MEDUSAE OF THE WORLD

VOLUME I

THE HYDROMEDUSAE

BY

ALFRED GOLDSBOROUGH MAYER

WASHINGTON, D. C.

PUBLISHED BY THE CARNEGIE INSTITUTION OF WASHINGTON

1910

CARNEGIE INSTITUTION OF WASHINGTON
PUBLICATION No. 109, VOL. I

Copies of this Book
were first issuec*

AUG9 1910

CONTENTS.

VOLUME I.

Introduction i~4

Synopsis of Genera of the Hydromedusas 5-16

Medusae Milleporinae 16

Anthomedusae 17-196

Genera of the Anthomedusae:

Pachycordyle 21 Zancleopsis 91 Podocoryne 135

Amaithasa 22 Pteronema 92 Turritopsis 143

Pennaria 23 Eleutheria 93 Oceania 146

Trichorhiza 28 Mnestra 96 Stylactis 149

Steenstrupia 29 Ctenaria 98 Thamnostylus 151

Hybocodon 37 Cladonema 98 Thamnitis 152

Microcampana 44 Dendronema 102 Lymnorea 153

Dicodonium 44 Protiara 105 Bougainvillia 155

Sarsia 47 Heterotiara 107 Nemopsis 172

Stnuridiosarsia 64 Stomotoca 108 Rathkea '75

llydrichthys 66 Dissonema 115 Chiarella 182

Eucodonium 68 Pandea 116 Bythotiara ..185

Ectopltura 68 Turns I2O Sibogita 186, vol. 2 4QI

Corynitis 71 Conis 130 Niobia 187

Slabberia 73 Calycopsis 130, vol. 2 491 Proboscidactyla. . . . . . 188

Margelopsis 80 Cytsis ... , : . .-...- ... 132 Willsia . . 193

Zanclea 85

Leptomedusae '..'.' .196-230

Genera of the Leptomedusae:

Thaumantias 198. Staurodiscus 213 Dichotomia m

Laodicea 201 1'tvchogena 214 Dipleurosoma 224

Melicertum 207 I'olyorchis 218 Toxorchis 228

Melicertissa 209 Spirocodon 219 Netocertoides. . . .229

Orchistoma 21 r Cannota 221 Monobrachium. . --230

Timoides . . ..212 Cuvieria 221

MEDUSAE OF THE WORLD.

THE HYDROMEDUSAE.

INTRODUCTION.

This work was commenced in 1892 at the suggestion of Dr. Alexander Agassiz
while I was a student in his marine laboratory at Newport. Dr. Agassiz's plan
was that we produce conjointly a work upon the Medusae, Siphonophora, and
Ctenophorae of the Atlantic coast of North America. In pursuance of this plan,
Dr. Agassiz sent me upon expeditions to Halifax, Nova Scotia; Eastport, Maine;
Charleston, South Carolina; and Tortugas, Florida. It was also my privilege to
accompany him as his assistant upon his expeditions to the Bahamas, and to the
coral regions of the tropical Pacific. During these years the majority of our
American species were captured and drawings of them made from life.

The description of all of the then known American Atlantic forms was com-
pleted by me in 1900, but unfortunately the pressure of other and more important
work prevented the revision of the manuscript by Dr. Agassiz, and thus it remained
in the Museum of Comparative Zoology at Harvard University until 1904, when
Dr. Agassiz generously returned it to me with permission to publish it in any
manner whatsoever. Thus the original plan was reluctantly abandoned.

During the four years that elapsed while the manuscript lay unstudied at
Harvard, new forms had been discovered along our coast; and Nutting and Hargitt
had published their papers upon the hydroids and medusae of the Woods Hole
region.

It was necessary to thoroughly revise the manuscript, and in order to render it
of greater service, I have attempted to extend the original work to include descrip-
tions of all known forms of medusae from all parts of the world. This extension
was made possible through the generous establishment by the Carnegie Institution
of Washington of a Marine Research Laboratory at Tortugas, Florida. Many
forms were collected upon a cruise of the Carnegie Institution yacht Physalia
from Boothbay Harbor, Maine, to Tortugas, Florida; and new or interesting
medusae have been obtained each season upon excursions over the tropical Gulf
Stream, and among the Bahamas. My official position in connection with the
laboratory has afforded me every possible facility in time and opportunity tor tilt-
prosecution of these studies, and I can not too kindly express my sense of gratitude
to the executive officers of the Carnegie Institution tor their generous support.

To Geheimrath Prof. Dr. Anton Dohrn, and to his able corps of associates at
the Stazione Zoologica, I am indebted for numerous kindnesses shown to me during
my visit to the laboratory at Naples from November to February, 1907 and 1908.

I have also enjoyed full privileges of study in the libraries of the Museum of
Comparative Zoology at Harvard University; in the Boston Society of Natural

1

2 MEDUSvE OF THE WORLD.

History; the American Museum of Natural History; Columbia University, the
National Museum at Washington, and the Museum of the Brooklyn Institute of Arts
and Sciences. Through these facilities I have been enabled to review nearly all
of the published works upon medusae, but the review of literature can not pre-
tend to completeness for 1907 and 1908, although all papers of those years which
the author could discover are recorded.

Moreover, Profs. William K. Brooks and Louis Murbach have been so kind
as to lend some of their original drawings, which are reproduced in this work, and
the following gentlemen have generously granted permission for the reproduction
of figures from their published works, thus enabling us to present text-figures of
many forms which would otherwise have been represented merely by descriptions:
Dr. Alexander Agassiz, Director of the Museum of Comparative Zoology at Harvard
University; Dr. Henry B. Bigelow, of Harvard; Dr. R. P. Bigelow, of the Massa-
chusetts Institute of Technology; Prof. Edward T. Browne, of the University of
London; Prof. Dr. Carl Chun, of Leipzig; Prof. Dr. S. Goto of Tokyo;
Geheimrath Prof. Dr. Ernst Haeckel, of Jena; Prof. C. W. Hargitt, of Syracuse
University; Prof. Dr. Cl. Hartlaub, of Helgoland; Prof. W. C. M'Intosh, of Aber-
deen University; Prof. Dr. Otto Maas, of Miinchen; Prof. C. C. Nutting, of Iowa;
Prof. Henry F. Perkins, of Vermont; and Prof. Dr. Ernst Vanhoffen, of Kiel.

I have always felt that each working naturalist owes it as a duty to science to
produce some general systematic work, and this has been an actuating motive
in the production of this book. But chiefly have I been moved to the task through
respect for the wishes of my generous friend and master in science, Alexander
Agassiz. Nor can one remain insensible to the rare grace of form and delicate
beauty of color of these creatures of the sea, associated as their study is with mem-
orials of the labors of a host of distinguished naturalists. Dry though these pages
must be to the reader, to the writer they are replete with memories of the ocean
in many moods, of the palm-edged lagoons of coral islands sparkling in the tropic
sun, of the cold, gray waters of the northern sea bestrewn with floating ice, of days
of withering calm in the heat of the torrid zone, and of adventure in the hurricane ;
all centering around the absorbing study of the medusa?. Love, not logic, impels
the naturalist to his work.

This book attempts to present a new classification of the medusae. With every
respect for Haeckel's great work, it has appeared to me that its subdivisions are
often too precise to be convenient, and too artificial to accord with nature. More-
over, many of Haeckel's genera are founded upon intergrading characters, and are
thus imperfectly separated. The young of many medusae appear in one genus,
and the adults in another. The aim of Haeckel's system is to emphasize distinctions,
whereas my aim is to indicate relationships. I therefore attempt to separate genera
upon positive, not upon relative, distinctions. For example, if we define one genus
as having a narrow manubrium (Margelis), and another as having a wide manu-
brium (Boiigamvillta), we must either institute a third genus for newly-discovered
medusae with manubria of moderate width or place them doubtfully in one or the
other of the genera of the extreme members of the series.

On the other hand, if we define one genus as having eight tentacles, and
another as having nine or more tentacles, there can be no confusion between them,
for the difference, although slight, is positive and numerical, not qualitative and
intergrading.

I have not described hydroids which do not produce free-swimming medusae,
although I grant it is wholly illogical to admit Podocoryne into a system which
excludes Hydractmia, or to include only those species of Stylactis which produce

INTRODUCTION.

medusae and to exclude those which produce sessile gonophores. Nevertheless
this should be clearly understood and must be accepted as an artificial limitation
of the work.

I have thus attempted to describe only such hydroids as are known to produce
medusae, and have endeavored to bring the systematic arrangement of the medusa
more nearly into accord with that of the hydroids. A strictly natural system includ-
ing both hydroids and medusae can not be constructed, for many of the hydroids
remain undetermined. Moreover, dissimilar hydroids (Svm-tjrv>if, StanriJinim
may give rise to similar medusae (Sarsia); or the reverse may be the case, as in the
medusae of Bongamvillia and Nemopsis, or that of the two sorts of medusa- (Sarsia
and Cladoncrna] which may arise from hydroids of Stauridia.

These and many other cases of a similar nature interpose a barrier to our
attempts to invent a natural system which includes all hydroids and medusa" within
its embrace. At present, I believe, we must content ourselves with a compromise
between a natural and an artificial arrangement, confiding in the belief that as
more and more of the hydroids are discovered it will become correspondingly
more possible to arrange the medusae in a natural system. After consultation with
Prof. C. C. Nutting we have mutually decided that the promulgation of such ;i
system is at present inadvisable. Such a system has, indeed, been proposed by
von Lendenfeld, 1884 (Zool. Anzeiger, Bd. 7), but has gained no acceptance.

Much confusion has been introduced through the habit, in vogue among
marine expeditions, of sending all of the medusa" to one specialist and the hydroids
to another. Thus the sessile and the reproductive stages of the same animals are
worked upon independently from different view-points by different men.

I am inclined to regard the Trachymedusae and Narcomedusa" as being
transformed actinutae, for they commonly develop through an actinula larva in
which the bell grows out as a collar-like, or intertentacular lappeted expansion
from the sides of the body after the tentacles have appeared, and the tentacles of
the actinula become those of the medusa. The medusa of the Anthomedusas and
Leptomedusae is formed upon a different plan, for the tentacles grow outward from
the bell-margin after the bell has developed. I believe, therefore, that the bell of
the Trachymedusae and Narcomedusae is not homologous with that of the Antho-
medusa and Leptomedusae. It is evident that the entodermal otoliths of the
Trachymedusa" and Narcomedusae are not homologous with the ectodermal otoliths
of Leptomedusae. Budding and alternation of generations occur in both classes
of veiled medusae.

I believe that the medusa-shape has been acquired independently in the
Trachyhna and Leptolma forms of veiled medusae.

