species are all figured, though mostly with few 
or no anatomical details. 
1 . Dianaa rotunda, p. 181, pi. vi. A, figs. 1 and 2. 

Judging from the view given of the peduncle, this appears to be a true Oceania, with a globular 
umbrella and eight marginal tentacula. 
3. Diancea conica, p. 182, pi. 6, A, figs. 3 and 4. 

The form of the body indicates a Circe, but the appearance of the peduncle is nearer that of 
Oceania, and the remark of the describers, that it approaches the Medusa {Oceania) pileata 
of Forskal, would confirm such a view. The umbrella is mitrate, and acute above. The margin 
appears to have twenty tentacula, with red ocelli. The peduncle is reddish. 

3. Dianeea exii/ua, p. 183, pi. vi. A, figs. 5 and 6. 

A small Geryonia, with very small cordate ovaries, and four marginal tentacula. 

4. Diaiuea exigua, Var., p. 183, pi. vi. A, figs, 7 and 8. 

Exactly like the last, but wanting the ovaries. Is this a male animal, or is it a Tima ? It is 
the Liriopa cerasiformis of Lesson (Acal. p. 332), who strangely associates it with Diancea 
proboscidalis, in his genus Liriopa. 

5. Dianaa bitentaculata, p. 184, pi. 6, A, fig. 9. 

A minute Geryonia or Tima, having two long tentacles and twelve short ones. This is the 
Saphenia bitentaculata of Lesson. 

6. Dianoea funeraria, p. 184, pi. \-\, A, figs. 10-15. 

This appears to belong to a genus closely allied to Circe, and is certainly a member of the family 
Circeadce. It is the Tholus funerarius of Lesson. 

7. j^qtwrea capillata, p. 185, pi. vi, B, fig. 1. 

Too imperfectly described and figured to be assigned to any well-defined genus with certainty. 

8. Phorcynia pileata, p. 186, pi. vi, C, fig. 1. 

A mutilated or badly-observed species, of what genus ? It is the type of Lesson's genus 
Pileola. 

1828. Dr. Fleming. ' History of British Animals.' 

Under the genus Geryonia are enumerated " G. cequorea," " G. hemispharica" {Thaumantias) , 
and " G. octona" {Oceania). The last previously described by Dr. Fleming, in the eighth 
volume of the ' Edinburgh Philosophical Journal.' 

1829. F. Eschscholtz. ' System der Acalephen.' 4to, Berlin. 

A standard work vipon this class, founded on extensive personal research. I have already 
commented on the errors of the classification. The illustrative figures are in outline, 
but strikingly faithful, so far as they go. The naked-eyed species represented are Melicertum 
penicillattim ; Eurybia exigua ; Tima flavilabris ; Cytceis tetrastyla ; Cunina campanulata 
and globosa ; yEguorea ciliata ; Polyxenia cyanostylis ; yEquorea globosa (a Stomo- 
brachium ?) ; jEgina rosea, and citraa ; Mesonema abbreviata ; Geryonia bicolor and rosacea. 
The descriptions are in German, each prefaced by a Latin diagnosis, too slight in most 
instances to serve the purpose of identification. No student of the Medusse should be 
without this book. 

1830. Lesson. 'Zoology of the Voyage of the Coquille' (under Duperrey). 

Most of the figures of Medusa; in this work represent covered-eyed species. In plate xiv of 
the Zoophytes, a few naked-eyed species are represented, but though "the plates are 
beautifully engraved and coloured, the original drawings must have been sadly defective, 
judging from the Cyanaea Bougainvillii {BottgainviUea Macloviana), the first that attracts 
our notice, every organ of which is misunderstood, and wrougly delineated. Fig. 4 of 
the same plate represents a Turris, under the name oi ^quorea mitra ; the peduncle and 
ovaries strangely misunderstood. Fig. 1. Bursarius Cytherm may be a naked-eyed form, 
but after the manner in which the two previously-cited species are represented, I cannot 



BIBLIOGRAPHY. 95 

venture to offer an opinion with any approach to certainty. The Dianaa cerebriformis of 
plate X is possibly a Cyaiuea. The Eudorte are evidently mutilated disks. 

1830. ' Magazine of Natural History,' vol. iii. 

Dr. Baird, in an interesting paper " On the Luminousness of the Sea/' figures a small 
Medusa {Geryonia ?) from the Straits of Banca : the figure and notes are insufficient. 

1831. 'Magazine of Natural History/ vol. iv, p. 285. 

In a note on the " Luminosity of the Sea/' by Mr. Samuel Woodward, a very minute 
Medusa of this order is figured from specimens taken between Lowestoft and Yarmouth. 
The figure represents either a young Sarsia or the medusoid of Tubularia. 