The colored plates contained in this volume consist of drawings, from life, of
medusae of the Atlantic coast of the United States. The text-figures, on the other
hand, are chiefly outline tracings from the illustrations of many authors; and are
presented in order to spare the reader the trouble of consulting numerous scattered
works of reference. These outline copies of previously-published drawings of
medusae were carefully traced from the originals by Mr. Carl Kellner, artist of the
Tortugas Marine Laboratory of the Carnegie Institution.

Other outline figures are from life, and the majority of these were drawn by
the author while studying at Mousehole, Cornwall, England, and at the Naples
Zoological Station during the autumn and winter of 1907 and 1908.

This book aims to be something more than an old-fashioned systematic treatise,
for it attempts to record, if not to review, all works upon the embryology, cytology,
oecology, physiology, etc., of all forms coming within the scope of the text.

MEDUSAE OF THE WORLD.

Many of the medusae are modified more or less profoundly by environmental
conditions, and this gives rise to numerous local races, the determination of the
relationships between which is all but impossible in the present state of our knowl-
edge. Experimental work in this field is in its infancy, yet the few observations of
Hallez on Bongainvillia and of Browne in his experiments in maintaining various
hydroids under different conditions in aquaria suggest that the results of more
extended studies will be of great benefit to the systematist in determining the
natural limitations of species. A striking example of the profound effect of environ-
mental changes is seen in the experiments of Goldfarb, 1906, upon Eudendrium,
wherein he showed that after the regenerative process had entirely ceased in the
dark, it could be recalled into activity by an exposure of only five seconds to the
daylight.

At present the natural classification of the species of such genera as Obclia,
Phialidnnn, Etttnna, Bougamvillia, etc., is impossible, and the Linnean system is
inadequate to the task of expressing their actual relationships. Indeed, with the
exception of the sponges and corals, there is no phylum of the animal kingdom more
difficult to classify than the medusae.

In a work of this magnitude there must needs be both errors and omissions,
and for these I can but present my apologies in advance of their discovery, trusting
that all such will soon be discovered and announced, and that science may be more
advanced than hindered through the publication of this book. I trust that none
of my esteemed contemporaries will feel aggrieved at such criticisms of their labors
as may appear in the following pages, for should anyone find cause for such offense,
he may console himself in the fact that I am obliged to confess to having found
more to criticize and amend in my own previously-published papers than in those
of any other living naturalist.

ANTHOMEDUS^:.

HYDROMEDUS/E OR VEILED MEDUS/E.

Medusae with a velum, or diaphragm, which partially closes the marginal opening of
the bell. When ripe the sexual products are found in the ectoderm.

With a double marginal nerve-ring, one above and one below the velum.

Without gastric filaments. Development either direct from actinula larvae or through
alternation of generations from hydroids.

Order 1. ANTHOMEDUS/E Haeckel, 1879.

Hydromedusae with gonads in the ectoderm of the manubrium. Without otocysts.
The hydroids are of the Tubularian order.

Family No. i, CODONID.&.

Anthomedusae with ring-like gonad encircling the manubrium. Four to six simple,
unbranched radial-canals. Simple, unbranched tentacles.

Subfamily No. i, SARSIANJE.
Some or all of the tentacles arise singly, not in clusters, from bell-margin.

PACHYCORDYLE, Weismann, i8%$ = Parvanernus ( ?), Mayer, I94-

Degenerate medusae without tentacles, radial or circular canals. Hydroid: Pachy-
cordyle Weismann.

AMALTH^EA, Schmidt, 1854.

Four rudimentary tentacles, four radial-canals, and a ring-canal. Hydroid: Corymorpha.

PENNARIA, Oken, 1815; Goldfuss, i%2O = Globiceps, Haeckel, 1879.
Similar to Amalthaa, but the hydroid is Pennana.

TRICHORHIZA, Russell, 1906.

Medusa resembles Pennaria, but the hydroid is Tnchorhiza.

STEENSTRUPIA, Forbes, iS^.6 = Eup/i\s(i, Forbes, i$4.% = Eupliysora, Maas, 1905.

Four radially situated tentacles, one of which is long, the other three short. Bell radially

symmetrical. Hydroid: Corymorpha.
HYBOCODON, L. Agassiz, \%()2 = Hybocodon J t-Amphicodon, Haeckel, 1879.

One or more well-developed tentacles arise from base of one of the four radial-canals.
With rudimentary tentacles at the bases of the three other radial-canals. Bell asym-
metrical. Hydroid: H ybocodon Agassiz.

MICROCAMPANA, Fewkes, 1889.

With one long and five short tentacles, arising at ends of six radial-canals.

DlcoDONlUM = Dicodonium + Dincmti, Haeckel, 1879.

Two well-developed and two rudimentary tentacles. No meridional lines of nettle-cells

over exumbrella.
SARSIA, Lesson, i^^.^ = CoJontutn + Sarsia + S'vnJii'tyon, Haeckel, 1879.

Four equally developed tentacles with abaxial, ectodermal ocelli. No meridional lines

of nettle-cells over the exumbrella. Hydroid: Syncoryne.
StauriJiosarsia new subgenus. Medusa similar to Sarsia, but the hydroid is Stuundia.

HYDRICHTHYS, Fewkes, 1888.

Medusa resembles Sarsia, but there are no ectodermal ocelli upon the tentacle-bulbs.
Hydroid: H vJrichthys, Fewkes; commensal or parasitic upon fish.

EUCODONIUM, Hartlaub, 1907.

Medusa resembles Sarsia, but stomach is mounted upon a gelatinous peduncle. The
four tentacles terminate each in a knob-shaped extremity.

ECTOPI.EURA, L. Agassiz, 1862 (sens. ampl.).

With two or four tentacles. Eight meridional lines of nettle-cells over the exumbrella.
Hydroid: Ectopleura.

6 MEDUS.E OF THE WORLD.

CORYNITIS, McCrady, 1857.

Four knobbed tentacles. Manubrium cruciform in cross-section. Ocelli upon tentacle-
bulbs. Hydroid: Syncoryne (Hargitt).

SLABBERIA, Forbes, lB^6 = Slabberia + Dif>urena + BathycoJon, Haeckel, 1879.

Four knobbed tentacles. Manubrium tubular and encircled by two or more ring-like
gonads. Hydroid: Syncoryne.

Subfamily No. 2, MARGELOPSIN^E.

Four radially placed clusters ot marginal tentacles. No oral tentacles. A ring-like
gonad encircles the stomach. Four simple, unbranched radial-canals. Hydroids are pelagic
Tubularians, and the medusae arise by budding from their sides.

MARGELOPSIS, Hartlaub, 1897.

Characters of the medusa are those of the subfamily. Hydroid: Margelopsis, in which
the tentacles are disposed in definite circlets.

PELAGOHYDRA, Dendy, 1903.

Medusa similar to Margelopsis. Hydroid: PelagohyJra. Its tentacles arise irregularly
from sides of the hydranth, and are not disposed in definite circlets.

Family No. 2, CLADONEMID^E.

Tentacles branch dichotomously or complexly, or give rise to a linear series of nematocyst-
bearing filaments along their abaxial sides. Gonads ring-like, or segregated upon the inter-
radial and adradial sides of the manubrium.

Subfamily No. i, PTERONEMINJE.

Manubrium without oral tentacles.

ZANCLEA, Gegenbaur, l8^6 = Zanclea + Gernmaria, Haeckel, 1879.

With two or four tentacles, each of which gives rise to an abaxial row of nematocyst-
bearing branches. With meridional rows of nettle-cells upon the exumbrella.
No ocelli upon the tentacle-bulbs. No brood-pouch above the stomach. Hydroid:
Gemmaria McCrady; Allman.

ZANCLEOPSIS, Hartlaub, 1907.

Similar to Znticlea, but without meridional lines of nettle-cells over the exumbrella. With
ectodermal ocelli upon the outer sides of the tentacle-bulbs.

PTERONEMA, Haeckel, 1879.

Similar to Zanclea, but with a brood-sac above the stomach. Four tentacles.

ELEUTHERIA, Ouatrefages, 1842.

Four to six bifurcated tentacles, and an equal number of simple unbranched radial-
canals. There is an ectodermal brood-sac above, but not connected with the stomach.
The medusa is hermaphroditic and the germ-cells develop in the brood-sac. Hydroid:
Clavatella Hincks.

MNESTRA, Krohn, 1853; Giinther, 1903.

Degenerate medusa parasitic upon Phylhrrhoe. Four to no tentacles, each with an
abaxial line of nettle-warts. Four perradial meridional lines of nettle-cells over
the exumbrella. Four radial-canals. Throat is blocked by a cavernated mass of
entoderm.

Subfamily No. 2, DENDRONEMIN^).

Manubrium with oral tentacles.

CTENARIA, Haeckel, 1879.

Two marginal tentacles with abaxial filaments. Four bifurcated radial-canals. Simple
oral tentacles. Brood-sac above the stomach.

ANTHOMEDUS.E. 7

CLADONEMA, Dujardin, 1843.

Four or five bifurcated or eight to ten simple radial-canals. Branched marginal tentacles.
Simple oral tentacles. No brood-sac above the stomach. Hydroid: Statin Jin
Dujardin.

DENDRONEMA, Haeckel, 1879.

Similar to ClaJonema, but with branched oral tentacles and with brood-sac above stomach.

Family No. 3, OCEANID.S, sensu Vanhoffen.

Anthomedusae in which the gonads are segregated and developed upon the interradial
or adradial sides of the manubrium. With unbranched marginal tentacles. Mouth with
four lips.

Subfamily No. i, TIARIN.E.

Unbranched radial-canals. Marginal tentacles separate; not grouped into clusters. No
oral tentacles. Tentacles hollow. When present the ectodermal ocelli are upon the abaxial
sides of the tentacle-bulbs. With the exception of Calycopsis all of the genera have four
radial-canals.

PROTIARA, Haeckel, i%jq = Halitiara, Fewkes, 1882.

Four radially placed, well-developed tentacles. Four interradial gonads with smooth
outer surfaces. With or without marginal cirri. External surfaces of gonads smooth.
Four cruciform, simple lips. No ocelli on the velar sides of the tentacles.

HETEROTIARA, Maas, 1905.

Eight or more tentacles. The ring-canal gives rise to blindly-ending centripetal vessels.

STOMOTOCA, L. Agassiz, \%()2 = Amf>hinema-\-Stomotoca + Codonorchis, Haeckel, 1879.

Two well-developed and many rudimentary tentacles. External surfaces of the adradial
gonads are thrown into transverse folds. Hydroid: Pengonimus.

DISSONEMA, Haeckel, 1879.

Similar to Stomotoca, but the gonads finally migrate outward along the four radial-canals.

PANDEA, Lesson, 1843.

Four or more tentacles. Gonads four interradial, folded ridges on the sides of the
stomach but these gonads are not completely separated in the four principal radii.
Hydroid: Dcndroclava ( ? ?).

TURRIS, Lesson, 184.3 = Tiara + Turris + Catablema, Haeckel, 1879.

Four or more tentacles. Four interradial horseshoe-shaped gonads on the stomach
wall. These are composed of more or less fused ridges or network-like swelling.
They are completely separated in the four principal radii. Hydroid: Clirculu Wright.

CONIS, Brandt, 1834; Haeckel, 1879.

Similar to Pandea, but the tentacle-bulbs give rise to abaxially-placed clubs which bear
ocelli.

CALYCOPSIS, Fewkes, 1882.

Sixteen simple, separate radial-canals. Eight transversely folded, adradial gonads.
Ring-canal simple.

Subfamily No. 2, MARGELINjE.

With four unbranched radial-canals. With oral tentacles, or nematocyst-knobs, upon
the lips. Tentacles solid. When present the ectodermal ocelli are upon the inner (velar)
sides of the tentacles.

CYTVEIS, Eschscholtz, iS2() = Cytxis + Cubogastcr, Haeckel, 1879.

Four simple marginal tentacles. With simple, unbranched, oral tentacles.

PODOCORYNE, Sars, \?>J t b = Dysmorf>hosa-\-Cytteandra, Haeckel, 1879.

Eight or more simple marginal tentacles. With simple, unbranched, oral tentacles.
Hydroid: Podocoryne. When present the peduncle above the stomach is solid and
gelatinous.

MEDUSAE OF THE WORLD.

TURRITOPSIS, McCrady, 1856.

Eight or more simple marginal tentacles. The entodermal walls of the radial-canals
above the stomach are composed ot vacuolated cells forming a peduncle-like base
for the stomach. The mouth is studded with a row of nematocyst-bearing knobs.
With ectodermal ocelli on the velar sides of the tentacles near their bases. Hydroid:
DenJroclava (Brooks).

OCEANIA sensu Kolliker, 1853; Gegenbaur (in part) 1856.

Medusa similar to Turritopsis, but with solid gelatinous, non-vacuolated peduncle above
the stomach. Hydroid: C lava-like.

STYLACTIS, Allman, 1864.

Degenerate medusae, with four to eight rudimentary marginal tentacles and no oral
tentacles. Hydroid: Stylactis.

THAMNOSTYLUS, Haeckel, 1879.

With two simple, marginal tentacles, and with branched oral tentacles.

THAMNITIS, Haeckel, 1879.

Four radially placed, simple, marginal tentacles, and branched oral tentacles.

LYMNOREA, Peron and Lesueur, iSog Limnorea + Thamnostoma, Haeckel, 1879.
Eight or more simple, marginal tentacles. Branched oral tentacles.

BOUGAINVILLIA, Lesson, \$4.T > = Margelis + Lizusa + Hif>pocrene, Haeckel, 1879.

With branched oral tentacles. The marginal tentacles are grouped in four radial clusters.
All of the tentacles are filiform. Hydroid: Bougainvillia.

NEMOPSIS, L. Agassiz, 1849.

Similar to Bougainvillia, but each cluster of marginal tentacles consists of a median
pair of clavate tentacles flanked by filiform tentacles. Hydroid: Bougainvillia.

RATHKEA, Brandt, lS^y = Lizzia + Lizella + Rathkea + Margellium, Haeckel, 1879.

With eight clusters of marginal tentacles. Simple or branched oral tentacles. Ring-
canal simple.

CHIARELLA, Maas, 1897.

Sixteen (eight double) clusters of marginal tentacles. The ring-canal gives rise to centrip-
etal vessels. Branched oral tentacles.

Subfamily No. 3, DENDROSTAURINjE.

With branched radial-canals. No oral tentacles. Marginal tentacles arise singly, and
are not grouped into clusters. Tentacles hollow. No cirri or marginal clubs.

BYTHOTIARA, Giinther, 1903.

Four bifurcated radial-canals and a ring-canal. Four interradial gonads.

SIBOGITA, Maas, 1904.

Four principal radial-canals, which branch complexly. Four interradial gonads. Ring-
canal present.

NIOBIA, Mayer, 1900.

Four principal radial-canals, two of which bifurcate so that six canals reach the circular
vessel. Four interradial gonads. The marginal tentacles develop into medusae,
and are cast off.

PROBOSCIDACTYLA, Brandt, \%Tfi=Dyscannota + Dicranocanna+ Willeta

-\-Proboscidactyla, Haeckel, 1879.

The four primary radial-canals give rise to simple or branched side branches. No ring-
canal. With intertentacular lines of nematocysts upon the exumbrella above the
bell-margin. Gonads on the adradial sides of the stomach extending outward
along the sides of the four main radial-canals.

LEPTOMEDUS.E.

9

WILLSIA, Forbes, 1846.

Similar to ProbosciJactyla, but with six or more primary radial-canals. The hydroid
belongs to the genus Lar Gosse.

Order 2. LEPTOMEDUSjE Haeckel, 1886.

Hydromedusae with gonads upon the radial-canals. When present the otoliths are of
ectodermal origin. The medusae arise through alternation of generations from Campanu-
larian hydroids.

Family No. i, THAUMANTIAD^).

Leptomedusae without lithocysts.

Subfamily No. i, MELICERTIN^.

With simple, unbranched radial-canals and an equal number of lips, without oral ten-
tacles. Cirri or marginal clubs may or may not be present.

THAUMANTIAS, Eschscholtz, \%29 = Tetrtinfmti + T/iiiii>tiiinti(is, Haeckel, 1879.

Four or more tentacles. Four radial-canals. No marginal clubs or cirri. Hydroid:
T haumantias.

LAODICEA, Lesson, i$4$ = Octonema + Laodice, Haeckel, 1879.

Four or more tentacles. Four radial-canals, with marginal clubs or cirri. Hydroid:
Cuspidella.

MELICERTUM, Oken, 1815; sensu L. Agassiz, i%ta=Melicertella+Melicertum, Haeckel, 1879.
With eight or more tentacles. Eight radial-canals. No marginal clubs or cirri. Hydroid:
Melicerturn Agassiz.

M.ELlCER.TlSSA = Mclicertissa + MflicfrtiJnitn, Haeckel, 1879.

Similar to Melicertum, but with marginal clubs or cirri between the tentacles.

ORCHISTOMA, Haeckel, 1879.

More than eight radial-canals. With or without marginal clubs or cirri.

TIMOIDES, H. B. Bigelow, 1904.

Four radial-canals. The ring-canal gives rise to blindly-ending centripetal diverticula.
Numerous tentacles and cirri. Four gonads on the four radial-canals. Stomach
mounted upon a peduncle.

Subfamily No. 2, POLYORCHIN.E.

The radial-canals give rise to branches which end blinJlv and do not connect with the
marginal ring-canal.

STAURODISCUS, Haeckel, 1879.

With four radial-canals, each of which gives rise to two side branches, which end blindly.

PTYCHOGENA, A. Agassiz, 1862, 1865.

With four radial-canals which give rise to numerous blindly-ending side branches. Gonads
leaf-shaped and developed upon the side branches of the radial-canals.

POLYORCHIS, A. Agassiz, 1862 to 1865.

With four radial-canals which give rise to numerous blindly-ending side branches.
Numerous sac-like, sausage-shaped gonads attached to the radial-canals and to
their side branches. Ring-canal simple. Bell-margin not cleft into lappets.

SPIROCODON, Haeckel, lS^g = Goniomeam/rus, Kirkpatrick, 1903.

Similar to Polyorchis, but the ring-canal gives rise to blindly-ending, centripetal branches,
and bell-margin is cleft into lappets.

10 MEDUS/E OF THE WORLD.

Subfamily No. 3, BERENICINJE.
The radial-canals give rise to branches which connect with the ring-canal.

CANNOTA, Haeckel, 1879.

With four radial-canals, each of which gives rise to two side branches which join the
ring-canal.

CuviERIA, Peron, 1807 = Berenice, Haeckel, 1879.

With four main radial-canals which give rise to numerous non-dichotomous side branches.
Gonads on the terminal ramuli of the canals.

DICHOTOMIA, Brooks, 1903.

With four main radial-canals which divide dichotomously two or more times. The
gonads extend outward from the stomach over the canals.

DIPLEUROSOMA, Axel Boeck, i$66 = Tetracannota, Mayer, 1900.

With three or more main radial-canals which give rise to non-dichotomous side branches.
Gonads on the canals adjacent to the stomach. Hydroid: Cuspidella (?).

ToxoRCHiS = Toxorchis + ClaJocanna, Haeckel, 1879.

Four or more main radial-canals which branch dichotomously one or more times. Gonads
on the outer branches of the canals near the ring-canal.

NETOCERTOIDES, Mayer, 1900.

Eight main radial-canals which branch dichotomously. Gonads extend outward from
the sides of the stomach along the radial-canals.

Family No. 2, EUCOPID^.

Leptomedusaewith lithocysts, and with less than eight radial-canals upon which the gonads
are developed.

Subfamily No. i, OBELIN^E.
With eight adradial lithocysts. Four radial-canals. Stomach without a peduncle.

EUCOPELLA, von Lendenfeld, 1883.

Degenerate medusae. No tentacles. No manubrium. Branched radial-canals. Hydroid:

Eucopella.

AGASTRA, Hartlaub, 1897.

Degenerate medusae. No manubrium. Simple, unbranched radial-canals. Hydroid:
Campanularia ( ?).

EUCOPE, Gegenbaur, 1856.

Basal bulbs of the tentacles are simple and hollow, and do not project inward into the
gelatinous substance of bell. Lithocysts on bell-margin. Hydroid: Campanulana.

OBELIA, Peron and Lesueur, 1809.

Entodermal cores of tentacles project inward into the gelatinous substance of the bell.
Otocysts on bases of tentacles. Hydroid : Obelia.

TIAROPSIS, L. Agassiz, 1849.

An ocellus with entodermal pigment above each lithocyst. Tentacle-bulbs simple and
hollow.

Subfamily No. 2, PHIALIN^.

With more or less than eight lithocysts. Four to five radial-canals. Stomach without a
peduncle.

CLYTIA, Lamouroux, \%i2 = Ef>enthesis, McCrady, 1857.

Sixteen tentacles alternating with sixteen lithocysts. Four radial-canals. No cirri.
Hydroid: Clytia.