1833. Quoy and Gaimard. ' Zoology of the Voyage of the Astrolabe.' (Expedition of Dumont 
d'Urville.) 

The Medusae are described in the fourth volume of the text, and figured in plate xxv. They 
are Carybdea bicolor, possibly an imperfect animal, but well figured ; Caryhdea bitentacuhita, 
and a covered-eyed species, Orythia incolor. Of the two former, the first was found near 
the Cape de Verde, the second, in the roads of Amboyna. 

1833. Milne Edwards on " Carybdea marsupialis." 

A valuable and excellently illustrated paper in the 28th volume of the ' Annales des Sciences 
Natiu-elles.' 

1833. Dr. Johnston. Description and figure of " Diancea Bairdii" {Tima Bairdii, mihi), in the sixth 

volume of the 'Magazine of Natural History' (Loudon's). 

1834. De Blainville. ' Manuel d'Actinologie.' 

The account of the pulmograda in this useful manual is a very full and excellent compilation 
of the knowledge up to the time, and among the plates are many useful figures copied 
from other works. De Blainville combines the genera JErjuorea, Mesonema, Polyweiiia, 
jEgina, and Cunina in one genus, ^quorea. He refers Melkerta penicellata of 
Eschscholtz to the genus Aglaura of Peron. Under Geryonia he unites Saphenia, 
Geryonia, and Diancea proper ; regarding Dianaa endractensis as the type of that genus. 

1834. J. T. Brandt. " Prodromus Descriptionis Aniraalium ab H. Mertensio observatum," in the 
' Recueil des Actes de la Seance publique de 1' Academic Imperiale des Sciences de 
St. Petersbourg,' 1833-4. 

In this valuable paper Brandt gives a synopsis of the radiate animals observed by ^lertens 
during his voyage. The following naked-eyed MedusiE are enumerated : 
Circe, Mertens (the genus characterised). 

1. C, camtschatica, Brandt. Kamtschatka. 
CoNis, Brandt (the genus characterised) . 

2. C. mitratu. Pacific " ab insulis Boninimensibus." 

3. iEQUOREA rhodolema, Brandt. Conception, in Chili. 
Stomobrachiota, Brandt (the genus characterised). 

4. S. lenticularis, Brandt. Atlantic, " ab insidis Malvinensibus. 

5. Mesonema macrodactyla, Brandt. Southern Ocean. 

6. Mesonema ccerulescens, in 50° lat., and 144° long. W. 
jEginopsis, Brandt, (the genus characterised). 

7. JE. horensis, Brandt. In Behring's Straits. 

8. PoLYXENiA flavobracMa, Brandt. 5° lat., 127° long. W. 

9. Geryonia hexaphylla, Peron. Pacific Ocean, in 36° 30' lat. and 211 long., ab ins. 

Boninsimensibus. 

10. Proboscidactyla ^^ar«cwTA«?«, Brandt. Camtschatica. 

11. HippocRENE BougainvilUi, Brandt. Behring's Straits. 

12. Staurophora Mertensii, Brandt. North Pacific. 



96 BIBLIOGRAPHY. 

1835. H. Ratlike, (Professor, of Dorpat). " BescLreibuug der Oceania Blumenbachii, einer bei Sevastopol 
gefundenen leuclitenden Meduse," in the ' Memoires presentes a rAcademie Imperiale 
des Sciences de St. Petersbourg,' vol. ii, with an excellent plate. 

The animal figured and described in this paper is a very remarkable one, and evident!}' 
sui generis. It is not a true Oceania, nor a member of the family Oceanida, but of 
that section of the Sarsiadm which will probably eventually assume the position of an 
independent group including Bouffainvillea and Lizzia. The umbrella is hemispherical, 
bordered by eight compound tentacular bulbs, of a bright yellow colour, from each of 
which rise three filiform, highly extensile, white tentacula. The peduncle is four-lobed, 
broad, striped with white and yellow, and opens by a mouth surrounded by four Ups, with 
tentacular and gland-tipped prolongations arranged in a pinnate fashion. 

1835. M. Sars. ' Beskrivelser og Jagttagelser over nogle moerkelige ellernyei havet ved den Bergenske 

kyst lebende dyr af Polypernes, Acalephernes,' Sic. 8vo, Bergen. 

This very interesting work is in the Norwegian language. In it Phorcynia cruciata and 
Thamnantias hemispheerica are mentioned as Norwegian species, and the following new 
species described and figured : 

1. Oceania ampiiUacea, p. 22, t. iv, fig. 8. 

2. Oceania octocostata, p. 24, t. iv, fig. 9. This is our Stomobrachiiim octo- 
costatum. 

3. Oceania saltatoria, p. 25, t. iv, fig. 10. This appears to be a Circe by its form. 
No ovaries are shown in the figure. It has sixteen tentacula. It is the Pandea 
saltatoria of Lesson. 