LEPTOMEDUS.E. 11

PHIALIDIUM, Leuckart, i%$() = 0<:eania, Agassiz, 1862, 1865.

Sixteen or more tentacles. More than sixteen lithocysts. Four radial-canals. No rudi-
mentary tentacles. No cirri. Hydroid: Campanulina.

PHIALUCIUM, Maas, 1905.

Similar to Phialidium, but with permanently rudimentary tentacle-bulbs. Hydroid un-
known.

BLACKFORDIA, Mayer.

Numerous tentacles and lithocysts. Entodermal cores of some or all of the tentacles
project inward into the gelatinous substance. Four radial-canals. No cirri. No
permanently rudimentary tentacles.

PSEUDOCLYTIA, Mayer, 1900.

Five radial-canals, five lips, five gonads. Numerous tentacles and lithocysts. No cirri.

GASTROBLASTA, Keller, \?>?>^=Multioralis, Mayer, 1900.

Two or more manubria. No cirri. The medusa propagates by fission.

EUCHEILOTA, McCrady, 1857.

Four or more closed vesicular lithocysts. Four radial-canals. Marginal or lateral cirri.
Hydroid: Campanulina ( ?).

MnROCOMA = Phialis + Mitrocoma + Mitrocomiurn + Mitrocomella, Haeckel, 1879.

Similar to Eucheilota, but the lithocysts are contained in open folds of the velum. No
entodermal ocelli such as are found in Tiaropsis.

STAUROPHORA, Brandt, lS^ = Staurostoma + Staurof>hora, Haeckel, 1879.

Mouth an open, cruciform, gutter-like slit extending down the four radial-canals.

Subfamily No. 3, EUTIMINJE.

With eight adradial lithocysts. Stomach mounted upon a gelatinous peduncle. Four
radial-canals.

SAPHENIA, Eschscholtz, 1829.

Two tentacles. Numerous cirri.

EUTIMA, McCrady, 1 857 = Eutima + Eutimeta + Octorchis + Octorchandra

-\-Eutnnalphcs, Haeckel, 1879.

Four or more tentacles. Numerous cirri or marginal warts. Four or eight gonads upon
the four radial-canals. Hydroid: Campanopsis (Claus, Brooks).

JLwTIMlUM = Eutimium + Octorchi(lium, Haeckel, 1879.

Similar to Eutima, but without cirri. Hydroid: Campanulina ( ?)

Subfamily No. 4, EIRENINvE.

With more than eight lithocysts. Stomach mounted upon a gelatinous peduncle. Four
or six radial-canals.

PHORTIS, McCrady, 1857.

Four or more tentacles. No cirri. Four radial-canals. Hydroid: P/iortis Brooks.

IRENOPSIS, Goette, 1886, non Ireniopsis, Mayer, 1894.

Six or more tentacles. Six radial-canals. Six lips. Six gonads.

EIRENE, Eschscholtz, i%2g= Irene +Irenittm, Haeckel, 1879.

Four or more tentacles. Numerous marginal warts or cirri. Four or eight gonads
developed upon limited parts only of the four radial-canals. Hydroid: Campan-
ulina ( ?)

TIMA, Eschscholtz, 1829.

Similar to Eirene, but with gonads upon the entire lengths of the four radial-canals.
Hydroid: Campanulina ( ?)

12 MEDUS.E OF THE WORLD.

Family No. 3, ^EQUORIDjE.

Leptomedusae with otocysts, and with eight or more radial-canals.

OCTOCANNA, Haeckel, 1879.

Eight radial-canals, 45 apart. Eight lips. No ocelli. (Is this a young jEquorea ?)

OCTOGONADE, Zoja, 1896.

Similar to Octocanna, but the marginal sense-organs have ocelli as well as lithocysts.

STOMOBRACHIUM, Brandt, lS^^ = Stomobrachium + Staurobrac/iium, Haeckel, 1879.

Eight or more simple, unbranched radial-canals, which arise at equal intervals from the
margin of the stomach. Four lips.

HALOPSIS, A. Agassiz, 1863, 1865.

Radial-canals arise in four groups from the four perradial corners of stomach. Four lips.

^EQUOREA, Peron and Lesueur, i$og = &(juorea + R/iegmatoJes + Mesonema

-\-Polycanna, Haeckel, 1879.

More than eight simple, unbranched radial-canals which arise separately from the mar-
gin of the stomach. More than four lips. Subumbrella smooth, without gelatinous
papilla-like protuberances. Hydroid: Campanulina.

ZYGODACTYLA, Brandt, 1835; sensu Agassiz, 1862.

Similar to jEquorea, but with interradial rows of papilla-like, gelatinous protuberances
upon the subumbrella.

, Haeckel, 1879.

With bifurcated or branched radial-canals which arise at equal intervals from the margin
of the stomach. More than four lips. No peduncle.

ZYGOCANNULA, Haeckel, 1879.

Similar to Zygocanna, but the stomach is mounted upon a gelatinous peduncle.

Order No. 3. TRACHYMEDUS.E Haeckel, 1866.

Medusae with a marginal velum, and with lithocyst concretions of entodermal origin.
With simple uncleft bell-margin.

Family No. i, OLINDIADJE.

Some or all of the tentacles project from the sides of the bell, above the margin, and have
adhesive disks. Gonads linear, sac-like, or folded, and developed upon the four or six radial-
canals. The tentacles arise separately and are not grouped in clusters.

GONIONEMUS, A. Agassiz, 1862, 1865.

All of the tentacles project from sides of bell in a zone slightly above bell-margin. All
have adhesive disks. Four radial-canals. Lithocysts external, on bell-margin. No
centripetal canals. Development through an attached hydra stage. (Perkins.)

CUBAIA, Mayer, 1894.

Similar to Gonionemus, but with two sets of tentacles, one arising from the bell-margin
and the other set projecting from the sides of the bell, as in Gonionemus.

VALLENTINIA, Browne, 1902.

Similar to Cubaia, but with lithocysts inclosed within the gelatinous substance of the
bell, adjacent to the ring-canal, and on the inner side above the velum. (Is this
a young Olindias])

OLINDIAS, F. Miiller, 1861.

Similar to / allentmia, but with blindly-ending, centripetal diverticula from the ring-canal.

TRACHYMEDUS.K. 13

OLINDIOIDES, Goto, 1903.

Similar to OlinJias, but with six radial-canals (two bifurcated and two simple). Six
gonads. Four lips. The exumbrella tentacles project at various levels from the
sides of the bell.

Family No. 2, PETASIDjE Haeckel, 1879.

Trachymedusae with tour radial-canals upon which the linear or sac-like gonads are
developed. Tentacles without adhesive disks. Four lips.

PET ASUS = Pftasus + Dif>etasus + Petas(ita + Pftacli>: uni, Haeckel, 1879.

Tentacles arise at equal intervals, not grouped into clusters. No centripetal canals.
Free marginal hthocyst-clubs.

AGLAUROPSIS, F. Miiller, 1865.

Similar to Petasus, but the lithocysts are vesicular, and project from the bell-margin
between the tentacles.

CRASPEDACUSTA, Lankester, i$%o=LimnocoJium, Allman, 1880.

Tentacles arise singly as in Pi-tnsus and Aglauropsis, but the lithocyst concretions are
each inclosed in a cavity within the gelatinous substance of the velum on the inner
(centripetal) side of the ring-canal. (The medusa lives in fresh water among water-
lilies.) The hydroid is devoid of tentacles.

MlCROHYDRA, PottS, 1885.

Is possibly identical with Limnocodiutn, but the mature medusa is unknown. The
young medusa has no lithocysts, and it arises by budding from a minute hydroid
which has no tentacles.

MJ^OTIAS, OstroumofF, 1896.

Tentacles arise at equal intervals, not in clusters. Numerous centripetal, blindly-ending
canals arise from the ring-canal.

GOSSEA, L. Agassiz, 1862.

Tentacles grouped into clusters. No centripetal canals. Lithocyst concretions free or
inclosed.

Family No. 3, LIMNOCNIDIDjE.

Numerous hollow tentacles which project singly, not in clusters, from the sides of the
bell in a zone slightly above the margin. Tentacles without adhesive disks. Numerous
inclosed lithocysts on the exumbrella side of the velum. Mouth a round opening. Gonads
developed diffusely in the ectoderm of the stomach-wall. Four (occasionally five or six) radial-
canals. Medusa-buds arise from the sides of the stomach, and are set free.

LIMNOCNIDA, Giinther, 1893.

Generic characters are those of the family. The only known species is /,. tanganjicd
from the fresh-water lakes of Central Africa, and the Niger river.

Family No. 4, PTYCHOGASTRIDJE.

Numerous more or less isolated clusters of tentacles, some of which bear adhesive disks.
Numerous free lithocyst-clubs. Eight radial-canals. Four lips. Stomach eight-lobed. These
stomach-lobes are in the radii of the radial-canals, and are bound to the subumbrella by
means of eight mesenterial partitions. The gonads are upon the eight stomach-lobes, and
each is more or less divided by the mesentery so there may be eight double (sixteen) gonads.

PTYCHOGASTRIA, Allman, iSfi^Pcctyllis + Pectis + Pectanthis, Haeckel, 1879.
The generic characters are those of the family.

Family No. 5, TRACHYNEMID^E.

Trachymeduss with eight simple radial-canals upon which the gonads are developed.
No mesenterial partitions in the subumbrella. Tentacles without adhesive disks. Ring-
canal simple without centripetal branches.

14 MEDUSA OF THE WORLD.

Subfamily No. i, RHOPALONEMINjE.

Trachynemidae in which the stomach lacks a peduncle.
RHOPALONEMA, Gegenbaur, iS^6=Trachynema (young medusa), Gegenbaur, 1854

= Trachynema + Rhopalonema + Marmanema, Haeckel, 1879.

With eight well-developed radial tentacles, and eight or more small cirrus-like or club-
shaped interradial tentacles. All tentacles arise in a single row. Eight gonads
localized on the eight radial-canals. Four lips.
SMINTHEA, Gegenbaur, 1856.

Similar to Rhopalonema, but with only eight tentacles, one at the foot of each of the

eight radial-canals.

HOMCEONEMA, Maas, i8g3 = Colobonema, Vanhoffen, igo2 = lsonema (in part), Maas, 1906.
Similar to Rhopalonema, but the tentacles are all of one sort. No small club-shaped or

cirrus-like tentacles. Four lips.
PANTACHOGON, Maas, 1893 (sens. ampl.).