4. Oceania (?) tuhulosa, p. 28, t. iv, fig. 11. — Sarsia tubulosa. 

6. Thamnantias nmlticirrata, p. 25, t. viii, fig. 12. 

7. Thamnantias (?) jjlana, p. 26, t. v, fig. 11. 

8. Cytais octopunctata, p. 28, t. vi^ fig. 14. — Lizzia octopunctata. 

1836. In the ninth volume of the ' Magazine of Natural History' (Loudon's) is a " Catalogue of the 

Species of Rayed Animals found in Ireland, as selected from the papers of the late 
J. Templeton, Esq., of Cranmore, with notices of Localities, and with some Descriptions 
and Illustrations by Robert Templeton, Esq." 

This list is of considerable value, and shows that the distinguished naturalist, from whose 
papers it was compiled, had taken great interest in the Acalephs. Of the species he 
enumerates, the following appear to belong to the Pulmograda Gyninopthahnata. 
" Piliscelotus. Body hyaline, hemispherical, the apex somewhat produced, and terminating 
in a fleshy, elongated, spindle-shaped appendix. Margin of the body with four moderately 
long tentacula, each tentaculum arising from a small tubercle. P. viireus (p. 302, f. 48). 
Hyaline, bell-shaped, with four brown tentacula arising from the margin, uearly equidistant ; 
the centre produced into a long, dark brown appendage, somewhat thickcued in the 
middle. Found in the pools on the limestone rocks, at the Whitehead, Juue 25th, 1812. 
Moving with a pretty quick but steady motion, by expanding and collapsing the body, 
which was so extremely transparent, that scarcely any part was visible but the dark brown 
appendage and the marginal tentacula. The marginal tentacula were dilated at their 
base." Anomalous as this creature is represented, I hardly doubt that it is other than 
Sarsia tubiilosa accidentally turned inside out, as I have elsewhere observed. A curious 
Medusa, having a simple umbrella without tentacula at the margin, is figured at Cut 47. 
It is described as " Ocyrhos (?), Peron (Cassiopeia (?), Lam.) cruciata. Hyaline, four arms, 
pale purple, corrugated ; eight darker, fine rays, and numerous dusky obsolete oues." The 
figure seems to represent a naked-eyed species, but it may be some mutilated Pulmograde 
of higher rank. It would be unsafe, without new observations, to admit this form into 
systematic lists. All the other forms mentioned by Templeton (except " Medusa 
scintillans of Macartney," which probably refers to TTiaumantias hemisphcerica) are 
members of higher groups, and some of them, such as his " yEquorea (?) radiata," 
monstrous and mutilated Aurelia. 



BIBLIOGRAPHY. 97 

1837. Cuvier's 'Regne Animal,' Commemorative Edition, illnstratcfl by his pupils. 

Among the plates of this beautiful work are thirteen representing Medusa;. Eight of 
these are devoted to undoubted steganopthahnatous species. The following naked-eyed 
forms, or else doubtful, are copied from the unpublished plates of Pcron : 

Phorcynia ist'wpliora ; Eulhnenes cycloplnjlla ; Equorcd I'lO'pvrea ; Lymnorea tr'uedra (covered- 
eyed?); Favonia hexamena ; Geryonia hexajihyUa {Dianiea proboscidalis?) ; Berenix cariso- 
chroma, B. euchroma (this genus seems to have affinities with WiUsia, but the structure of the 
peduncle and ovaries is not indicated in the drawing) ; Geryonia dinema (possibly 
belonging to a genus of the Sarsiadte, as well as Orythia viridis) ; Orytlda minima (this 
appears to be an immature Cyaiuea) ; Eudora undulosa (steganopthahnatous ?) ; Carybdea 
periphylla (surely not of this genus, and possibly steganopthahnatous). 

Figures are also given of Geryonia bicolor, copied from Eschscholtz ; G. Dubunlii [balearica) 
[possibly an Orytbia], after Quoy and Gaimard; G. tetraphy/la, after Cliaraisso and 
Eysenhardt ; Carybdea marsupialis, aud ^Eqiwrea violacea, after Milne Edwards. 

1837. Lesson. ' Prodrome d'une Monographie des Meduscs.' 
I have never seen this work. Was it ever printed? 

1837. Ehrenberg, in the ' Transactions of the Berlin Academy,' for the year 183."), vol. viii, gives 

two good figures of naked-eyed Medusae, the one " Oceania pileata," and the other 
" Melicertum campamdatum" (really Stomobrachium octocostatum), both from Norway, 
and already noticed in the account of our native species of the genera to which they 
belong. 

In the same paper there is a catalogue of the Medusse of the Red Sea, but all the species 
enumerated are steganopthahnatous. 

1838. J. F. Brandt. " Ausfiihrliche Beschreibtnig der von C. H. INIertens auf seiner Weltumsegelung 

beobaehteten Schirmquallen, nebst allgemeinen Bemerkungeu liber die Schirmquallen 
iiberhaupt," with 34 coloured plates, in the ' Memoirs of the Imperial Academy of St. 
Petersburg,' 6th series, ' Sciences Naturelles,' 2d vol. 

One of the most valuable and beautifully illustrated memoirs upon the Medusse extant. 
The naked-eyed species figured in it are Circe Camtscatica, Conis niitrata [a Turris?), 
JErjuorea rhodokma, Stomobrachium lenticulare, Mesonema macrodactyla, carulescens and 
dubiiim, jrE(jinopsis Laurentii, (if the Pohjenia Alderi of this work prove not to belong to that 
genus to which I have referred it, ^Egitiopsis may prove its proper place,) Geryonia 
hexaphylla, Proboscidactyla flavicirrhata (a genus of WiUsiada;), Hippocrene Bougainvillii, 
and (?) Staurophora Mertensii. The figures are from the drawings of Mertens, and bear 
every mark of being faithful representations. The remaining species are Steijanoplhahtiata, 
and are by far the best figures hitherto published of JSIedusse of that order. 

18-10. The second edition of Lamarck's ' Auimaux sans Vertebres/ edited by Deshaj'es and Milne 
Edwards. 

The Medusaj are contained in the third volume of this edition, and have beeu revised by 
M. F. Dujardin. The additional notes are very good, and serve to make the work a 
useful manual, as they embody the labours of recent writers, especially Eschscholtz and 
Brandt. 

1841. Milne Edwards described and figured with admirable accuracy the Mquorea violacea, in the 16th 
volume of the second series of the ' Annales des Sciences Naturelles.' 

1841. Augustus A. Gould, M.D. 'Report on the Invertebrata of Massachusetts,' Cambridge, 
U. S., 8vo. 

Three species of naked-eyed Medusae are enumerated as inhabiting the shores of the 
United States, viz., " Oceania tubulosa" (i. e. Sarsia tubulosa), " Hippocrene Boiiffainvillii" 
(more probably Boiigainvillea britannica), and "Stomobrachium lenticulare :" the two latter 
identified with Brandt's species of those names. 

13 



98 



BIBLIOGRAPHY. 



1.841. Dr. Davis on " Cyatuea coccinea," {Turris neglecta,) 
Natural History.' 



in the Seventh Volume of the ' Annals of 



1841. E. Forbes, on anew Hippocrene, and some new species of Thaumantias in the 'Annals of Natural 
History.' 

1843. Lesson. ' Histoire Naturelle des Zoophytes ; Acalephes/ a thick 8vo volume, forming part of 
the ' Nouvelles Suites h Buffon.' 

This work is oue of the most useful, and yet one of the most provoking, in its department 
of natural history ; useful, because it brings together, verbatim, everything that has been 
written upon the Medusae in France ; provoking, because every attempt in it at an 
arrangement or digest of the matter so collected serves only to make the obscure more 
obscure, and the crude more crude. It is executed without any judgment, though with 
considerable industry. Of what has been done outside of France it is a most imperfect 
account. Nevertheless, for the present, it is indispensable to the student of the Medusse, 
and includes the fullest list published of species and references. A few plates in which 
some interesting species are figured are appended. I have compiled the following 
Catalogue of Medusse, either naked-eyed, or possibly so, enumerated by Lesson, with the 
■ localities given in his work, and the name of the original observer. 



NAKED-EYED MEDUSA ENUMERATED BY LESSON. 



Figured in works referred to by Lesson. 



f Figured in this Monograph. 



Name in Lesson's Work. 


Locality. 


Observer and Remarks. 


* ? 


Eulimenes sphaeroidalis, Peron. 


South Atlantic. 




Peron. 


* ? 


Eulimenes cyclophylla, Peron. 


South Atlantic. 




Peron and Lesueur. 


? 


Eulimenes heliometra. Lesson. 


Peru. 




Lesson. 


* 


Phorcynia cudouoidea, Peron. 


Australia. 




Peron and Lesueur. 


* 


Phorcynia petasella, Peron. 


lies Furneaux. 




Peron and Lesueur (figure unpublished). 


* 


Phorcynia istiophora, Peron. 


S. Australia. 




Peron and Lesueur (fig. unpub.) 




Phorcynia cruciata, Lin. 


Norway. 




Linnaeus (figure much wanted). 


* 


Pileola pilcata, Q. and G. 


Gibraltar. 




Quoy and Gaimard (Genus?). 


* 


Marsupialis Plauei, Less. 

{Caryhdea marsupialis, Lam.) 