Gonads not localized as in Homceonema and Rhopalonema, but developed diffusely over

the radial-canals. Four lips.
HALICREAS, Fewkes, i$82 = Halicreas + Haliscera, Vanhoffen, 1902

= Isonema (in part), Maas, 1906.

The mouth is a simple round opening, without four lips. (In all known species the
radial-canals and ring-canals are very broad and flat.) Wart-like protuberances
may be present upon the sides of the exumbrella. Radial tentacles large, interradial,
small. Tentacles arise in a single row.
BOTRYNEMA, Browne, 1908.

Similar to Halicreas, but the tentacles are grouped in linear clusters in a single row

around the bell-margin.
CROSSOTA, Vanhoffen, 1902.

The tentacles arise in several rows from the bell-margin. Mouth with four lips.

Subfamily No. 2, AGLAURINjE.

Stomach mounted upon a peduncle.
AGLAURA, Peron and Lesueur, 1809.

Eight gonads upon the peduncle above the stomach. Sexes separate. Development direct.
AGLAmHA = /fglantha + 4glisi:ra, Haeckel, 1879.

Eight gonads upon the subumbrella, or at the turning points of the eight radial-canals

between the peduncle and the subumbrella. Sexes separate.
AMPHOGONA, Browne, 1904.

Similar to Aglantha, but medusa is bisexual, four of gonads being male, and four female.
STAURAGLAURA, Haeckel, 1879.

Four gonads, one upon each alternate radial-canal.
PERSA, McCrady, 1857.

Two gonads on two of the radial-canals, 180 apart. The six other radial-canals are sterile.

Family No. 6, GERYONIDjE.

Trachymedusae with four or six radial-canals upon which the flat, expanded, leaf-like
gonads are developed. Stomach mounted upon a gelatinous peduncle. The ring-canal
gives rise to blindly-ending centripetal canals.
LIRIOPE, Lesson, i%^ = Linantha -\-Lirwpc + Glossocodon -\-Glossoconus, Haeckel, 1879.

Four radial-canals. Four gonads. Four lips. With four primitive, solid, radial, and four
solid interradial, and four hollow, flexible, radially-placed tentacles; all twelve of
which may be found upon the medusa at one and the same time. Development
direct through a free-floating, actinula-like larva.
GERYONIA, Peron and Lesueur, i8og = Geryones + Geryonia + Carmaris

+ Carmarina, Haeckel, 1879.
Similar to Liriope, but with six radial-canals, six gonads, six lips, etc.; instead of four.

NARCOMEDUS.E. 15

Order No. 4. NARCOMEDUS^ Haeckel, 1879.

Veiled medusae with bell-margin cleft into intertentacular lappets. With free lithocyst-
clubs, containing concretions of entodermal origin. These medusx develop from actmula
larvae either directly or by budding. The bell grows outward from the sides ot the body of the
actinula, or the medusa-bud, leaving the tentacles stranded in the partially closed-over clefts
between the lappets of the bell. The Narcomedusae are thus medusiform, actinula-like
animals, the bell of which is not homologous with that of the Anthomedusae or Leptomedusae.

Family No. i, SOLMARIDjC.

Narcomedusae in which the outer margin of the stomach is plain, entire, and without
peripheral stomach-pouches. Saccules may, however, arise from the subumbrella floor of
the stomach.

o/)'fo//>a (young) + Sohnaris, Haeckel, 1879.
Without subumbrella saccules. Gonad is a simple annulus in ectoderm of subumbrella

floor of stomach.
PEGAJXTBA=Pegasia+Polyxenia+Pegantha+SoImoneta (in part), Haeckel, 1879.

With out-pocketings on the subumbrella floor of the stomach. The gonads are developed
in these subumbrella saccules.

Family No. 2, jEGINID^E, sens. ampl.

Narcomedusae in which the central stomach gives rise to simple or cleft marginal out-
pocketings in the radii of the tentacles.

CUNANTHA, Haeckel, 1879.

Four tentacles. Four peronial strands in the tentacular radii. Four simple, uncleft,
peripheral stomach-pouches in the radii of the tentacles. This "genus" is prob-
ably only a young stage of JEgina.
JEciNA, Eschscholtz, i$2g = Cunarcha + gina + SolmunJus, Haeckel, 1879.

Four tentacles. Four peronial strands. Four cleft ( = eight peripheral) stomach-pouches,
outer margins of which may be still further divided.

SoLMUNDELl.A=dlgi>iella + SolmunJella, Haeckel, 1879.

Two tentacles. Four peronial strands. Four cleft ( = eight peripheral) stomach-pouches.
An apical (exumbrella) sense-organ is present in larva, but does not persist in adult.
Derived from jEgina by the disappearance of half ot its tentacles.

HYDROCTENA, DawydofF, 1903.

Two tentacles. No peronial strands. Two simple, uncleft stomach-pouches in the
tentacular radii. There is an apical (exumbrella) sense-organ consisting in a
ciliated pit containing two lithocyst-clubs. A median axial canal extends upward
from the stomach to the bottom of the sensory pit.

CUNOCTANTHA, Haeckel, 1879.

Eight tentacles. Eight peronial strands. Eight simple, uncleft stomach-pouches in the
tentacular radii.

unoctona + ^Eginura, Haeckel, 1879.

Eight tentacles. Eight peronial strands. Eight cleft ( = sixteen peripheral) stomach-
pouches. The outer margins of these pouches may be still further divided so as to
give thirty-two marginal pouches.

S, Brandt, 1835.

Four tentacles. Eight peronial strands. Eight cleft ( = sixteen peripheral) stomach-
pouches. Derived from jEginura by the disappearance of half of its tentacles.

CUNINA, Eschscholtz, 1829.

Nine or more tentacles, and an equal number of peronial strands. Peripheral stomach-
pouches simple, uncleft and equal in number to the tentacles, in the radii of which
they are developed. With otoporpae above the sense-clubs.

16 MEDUS.E OF THE WORLD.

SOLMISSUS, Haeckel, 1879.

Similar to Cunina, but without otoporpae.

CvNlSSA = Cnnissa + ^EginoJorus, Haeckel, 1879.

Nine or more tentacles, and the same number of peronial strands. Peripheral stomach-
lobes twice as numerous as the tentacles, being cleft in the tentacular radii.

jEciNODiscus, Haeckel, 1879.

Eight tentacles, sixteen peronial strands. Sixteen cleft ( = thirty-two peripheral) stomach-
pouches.

MEDUS/E MILLEPORIN^;.

Under this heading we may place the degenerate, free-swimming medusae of
Millepora. They have no velum and are thus separated from the veiled medusae
or Craspedotae. Not only is the velum absent, but the medusa is also devoid of a
peripheral canal system and of marginal tentacles.

The medusae Milleponna? and Craspedotae are doubtless derived from a
common ancestral phylum, but have departed widely, one from the other, so that
the Craspedotae are constantly characterized by a diaphragm-like membrane, or
velum, which partially closes the opening of the bell-cavity at the tentacular margin;
whereas this structure is absent in the Milleporinae.

The only known forms of Medusae Milleporinae are those of Millepora.

Millepora alcicornis (Medusa).

Millepora (medusa*), HICKSON, 1900, Proc. Roy. Soc. London, vol. 66, p. 3, figs. i-io. DUERDEN, 1899, Journal of the Institute
of Jamaica, March, 1899. HICKSON, 1906, Cambridge Natural History, vol. I, Coelenterata, p. 259, fig. 129.

Each medusa lies in an ampulla, or cavity, of the corallum, and is attached by a narrow
stalk to the innermost part of the wall of the cavity. The bell is 0.4 to 0.6 mm. in diameter
and is devoid of radial or circular canals or velum. It consists of a median lamella of entoderm
covered on the exumbrella and subumbrella sides by an ectodermal epithelium. There are
no tentacles, but instead there are 4 or 5 swollen masses of nematocysts 90 or 72 apart, near
the bell-margin, but projecting from the sides of the exumbrella. The manubrium is greatly
swollen, and fills the greater part of the bell-cavity. A mouth may (or may not ?) be present.
There are I to 5, usually 3 or 4, large ova in the ectoderm ( ?) of the manubrium. The central
entodermal cavity of the manubrium consists of an axial chamber which often gives rise to 4
perradial pouches which project into the entoderm of the manubrium, and end blindly. In
some medusae these pouches do not exist, in other specimens there are but 2 or 3, but probably
the most common condition is that of 4 pouches 90 apart. The fully-developed ova occupy
positions alternating with the pouches. Dr. Duerden observed that these medusae are set
free and swim slowly about with infrequent feeble pulsations. Soon after liberation the ova
begin to be discharged into the water and the medusa dies after 5 or 6 hours of life, having
discharged all of its eggs.

The Pacific millepores also produce medusae (see Hickson, S. J., 1891, Quart. Journ.
Micros. Sci., 1898; Proc. Zool. Soc. London; and Philosoph. Trans. Roy. Soc. London, B,
vol. 179).

OKOKIf A.Vmn.MKDI S.K. 17

VEILED MEDUS/E.
MEDUSAE CRASPEDOT^E Gegenbaur, 1856.

Cr\ftacar/iir, ESC'HSCHOLTZ, 1829, Syst. drr Acalc'phac.

Gymnopthalmee t FORBES, 184^, British Naked-eyed MeJuvr.

CnispeJala, GEC-.ENBAI'R, 1856, Zeit. fiir wisscn. Zool., Bd. X, p. 217.

Craspcdotf, HAECKEL, 1879, Syst. der Medusen. MAAS, 1X9}, Ergi-h. ilcr Plankton-Expedition] Bd. 2, K.c.

CHARACTERS OK THK VKII.ED MEDUSJE.

Medusae with a velum or diaphragm which partially closes the marginal opening of the
hell-cavity; with ectodermal gonads (i. e., the sexual products u'lifn ripe are commonly found
in the ectoderm). Without gastric filaments, \\irh a douhle marginal nerve-ring, one ahmi-
and one below the velum. Development either direct or through alternation of generations
from hydroids.

Order ANTHOMEDUS^E Haeckel, 1879.

Oceanitit (in part), GEUENBAUR, 1856, Zeit. fur wissen. Zool., Bd. 8, pp. 218, 219.

Tubularix (in part), ACASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 337.

AnthomeJustr, HAECKEI., 1879, System der Medusen, p. 3. VANHOFFEN, 1891, Zoologischer An/eiRtr, Bd. 14, pp. 439, 442.
MAAS, 1905, Craspedoten Medusen der Siboga Expedition, p. 5; 1897, Mem. M. C. Z. at Harvard Coll., vol. 23, p. 9.
HARTLAUB, 1892, Nachrichten kgl. Gesell. \Vissenschaftcn Gottingen, p. 17; 1897, Nordisches Plankton, Nr. 12, p. 4.
HARGITT, 190^, Bull. U. S. Bureau of Fisheries, vol. 24, p. 29.