Mediterranean. 




Plancus (well figured by Milne Edwards). 


* 


Marsupialis alata, Beyn. 


Atlantic. 




Reynaud (figured in Lesson's 'CenturieZoo 
logique.') 




Marsupialis flageUata, Lesson. 


New Guinea. 




Lesson (figure wanted). 


* 


Bursarius cytherene, Lesson. 


New Guinea. 




Lesson (very doubtful). 


* 


Mitra Rangii, Lesson. 


(W?) African Seas. 




Rang. 


* 


Eurybia exigua, Eschsch. 


South Sea, between the 


tropics. 


Eschscboltz. 


* 


Cytteis tetrastyla, Esch. 


Atlantic, under the Equator. 


Eschscboltz. 


* 


? Campanella cbaraissouis. Lesson. 

(Medusa campaniilata, E. and C.) 


Pacific. 




Eyseuhardt and Chamisso, insufficiently known 




Campanella Fabricii. 


Greenland and Baffin's 


Bay. 


0. Fabricius (no figure). 




(Medusa campanula, 0. F.) 








* 


Scypbis mucilaginosa, Chamissoa.ndiEysenh. 


Pacific. 




Chamisso (too imperfect). 



BIBLIOGRAPHY. 



99 



Niime in Lesson's Work. 



Locality. 



Observer and Remarks. 



* Turris papua, Lesson. 
•f- Turris borealis, Less. 

{T. digitate.) 
f * Turris neglecta, Less. 

* Circe camtschatica, Brandt. 

* Circe anais, Less. 

* Circe elongata. Less. 

* Tiara papalis, Lesson. 

{Medusa pileata, Forskal.) 

* Tiara Sarsii, Lesson. 

(O. ampullucea, Sars.) 

* Tholus funerarius, Q. and G. 

* Pandea conica, Q. and G. 

* Pandea rotunda, Q. and G. 

* Pandea saltatoria, Sars. 

* Bougainvillca macloviana. Lesson, 
f * Bougainvillea bi-itannica, Forbes, 
f * Bougainvillca octopunctata, Sars. 

* Proboscidactyla flavocirrata, Brandt. 

* Melicertum penicillatum, Esch. 

* Aglaura hemistoma, Peron. 

* Laodicea crucifera, Lesson. 

(Medusa criiciata, Forsk.) 

* Microstoma ambiguus, Lesson. 

* Berenix euchroma, Peron. 

* Berenix tbalassina, Peron. 

* Berenix Cuvieri, Peron. 

* ? Staurophora Mertensii, Brandt. 

* Pegasia dodecagona, Peron. 

* Pegasia cylindrella. 

* Foveolia moUicina, Fors/i. 
Foveolia pilearis, Gmelin. 
Foveolia bunogaster, Peron. 
Foveolia diadema, Peron. 
Foveolia lineolata, Peron. 
Foveolia pulvinata. Lesson. 
Cunina campanulata, Escli. 
Cunina globosa, Esch. 
jEgina citrea, Esch. 
jEgina rosea, Esch. 
^gina grisea, Q. and G. 
^gina semi-rosea, Q. and G. 
^gina capillata, Q. and G. 
^gina nivea. Lesson. 
^gina corona, Lesson. 
Mgina carolinarum, Lesson. 
jEginopsis Laurentii, Brandt. 
^quorea Forskalea, Peron. 
.Slquorea ciliata, Esch. 
j^quorea violacea, MUne Edw. 



Coast of the L of Waigeou. 
Arctic and Boreal Seas. 

Celtic Seas. 
Karatschatka. 
(W ?) Africa. 
(W?) Africa. 
Mediterranean. 

Norway. 

Gibraltar. 

Gibraltar. 

Gibraltar. 

Norway. 

North Pacific and Behring's Straits. 

North Atlantic. 

Norway. 

Kamtschatka. 

California. 

Mediterranean. 

Mediterranean. 

Island of Waigeon. 

Equatorial Atlantic. 

Arnheim's Land. 

Australia ? 

Pacific. 

S. Atlantic. 

Arnheim's Land. 

Mediterranean. 

Atlantic Ocean. 

Mediterranean. 

South Atlantic. 

Mediterranean. 

Indian Seas (?). 

Azores, 

South Seas. 

North Pacific. 

North Pacific. 

Admiralty Isles. 

New Guinea. 

Gibraltar. 

Indian Seas (?). 

Indian Seas (?). 

Pacific. 

Behring's Straits. 

Mediterranean and Atlantic. 

N. W. America. 

Mediterranean. 



Lesson. 

0. Fabricius (Tnrris). 

Davis (Turris). 
Mertens (Circe). 
Rang (Circe). 
Rang ( Circe). 
Forskal (Oceania). 

Sars (Oceania). 

Quoy and Gaimard (Tholus). 

Quoy and Gaimard. 

Quoy and Gaimard. 

Sars (Circe?). 

Mertens (BovgainviUea). 

Forbes (Boiigainvillea) . 

Sars (Lizzia). 

Mertens. 

Eschscholtz. 

Peron, Risso (figure unpublished). 

Forskal. 

Lesson. 

Peron and Lesueur. 

Peron and Lesueur. 

Peron and Lesueur. 

Mertens. 

Pei'ou and Lesueur. 

Peron and Lesueur (figure unpubUshed). 

Forskal. 

Peron. 

Peron. 

Peron. 

Lesson (?). 

Eschscholtz. 

Eschscholtz. 

Eschscholtz. 

Eschscholtz. 

Quoy and Gaimard. 

Quoy and Gaimard. 

Quoy and Gaimard. 

Raynaud. 

Raynaud. 

Lesson. 

Mertens. 

Forskal. 

Eschscholtz. 

Milne Edwards. 



100 



BIBLIOGRAPHY. 



Name in Lesson's Work. 


Locality. 


Observer and Remarks. 


* 


jEquorea globosa, Eseli. 


South Seas, between tropics. 


Eschscholtz. 


* 


iEqiiorea eurodina, Peron. 


Bass's Straits. 


Peron and Lesueur (tig. unpub.) 


* 


.Slquorea cyanea, Peron, 


Arnheim's Land. 


Peron and Lesueur (fig. unpub.) 


* 


J5quorea thalassina, Peron. 


Arnbeim's Land. 


Peron and Lesueur (fig. unpub.) 


* 


yEquorea stauroglypha, Peron. 


Frencb Coasts of British Channel. 


Peron and Lesueur (fig. unpub.) 


* 


.^quorea allantopbora, Peron. 


French Coasts of British Channel. 


Peron and Lesueur (fig. unpub.) 


* 


J<;quorea Risso, Peron. 


Mediterranean. 


Peron and Lesueur, Risso (fig. by Risso). 


* 


.lEquorea amphicurta, Peron. 


De Witt's Land. 


Peron and Lesueur (fig. unpub.) 




jEquorea bunogaster, Peron. 


Van Arnheim's Land. 


Peron. 


* 


jEquorea sphseroidalis, Peron. 


Endracbt's Land. 


Peron and Lesueur (fig. unpub.) 


* 


iEquorea pliospboriphora, Peron. 


Arnheim's Land. 


Peron and Lesueur (fig. unpub.) 


* 


jEquorea rbodoleina, Brandt. 


Conception, in Club. 


Mertens. 


t 


* ^quorea octocostata, Sars. 


Norway (British Seas). 


Sars. 




' jEquorea atlantica, Peron. 


North Atlantic. 


Locfling. 




{Medusa ee^vorea, Locfliag.) 








' jEquorea danica, Peron. 


Scandinavia. 


0. F. Mailer. 




{Medusa (Bquorea, MiJller, Gm.) 








' .bquorea groenlaudica. 


Greenland. 


0. Fabricius. 




{Medusa cequorea, 0. Fabricius.) 






* 


Polyxenia cyanostylis, Eschs. 


N. Atlantic. 


Eschscholtz. 


* 


Polyxenia purpurea, Peron. 


Endracbt's Land. 


Peron and Lesueur. 


* 


Polyxenia pleuronota, Peron. 


Arnheim's Land. 


Peron and Lesueur (fig. unpub.) 


* 


Polyxenia undulosa, Peron. 


Arnbeim's Land. 


Peron and Lesueur (fig. unpub.) 


* 


Polyxenia flavobracbia, Brandt. 


South Sea. 


Mertens. 


* 


Stomobrachiota lenticularis, Brandt. 


Falkland Isles. 


Mertens. 


* 


Mesonema ccelumpensile, Modeer. 


Mediterranean and Atlantic. 


Forskal. 


* 


Mesonema abbreviata, Esch. 


Straits of Sunda. 


Eschscholtz. 


* 


Mesoneraa pileus, Lesson. 


African Seas ? 


Lesson. 


* 


Mesonema macrodactyla, Brandt. 


South Sea. 


Mertens. 


* 


Mesonema ccerulescens, Brandt. 


35° N. lat., 144° Ion., W. 


Mertens. 


* 


Mesonema dubium, Brandt. 


Chili. 


Mertens. 


* 


Oceania pbosphorica, Peton. 


French coasts of the Channel. 


Peron. 




Oceania lineolata, Peron. 


Mediterranean. 


Peron, Risso. 




Oceania flavidula, Peron. 


Mediterranean. 


Peron, Risso. 




Oceania Lesueur, Peron. 


Mediterranean. 


Peron, Risso. 