CHARACTERS OF THE ANTHOMEDUSyE.

Hvdromedusae in which the gonads are contained within the ectoderm of the manubnum.
There are no marginal otocysts or sensory clubs. The hydroids are of the Tubulanan order.
We may distinguish three families:

(1) CodoniJif with simple tentacles, and with the gonad in the form of one or more rings
encircling the manubrium. No oral tentacles.

(2) CladonemiJtr with feathered or branched marginal tentacles. Gonads ring-like, or
more or less separated. With or without oral tentacles.

(3) OceaniJir with gonads confined to the interradial or adradial sides of the manuhnum
and separated one from another in the meridians of the principal radii. With or without
oral tentacles. With unhranched marginal tentacles.

The bell in the Anthomedusae is without marginal lappets, and usually dome-shaped.
The tentacles arise from the bell-margin, not from the sides of the bell. The velum is simple
and annular, and is provided with circular muscles serving to produce the periodic con-
tractions of the bell. The hell-margin is simple and entire, and there are no otocysts, otohth-
clubs, or club-shaped sense-organs. Marginal cirri rarely present. In the young medusa the
exumbrella is more or less besprinkled with nematocysts, but these usually disappear or
become less conspicuous in the adult; although definite nematocyst tracts often persist over
the exumbrella, this being far more commonly seen in Anthomedusas than in other orders of
Hydromedusae.

The tentacles usually arise singly from the bell-margin, and are usually simple and
unbranched. Ectodermal ocelli are often found upon the tentacle-bulbs, and Romanes
showed that in Sarsia these subserved a visual function, and that the medusa was strongly
attracted by rays between the red and violet. The main shafts of the tentacles are thickly
covered with nematocysts, which may be mounted upon filaments or converted into adhesive
organs, as in the Cladonemidae. The tentacle-bulbs are hollow, and connected with the
gastrovascular system ot the medusa.

The radial-canals are usually simple, but in the Tiarinae they often give rise to short,
blindly-ending diverticula, which may be glandular in function, (ilandular swellings are
found in the walls of the radial-canals of L\tnn<ir,-ii iil,-\iinJri, and probably in those of /)y-
inorpliosa dubia and Sliikbi-ria hnltcrtita. A ring-canal is present in all genera excepting

18 MEDUS.E OF THE WORLD.

Pachycordyle and the Williadi, where it has become filled with a more or less solid core of
entoderm cells. The ring-canal is usually simple, and rarely gives rise to blindly-ending
diverticula.

The proboscis, or manubrium, is usually flask-shaped, and the mouth in most of the
Codonidae is a simple round, or cruciform, opening, but in the Oceamdae it is surrounded by
lips which are more or less folded or crenated. Oral tentacles are found in the Margelmae,
and in some Cladonemidae. The stomach is often mounted upon a solid, conical peduncle,
but in Turritopsis the peduncle is made up of large, highly-vacuolated, entodermal cells which
constitute the walls of the proximal parts of the 4 radial-canals. The medusae are carnivorous,
feeding upon Crustacea, fishes, and other medusae or Siphonophorae.

In the Codonidae the gonad encircles the manubrium in a ring-like manner, the mature
genital products being found in the ectoderm on the sides of the stomach. In the genus
Slabbena there are 2 or more of these genital rings. In the Oceamdae, however, the gonads
are restricted to the interradial, or adradial, sides of the stomach. Haeckel, 1879, believed
that they were radial in position, corresponding with the radial-canals, but VanhofFen, 1891,
showed that this was an error, for they are commonly interradial, or on both sides of each
radial line. In Eleutheria, according to Hartlaub, and possibly in other Cladonemidae, the
genital products are developed in the ectoderm of a peculiar brood-pouch, which is not
connected with the stomach, but is invaginated from the general ectodermal wall of the
subumbrella. In the Williadi and in Nemopsis the radial corners of the stomach extend
outward along the radial-canals, and the gonads develop upon the sides of these pouches.
Eleutheria is successively hermaphroditic, either sex preceding, but in all other Anthomedusae
the sexes are separate. The mature genital products are usually found in the ectoderm,
although they often originate in the entoderm. The eggs are cast out into the water by the
breaking down of the ectodermal walls ot the manubrium, but in some species of Bougatnvillia,
or in Margelopsis, or Hybocodon, the larvae may be retained until they have passed into the
planula or even into the actinula stage. None of the Anthomedusae are known to develop
directly from the egg into medusae, but the hydroid stages of many genera remain unknown.
Wherever the sexual development is known it is through hydroids of the Tubulanan order, in
which the medusae bud out separately and are not protected in special capsules or sporangia.

Asexual development of medusae is found in Codonidae, in medusae of the genera Hv/>o-
codon, Slabberia, and Sarsia, and in the Oceanidae in the genera Cytieis, Dysmorphosa, Bou-
gainvillia, Rtithkea, and in the Williadi. In some of the Williadi the medusa-buds are borne
upon stolons which arise either from the radial corners of the stomach, or from the forks of
the radial-canals. In the case of Sarsia and Hybocodon the budding medusae develop as in
hydroids, the entoderm of the parent medusa forming the entoderm of the budded medusa,
and the ectoderm of the bud being formed from the ectoderm of the parent. In Rathkea,
Chun, 1895, discovered that the budding medusa is formed out of the ectoderm of the parent,
although a connection is finally established between the entoderm of the bud and that of the
parent a short time before the hud is set free. In Bougainvillia tuobe, on the other hand,
I find that the budding medusa is formed entirely from the ectoderm of the parent, no con-
nection being established between the entoderm of the bud and that of the parent. In the
genus Niobia the tentacle-bulbs develop into medusae and are set free.

The majority of the Anthomedusae undergo a considerable development while swimming
freely in the water. In some cases, however, the medusae may, at times, become mature and
even discharge their genital products while still attached to the hydroid. This is seen in
Pennaria, Podocoryne, Sarsia, and Stylactis. Among the most short-lived medusae are those
of Pachycordyle, which have neither tentacles, marginal sense-organs, nor radial nor circular
canals.

All of the Anthomedusae are inhabitants of salt water. They are rare in the open ocean
far from land, but are abundant along coasts, especially along continental shores, and many
of them thrive in harbors where the water may be more or less brackish. They appear to be
the simplest, and phylogenetically the oldest, of the Hydromedusae.

ORDER ANTHOMEDlS.i:. 19

Family CODONIDjC Haeckel, 1879, sens, amend.

Sarsiadat (in part), Founts, 1848, British Naked-eyed Medusa;, p. 55.

Sarsiada+ Tubularidtf, McCRADY, 1857, Gymn. Charleston Harbor, p. 21.

Sarsiadtf (in part)-t- Tubularidx+ Pennaridj: (in part), AGASSIZ, L., 1862, Cont. Nat. Hist. (-'. S., vol. 4, pp. ^9, 542.

SarsiadtE+ Orthocorynidx + Tubulands + Pennanda, AGASSIZ, A., 1865, North Amer. Acalephx, pp. 175, 183, |S6, 189.

Codonidz, HAECKEL, 1879, Syst. der Medusen, p. 9. VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14^.442. HARGITT, 1904, Bull.

Bureau of Fisheries U.S., vol. 24, p. 29. HARTLAL-B, 1907, Nordisches Plankton, Nr. 12, pp. 5, 6.
Codonina+ Cor\nid<z + Tubularidtt, VON LENDENFELD, 1884, Zool. Anzeiger, Bd. 7, pp. 446, 447.

FAMILY CHARACTERS.

Anthomedusae in which the gonad is ring-like and encircles the manubrium. Mouth with-
out oral tentacles or prominent lips. The 4 to 6 radial-canals are simple and unhranched.
\Yhen ocelli are present they are found upon the outer sides of the tentacle bulbs. There
are no marginal otocysts. The tentacles are neither branched nor feathered.

Medusae of the genera Sarsia, Steenstrupia, Ectopleura, Pennaria, Amalthtfa, Trichorhiza,
Hyhocodon, H yJnc/ithys, Coryrutts, and Margt-lopsis are known to develop asexually through
alternation of generations from Tubulanan hydroids. Direct development of medusae from
the egg is unknown in this family, although medusae are budded asexually from the manubrium
of some species of Sarsia, from the basal bulbs of the tentacles of HybocoJon, and from those
of Sarsia. Actinula larvae are set free from the manubrium of Margelopsis and Hybocodon.

Haeckel, 1879, considered the presence or absence of an apical projection upon the bell,
and the presence or absence of an axial canal extending upward from the stomach into this
apex, to be a criterion for the separation of genera. It should be borne in mind, however,
as was pointed out by VanhofFen, 1891 (Zool. Anzeiger, pp. 439-446), that the young medusae
often lack an apical projection and axial canal, whereas the mature individuals possess these
characters. They are also highly variable in development. We have therefore considered
these characters to be of specific but not generic value.

A natural classification of the Codonidae can not be based upon the characters of the
medusae alone, for the medusae of Pcnnana and Amaltheea are similar, while their hydroids
are distinct, the former being Pcnnana and the latter Carymorpha, and an equally remarkable
condition is presented by hydroids belonging to the two genera Syncoryne and StaunJi/i,
both ot which give rise to medusae belonging to the genus Sarsia. This peculiar case is still
further complicated by the fact that at least one species of StauriJia hydroid gives rise to a
ClaJonema medusa.

In HyJrichthys we meet with an extraordinary case of parasitism or commensalism of
the hydroid, and in Margelopsis we find a free-floating hydranth, recalling the hypothetical
ancestral form of the Siphonophorae. Margclnpsis is also interesting in that its medusae form,
apparently, a connecting link between the Codonidae and the Bougamvilleae, although they
are more closely related to the Codonidae than to the last-named family. Indeed their
resemblances to the Bougamvilleae are probably due to mere parallelism and not to blood-
relationship.

The Codonidae may conveniently be divided into two subfamilies, as follows:

(1) Srirsi/inn, with simple, unbranchtd, marginal tentacles, which arise singly from the bell-margin.

One or all of the tentacles may he degenerate or absent.

(2) Margelopstna, with marginal tentacles grouped into clusters.

20 MEDUS.E OF THE WORLD.

The following table defines the characters of the genera ot Codomdae.

SUBFAMILY No. i. SARSIAN^E.

With simple marginal tentacles which arise singly from bell-margin. One or all of the tentacles may be rudimentary or absent.

Tribe I. ANEMIDI.

Without tentacles or chymifcrous canals.

Pachycordyle, WEISMANN, 1883; HARCJITT, \^o^=Parvanenius ( ?) MAYER, 1904.

Medusa without tentacles, marginal sense-organs, radial-canals or circular canal. Hydroid: Pachycordyle ', WEISMANN.

Tribe II. AMALTHIADI.

The hydroid is Corymorpha. Medusae have 4 radially placed, rudimentary tentacles and 4 radial-canals.
Amalthtza, SCHMIDT, 1854. M.edus*= Amafthaa, HAECKEL.

With 4 rudimentary tentacles, 4 radial-canals, and ring-canal. Genital products on manubrium. Hydroid: Corvmorpha.

Tribe III. PENNARIDI.

Medusae similar to those of Tribe II, but hydroid is Pcnnaria.
Pennaria, OKEN, 1815; GOLDFUSS, 1820.

With 4 rudimentary tentacles, 4 radial-canals, and ring-canal. Genital products within manubrium. Hydroid: Pennaria.

Tribe IV. PSEUDOPENNARIDI.

Medusa similar to those of Tribes II and III, but hydroid is Trichorhiza.
Trick or kiza, RUSSELL, 1906.

Medusa similar to Pcnnaria, but hydroid is Trichorhiza.

Tribe V. EUPHYSIDI.

With small or rudimentary tentacles situated at bases of all but one radial-canal, and with one or more well-developed tentacles
situated at base of remaining radial-canal. Bell may or may not possess an apical projection, or an axial canal extending
into it from stomach-cavity.
Sleenstrupia, FORBES, 1846= Euph \sa, FORBES, 1848; Eufthysora, MAAS, 1905.

The bell is radially symmetrical and the 4 radial-canals are all of equal length, i long and 3 short tentacles. Hydroid:

Corymorpha.
ffybocodon, L. AGASSIZ, 1862= Amphicodon, HAECKEL, 1879.

The bell is asymmetrical. One of the 4 radial-canals is long, the one opposed to it short, and the two other canals of inter-
mediate length. One long tentacle arises from the base of the long radial-canal, while the other 3 tentacles are short
In old medusae of Hybocodon one is apt to find 2 or more tentacles at the base of the long radial-canal, and the medusa,
is in the Amphicodon stage. Hydroid: Hybocodon.
Microcampana y FEWKES, 1889.

With 6 radial-canals and 6 tentacles, one of which is well-developed and 5 arc rudimentary. Hydroid unknown.

Tribe VI. DICODONIDI.

With 2 well-developed, diametrically opposite tentacles, and 2 rudimentary tentacles. With or without an apical projection

and with or without axial canal extending upward from the stomach into apex of bell.
Dicoelonium s HAECKEL, 1879.

With 4 simple radial-canals and 4 radially placed tentacles; 2 of the tentacles are long, and 2 short. Gonad is ring like
and surrounds stomach. No lines of nettling cells over exumbrella.

Tribe VII. SARSIADI.

Bell radially symmetrical. With 4 equally-developed, unbranched, marginal tentacles. With or without an apical projection,
and with or without an axial canal extending upward from stomach into apex of bell, (i) The manubrium is encircled
by a single ring-like gonad. (2) With 2 or more ring-like gonads upon the manubrium.
Sarsia, LESSON, i$4$ = S\ndict\on t A. AGASSIZ, 1862; Codonium t HAECKEL, 1879.

Manubrium slender and tubular. Each tentacle-bulb bears an abaxial, ectodermal ocellus. Hydroid: S \ncoryne.

Also Staiindta.
Hydrichthys, FEWKES, 1888.

Medusa similar to Sarsia, but without ocelli upon the tentacle-bulbs. Hydroid: Hvdrichthys.
Eneodonium, HARTLAUB, 1907.

Medusa similar to Sarsia, but stomach is mounted upon a gelatinous peduncle, and tentacles end each in a knob.
Ectopleura, L. AGASSIZ, 1862.

8 longitudinal lines of nematocysts extend from bases of tentacles to apex of bell. Hydroid: Ecto pleura.
Corynitis, McCRAov, 1857.

Manubrium cruciform in cross-section, and bound to the 4 radial-canals by hollow mesenteries. 4 radially placed marginal
tentacles which are covered with wart-like clusters of nematocysts. Their basal bulbs bear ocelli. Hydroid: Syncoryne.
Slabber ia, FORBES, 1846= Difwrena, McCRADY, 1857. Dif>urena + Bathvcodon, HAECKEL.

Similar to Sarsia, but with two or more ring-like gonads upon the manubrium. Hydroid: S\ncor\ne.

SUBFAMILY No. 2. MARGELOPSINJE.

With 4 radiallv-placed clusters of marginal tentacles. No oral tentacles. Gonads ring-like, and encircling the manubrium.
MargelopsiSy HARTLAUB, 1897.

Marginal tentacles are grouped in 4 radially-situated clusters. Gonad ring-like, and surrounds manubrium. No oral
tentacles, mouth is a simple round opening. Medusae develop by budding from a free-floating hydranth, A/drgr/o/fj/i.
This interesting genus recalls the Bougainvillidi in its radially situated clusters of marginal tentacles. It resembles the
Codonidts in its ring-like gonad, and in the absence of prominent lips and oral tentacles.
Pelagohydra, DENDY, 1903.

Medusa similar to Margelopsi* but hydroid is Pelagohydra, and differs from hydroid of Margelopsn in that tentacles arise
at irregular intervals from the sides of the hydranth and are not arranged in definite verticils as in Margelopsis.
Hydroid is pelagic.

PLATE i.

Fig. I. PaehycorJyle degeneratus, male. Nassau Harbor, Bahama Islands,
July 19, 1903.

Fig. 2. Budding hydranth of Pennaria tiarella with female medusa-bud
still attached. Agassiz Laboratory, Newport, Rhode Island,
July 5, 1895. Found attached to eel grass in shallow water.

Fig. 3. A recently liberated male medusa of Pennaria tiarella. Agassiz
Laboratory, Newport, Rhode Island, July, 1895.

Fig. 4. Hydroid stock of Pennaria tiarella, natural size. Agassiz Labora-
tory, Newport, Rhode Island, July, 1895.

Fig. 5. Pennaria tiarella. From the coral reef at Tortugas, Florida, May 7,
1905.

fig. 6. Steenstrupia vtrgulata. From a drawing made from life by Dr.
Alexander Agassiz at Nahant, Massachusetts, August 21, 1862.

Fig. 7. Steenstrupia rubra, male. Oregon Inlet, Pamlico Sound, North
Carolina, November 12, 1904.

Fig. 8. HyhoeoJon forbesii. Tortugas, Florida, May 25, 1905.

Fig. 6, from a drawing by Dr. Alexander Agassiz
Figs, i to 5, 7, 8, from life, by the author.

AXTHOMKIHS K I'A( IIV( -iililiVLK. '_' I

DESCRIPTION OF GENERA AND SPECIES OF ANTHOMEDUSjE.

Genus PACHYCORDYLE Weismann, 1883.

Pachycordylt, WEISMANN, 1883, Entstehung Seiualzellen Hydromedusen, pp. 87, 217. HAKI.II r, 1404, Mitth. Zool. Station

Neapel, BJ. |6, p. 553.
Paminrmus, MAYER, 1904, Mem. Nat. Sci. Museum Brooklyn Institute Arts and Sti., v.il. i, No. i, p. 6.

GENERIC CHARACTERS.

Codonidae without tentacles, radial-canals, or circular vessel. Manubrium surrounded
by a ring-like gonad. The hydroid-stock is Pachycordyle.

Except the medusas of Millepora, these are the most degenerate and short-lived of the
free-swimming Hydromedusae. They are even more degenerate than are the medusae of
Corymorpha and Pennaria, and may be compared with Eucopflla (R. von Lendenfeld, 1883,
Zeit. fur wissen. Zool., Bd. 38, p. 497) and Agastra (C. Hartlaub, 1897, Wissen. Meeresunter-
such. Biologisch. Anstalt auf Helgoland, Neue Folge, Bd. 2, Heft i, Abt. 2, p. 504, taf. 22,
figs. 5, 8 10). In these last-named medusae we find neither manubrium nor marginal tentacles,
hut there are 8 otocysts and a velum, and the radial and circular vessels are well developed.

Pachycordyle weismanni Hargitt.

Pachvcorii\lf lutnmannt, HAROITT, 1904, Mittli. Zool. Station Neapel, Bd. 16, Heft 4, p. 553, plate 21, figs. 1-8.

( ?) Pachyconlyle napolilana, WEISMANN, 1883, Die Entstehung der Seiualzellen bei den Hvdromedusen, Jena, pp. 87, 217.

Medusa pyriform, 2 mm. high, 1.3 mm. wide. Tentacles and marginal sense-organs
lacking. No radial-canals. Ring-canal a mere fissure with vestiges of an entodermal linin
near the margin. Velum narrow, with a small opening. Manubrium large, conical, and with-
out a peduncle. Ripe ova are in the entoderm, and are discharged very soon after medusa is
liberated. Mouth lacking. Manubrium orange or dark-brown, other parts colorless. The
medusa swims with a short, jerky motion, but lives only one or two hours.

The hydroid is found in the Bay of Naples growing upon the shell of Fusus rostratits.
The colony arises from a delicate, reticulated hydrorhiza. Hydrocaulus sparingly branched,
3 to 8 mm. high. Perisarc dull yellowish-brown, not extending beyond base of the hydranth.
Hydranths club-shaped, with subconical hypostome. 8 to 16 irregularly arranged filiform ten-
tacles, delicate and thread-like when expanded. Body of hydranth orange or reddish,
hypostome whitish. Not more than 2 or 3 medusa-buds are borne on the side of the stem
of each budding polyp. Occasionally the medusa-buds develop on the side branches, more
commonly on the main stems. The ova originate in the entoderm where they remain until
discharged into the water from the manubrium of the medusa. This species may possibly be
the female torm of VVeismann's Pachycordyle napolitana. On Weismann's hydroid, how-
ever, there were no free medusae, only sessile gonophores. All of Weismann's specimens were
males, while Hargitt's were females, and it is possible, as Hargitt states, that the females
only give rise to free-swimming medusae.

Pachycordyle degeneratus.

Plate I, fig. I.