* 


Oceania dinema, Peron. 


Frencb Coasts of the Channel. 


Peron and Lesueur (fig. unpublished). 




' Oceania bimorpba, MiiUer. 


Baffin's Bay and Norwegian Seas. 


0. Fabricius. 


* 


1 Oceania tetranema, Peron. 


Holland (Slabber). 


Slabber. 


* 


1 Oceania sangninolenta, Peron. 


Holland (Slabber). 


Slabber {Turris coccinea). 


* ? Oceania danica, Peron. 


Norway. 


Thanmantias hemisphcerica, of Midler. 




1 Oceania paradoxa, Peron. 


Mediterranean. 


Peron. 


* 


? Oceania microscopica, Peron. 


Holland (Slabber). 


Slabber. 




1 Oceania heteronema, Peron. 


Havre. 


Peron. 


* 


? Melicerta perla. Slabber. 


Holland. 


Slabber, perhaps a young Pelayiu. 




Melicerta pleurotoma, Peron. 


De Witt's Land. 


Peron. 




Melicerta fasciculata, Peron. 


Mediterranean. 


Peron. 


* 


Melicerta morcbella, Lesson. 


1 


Lesson. 


t 


* Sapbeuia dinema, Peron. 


French and British Coasts. 


Perou and Lesueur. 


:!: 


Sapbenia bitentaculata, Q. and G. 


Gibraltar. 


Quoy and Gaimard {Geryonia or Tima). 


* 


Sarbeuia balearica, Q, and G. 


Mediterranean. 


Quoy and Gaimard. 



BIBLIOGRAPHY. 



101 



Name in Lesson's Work. 



Locality. 



Observer and Remarks. 



* Dianea endractensis, Q. and G. 
Dianea viridula, Peron, 
Dianea gibbosa, Peron. 

* Orythia viridis, Peron. 

* Orythia minima, Cuv. 

* Geryonia tetraphylla, Eijs. and Cham. 

* Geryonia bicolor, Eseh. 

* Geryonia rosacea, Esch. 

* Geryonia exigua, Q. and O. 

* Liriopa proboscidalis, Forskal. 

* Liriopa eerasiformis. Less. 

* Xantbia agaricina. Lesson. 
t * Sarsia tubulosa, Sars. 

* Tima ilavilabris, Esch. 

* Tbaumantias cymbaloidea, Slabber. 
■\ * Tbaumantias bemisphserica, Gron. 

* Tbaumantias lucida, Mac. 

* Tbaumantias plana, Sars. 

* Tbaumantias multicirrata, Sars. 
•j- * Tbaumantias pileata, Forbes. 

f * Tbaumantias Thompson!, Forbes, 
f * Tbaumantias punctata, Forbes, 
■f * Tbaumantias sarnica, Forbes. 
? * Linuche unguiculata. Swart.:. 
Usous roseus. Lesson. 

* Lymnorea triedra, Peron. 

* Favonia octonema, Peron. 

* FaTonia hexanema, Peron. 



S. W. Coast of Australia. 

French Coasts of Channel. 

Mediterranean. 

Endracbt's Land. 

Belgium. 

Indian Ocean. 

Brazil, of Cape Frio. 

S. Sea, between the Tropics. 

Gibraltar. 

Mediterranean. 

Gibraltar. 

? 

Norway (Britain, N. America). 

Azores. 

Holland. 

Norway (British Seas). 

Britain. 

Norway. 

Norway. 

Britain. 

Britain. 

Britain. 

Britain. 

Jamaica. 

? 

Bass's Straits. 
Australia. 
Equatorial Atlantic. 



Quoy and Gaimard. 

Peron. 

Peron, Risso. 

Peron and Lesueur. 

Baster. 

Eysenhardt and Chamisso. 

Escbscboltz. 

Escbscboltz. 

Qaoy and Gaimard {Geryonia). 

Forskal (Diantsa). 

Quoy and Gaimard (a Tima ?) . 

Lesson. 

Sars. 

Escbscboltz. 

Slabber (only T. hemisjihwrica). 

Miiller, Gronovius, &c. 

Macartney, (var. of last). 

Sars. 

Sars. 

E. Forbes. 

E. Forbes. 

E. Forbes. 

E. Forbes. 

Swartz (perhaps the fig. of a higher Medusa). 

Lesson. 

Peron and Lesueur. 

Peron and Lesueur. 

Peron and Lesueur. 



1844. Dr. J. G. F. Will, of Erlangen. ' Horse Tergestinse/ 4to, Leipsig. 

Some of the best researches on the Acalephse, as yet given to the public, miuute, accurate, 
and philosophical. The naked-eyed species described and figured are Pohjxenia leucostyla ; 
Cytceis polystyla ; Cytoeis (?) ; Geryonia planata ; Geryonia pellucida, [a Geryonopsis,) and 
Thaumantias leucostyla ; all observed in the Adriatic. 

1846. H. Sars. ' Fauna littoralis Norwegise.' 1st pt. in folio, with 10 plates. 

The first memoir in this valuable and beautifully illustrated work is devoted to the Medusa- 
producing polypes (Syncoryna, Podocoryna, Perigonomus,) and to the history of the gemma- 
tion of Cytais octopunctata, the Lizzia octopunctata of this Monograph. 

1846. E. Forbes. " On the Pulmograde Medusaj of the British Seas." 

Communicated to the British Association at Southampton, and including the outline of the 
present treatise. It was printed in the Proceedings of the Association, and ' Annals of 
Natural History ;' also translated in the ' Annales des Sciences NatureUes.' 



INDEX OF GENERA AND SPECIES 

DESCRIBED IN THIS MONOGRAPH, 



WITH THEIR SYNONYMS. 



*^* The names of spurious species and synonyms are in Roman letters. 





PAGE 




PAGE 


.iEquorea octocostata 


. 30 


Euphysa aurata 


■ . ,"KL 


Aurelia aurita . 


. . 75 


Geryonia appendiculata . 


. 30 


campanula 


. 76 


dinema 


. . 25 


cruciata 


. . 76 


hemisphffirica 


. 49 


gramilata . 


. 76 


octona . . . . 


. . 27 


lineolata 


. . 75 


Geryonopsis delicatula 


. 39 


purpurata . 


. 75 


Hippocrene Britannica 


. . 62 


radiolata 


. . 76 


octopunctata 


. 64 


rosea 


. 76 


Lizzia blond'ma . . 


. . 67 


surirea 


. . 76 


octopunctata 


. 64 


Biblis aquitanise 


. 76 


Medusa aurita .... 


. . 75 


Bougainvillea Britannica 


. . 62 


campanula 


. 76 


nigritella . 


. 63 


capillata 


. . 77 


octopunctata 


. . 64 


cruciata 


. 76 


Cassiopea lunulata 


. 77 


cymbaloidea . 


. . 49 


Borlasea 


. . 77 


digitale 


. 21 


rhizostomoidea 


. 77 


duodecilia 


. . 62 


anglica 


. . 77 


fusca 


. 77 


Campauella dinema 


. 25 


hemisphserica 


. . 49 


Chrysaora hysosceUa . 


. . 77 


hysosceUa . 


. 77 


Circe rosea .... 


. 34 


lucida .... 


. . 49 


Cyanea capillata 


. . 77 


lunulata 


. 77 


Lamarckii 


. 78 


ocilia .... 


. . 68 


coccinea 


. . 24 


purpurata . 


. 75 


Cytaeis octopunctata 


. 64 


tuberculata . 


. . 77 


Diansea Bairdii 


. . 37 


Melicerta digitale . 


. 20 


dinema 


. 25 


INIelicertum campanulatum . 


. . 30 


digitale 


. . 21 


Modeeria formosa . 


. 70 


Eirene digitale 


. 21 


Oceania episcopalis 


. . 27 



104 



INDEX. 





PAGE 




Oceania globulosa .... 


29 


Thaumantias convexa 


hemisplijerica 


49 


cymbaloidea 


octocostata 


30 


globosa 


octona ..... 


27 


gibbosa 


sanguinolenta 


23 


hemispharica 


tetranema .... 


23 


inconspicua 


tubulosa .... 


55 


lineata . 


turrita ..... 


28 


lucida 


Pelagia cyanella .... 


76 


lucifera 


Polyxenia Alderi .... 


32 


maculata . 


Rhizostoma jmlmo .... 


77 


melanops 


Aldrovandi .... 


77 


Milleri 


Cuvieri .... 


77 


Octona 


Saphenia dinema .... 


25 


pileata 


Sa7-sia gemmifera .... 


57 


pilosella 


pulchella .... 


57 


punctata 


prolifera .... 


59 


quadrata 


tubulosa .... 


55 


sarnica 


Slabberia halterata 


53 


Ttiompsoni 


SteenMrupia fiaveola .... 


74 


Tima Bairdii 


rubra .... 


73 


Turris digitalis . 


Stomobrachium octocostatum 


30 


neglecta 


Thaumantias ceronautica . 


44 


Wilhia stellata . 



PAGE 

. 47 

. 49 

. 46 

. 47 

. 49 

. 52 

. 48 

. 49 

. 52 

. 45 

. 45 

. 30 

. 44 

. 47 

. 42 

. 53 

. 43 

. 48 

. 49 

. 37 

. 21 

. 23 

. 19 



THE END. 



C. AND J. ADLARD, PRINTERS, BARTHOLOMEW CLOSE. 



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