Paroanemu* degtneretui t MAVFR, 1904, Memoir-. N.it. Sn. Museum Rrooklvn Institute of Art^. and Sci., vol. i. No. i, p. fi, plate
',. "H- "

Bell thin-walled, about 0.75 mm. high and 0.3 mm. in diameter. Bi-ll-walls quite rigid,
velum powerful and well developed. There are neither tentacles, radial-canals, circular
vessel, nor marginal sense-organs. Manubrium spindle-shaped, and about a third as lon
as height of bell. Fluids within the stomach-cavity are maintained in rapid motion, appar-
ently through the action of cilia. Near aboral end of bell is a deep conical cicatrice which
apparently marks the place of last connection between the medusa and its hydroid stock.
Bell is translucent and milky in color, while manubrium is cream-colored. Only 5 specimens,
all males, were found in Nassau Harbor, Bahamas, on the nights of July 18 and 19, 1903.

22 MEDUS/E OF THE WORLD.

They swam actively in arcs of circles, but all died early in the morning although maintained
in large glass dishes filled with pure sea-water. They appeared to be mature, for sperm was
discharged constantly from the sides of the manubrium.

The hydroid is unknown, and possibly the medusa may not belong to the genus Pachy-
corJyle, though in the medusa stage it conforms with the generic character of the medusae
which are set free from Pachycordyle iveismanni of Naples.

Genus AMALTH^EA Schmidt, 1854.

Corvinorplia (hydroid), SARS, M., 1835, Beskriv og Jagttagelser, p. 7.

Corymorpha (medusa and hydroid), STEENSTRUP, 1854, Vidensk. Meddel. Nat. For. Kjobenhavn, p. 46.

Amahhxa (medusa), SCHMIDT, 1854, Handatlas vergl. Anatomie, p. 13. HAECKEL, 1879, Syst. der Medusen, p. 38.

Amallhiza, ALLMAN, 1872, Monog. Tubularian Hydroids, p. 393. LOMAN, 1889, Tijdschrift, Nederland. Dierk. Vereen, Ser. 1,

Deel. 2, p. 270.
Corymorpha, HARTLAUB, 1905, Zoolog. Jahrb., Abth. Syst., Suppl. 6, p. 543; 1907, Nordisches Plankton, Nr. 12, p. 75.

GENERIC CHARACTERS OF THE FREE MEDUSA.

Codonidae with 4 rudimentary tentacle-bulbs, 4 radial-canals, and a ring-canal. Hydroid
is Corymorpha, but distinguished by the character of its medusae. The majority of species
of Corymorpha hydroids do not give rise to free-swimming medusae. Hydroid of Arnalthtta
might therefore be called Amalthaa. The medusae of Amalthcra resemble those of Pennaria,
but the manubrium is usually longer, and extends beyond the velar opening.

Hartlaub, 1907, includes the medusae of Steenstrufia, Euphysa, and Amalthaa in the
genus Corymorpha, for they all arise from identical hydroids; while the medusae differ as
follows: Steenstrufia, 3 short, and I long tentacle, and with an apical projection and axial
canal above the stomach. Euphysa, similar to Steenstrufia, but without an apical projec-
tion or axial canal. Amalthxa with 4 rudimentary tentacles.

Amalthrea sarsii Allman (Medusa only).

Corymorpha sarsii, STEENSTRUP, 1854, Vidensk. Meddel. Naturhist. For. Kjobenhavn, p. 48. BONNEVIK, 1898, Zeit.fiir wissen.

Zool., Bd. 43, p. 476. HARTLAUR, 1907, Nordisches Plankton, Nr. 12, p. 86, fig. 82.

Amatthtea sarsi:, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 393. HAECKEL, 1879, Syst. der Medusen, p. 38.
( ?) Arnaltluea amtebigera, HAFCKEL, Ibid., p. 38, taf. I, figs. 10, ll.

It is not certain that this hydroid gives rise to free-swimming medusae, though when arti-
ficially set free the medusae swim about actively.

The bell of the attached medusa-bud is elongate with a subconical basal apex and with
fairly thick bell-walls. It is 4 mm. high. There are usually 4 equally developed rudimentary
tentacles, but occasionally one of the tentacles is longer than the others. Velum well developed.
4 straight, slender radial-canals. Manubrium spindle-shaped. Mouth a round opening
studded with nematocysts. In the male the manubrium projects one-third its length beyond
velar opening, but in the female it is shorter and the mouth is at the level of velar opening.
Ova are few in number. When immature they are amoeboid, but when older they become
spherical and project over the surface of the gonad, attached by short pedicels. The gonad
encircles the entire manubrium. Manubrium straw-yellow, tentacle-bulbs light-red. Found
oft coast of Norway. Hydroid: Corymorpha sarsn.

Amalthaea uvifera Schmidt.

Amalthira uvifera, SCHMIDT, 1854, Nordisches Plankton, Nr. 12, p. 88, fig. 83.

Corymorpha uvifera, HARTLAUB, 1907, Handatlas der vergleich. Anatomie, p. 13, taf. 9, fig. 2.

This medusiform gonophore is not known to be set free. It is borne upon branched
stolons which arise from the sides of the polypite above the basal circlet of long tentacles.
Each stolon is tree-like and gives rise to 30 to 40 grape-like medusa-buds, so that each
hydroid gives rise to more than 100 buds in various stages of development.

In the medusa-bud (when about to be set free ?) there are 4 large globular, rudimentary
tentacle-bulbs. The bell is higher than a hemisphere, with walls thin at apex. Manubrium

ANTHOMKDUS.K AMAI.TII.KA, I'KN \.\HI.\. -'.',

spindle-shaped, as long as the depth of the bell-cavity. The hydroid is C.or\in<n [>hn uvifern,
found at Loppen Island, ahout 10 miles from Hammerfest, at a depth of about I fathom.

Amalthaea vardoensis Loman.
Amahh<ta vardiitnsis, LOMAN, 1889, Tijdschr., Nederland. Dierk. Ver., Ser. 2, Dcel. 2, p. 271, teit-fig. 5, taf. 13, figs. 1-9, 15.

Hydroid about 50 mm. high. The thin, transparent perisarc extends only over lower halt
of stem. The terminal polypite is sharply set off from the stem by a constriction at its base.
Nearly 50 large tentacles in basal circlet at wide base of polypite. Over 100 very short tapering
oral tentacles in 7 to 9 circlets. 16 to 20 peduncles arise in a circlet from the sides of the
polypite between the basal and oral tentacles. Each peduncle bears a number of medusa-
buds, which have 4 radial-canals and 4 equally developed, very short tentacle-bulbs. The
hydroid is translucent rose-colored. Found in Busse Sound at Vardri, 71 N. lat., Norway.
For details of histology, see Loman.

Amalthaea (?) Hybocodon(?) januarii Steenstrup.

Corymorpha januarii, STEENSTRUP, 1854, Vidensk. Meddcl. Nat. For. Kjbbenhavn, p. 46. SARS, 1861, Annals and Mag. Nat.

Hist., vol. 8, p. 356.
Amalthtfa januarii, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 394. HAECKEL, 1879, Syst. der Mi-JuM-n, p. 39.

Described by Steenstrup from a single imperfect hydroid found in the harbor of Rio
Janeiro, Brazil. The hydrocaulus is about 150 mm. long and 8 mm. wide. About 80 tentacles
in the proximal circlet, and these are about 50 mm. long. The oral circlet was imperfect
and can not be described. There were about 40 branched peduncles above the basal circlet of
tentacles. These bear numerous medusa-buds having 4 equal tentacle-bulbs, but oblique
margins. The free medusae are unknown.

Genus PENNARIA Oken, 1815.

Pennaria (in part), OKEN, 1815, Lehrbuch der Naturgesch., Bd. I, p. 94.

Pennaria (hydroid), GOLDFUSS, 1820, Handbuch der Zoologic, p. 89.

Pennarin (medusa), MI/CRADY, 1857, Gymn. Charleston Harbor, p. 50.

Pennaria + Halocordyle, ALLMAN, 1871, Monog. Tubul. Hydr., pp. 363, 368.

Pennaria, AOASSIZ, L., 1861, Cont. Nat. Hist. U. S., vol. 4, p. 278. AC;ASSIZ, A., 1865, North Amer. Acal., p. 187. HARTLAUB,

1907, Nordisches Plankton, Nr. 12, p. 72. PEEBLES, 1903, Archiv. Entwick.-mech., Bd. [4, p. 55 (regeneration).
Pennaria, AGASSIZ and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 32, p. 161 .--CLARKE, 1907, Mem.

Museum Comp. Zool. Harvard Coll., vol. 35, p. 6.
GlobicfpSf AC.ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344. VANHOF* EN, 1891, Zool. An/eiger, p. 443. HAECKF.I , 1879,

Syst. der Medusen, p. 39.

Globicfps (hydroid), AVERS, 1852, Proc. Boston Soc. Nat. Hist., vol. 4, p. 193.
Rucorvne (hydroid), LEIDV, 1855, Journ. Acad. Nat. Sci. Phila., ser. 2, vol. 3, part. 2, p. I VI- plate 10, figs. 1-5.

The type-species ot this genus is Pennaria ilistielia ot Goldfuss, 1820, from the Mediter-
ranean. Oken's "Pennaria" is wholly indefinite, including as it does Plumularia, Aglao-
phenia, etc.

(JKNERIC CHARACTERS.

Codonidae with 4 permanently rudimentary tentacles, which are reduced to mere basal
bulbs. The hydroid stock is a Pennaria.

Medusae of the genus Pennaria may become mature, and discharge their genital products
while still attached to the hydroid, and at the same time other individual medusae from the
same stock may be set free in an immature state. Generally, however, the genital products are
discharged a few hours after the medusae are set free into the water.

There is no generic difference between the medusae of Pennaria and medusae belonging
to the genus Amalthtsa, but their hydroids are different. The hydroid stock of Amalthaa
is Corvnifjrpha, and is closely related to the hvdroids of the medusa genera Hybocodon, St., n
strupia, and Ectopleura. The hydroid of the medusa genus Pennariti McCrady is Pennaria
Goldfuss (Globiceps Ayers). On account of this decided difference in their hydroids we
have separated Pennaria from Amaltheea = (Corymorpha\ for their apparent similarity is
only a case of parallelism.

24

MEDUSAE OF THE WORLD.

Pennaria disticha Goldfuss (European).

( ?) Pennaria marina, IMPERATO, 1599, Dell' historia naturale libri ventoto, Napoli, p. 747.

Pennaria cavoiini, SCHNEIDER, 1892, Jena. Zeitsch. Naturw., Bd. 20, p. 435, taf. 14, figs. 49-54 (histology). Du PLESSIS, 188
Mittheil. Zool. Sta. Neapel, Bd. 2, p. 147. PHTET, 1893, Revue Suisse Zool., tome i, p. 12, plate I, figs. 7-9. WEI

, 1881,

is-

1903,
colonies).

colonies).
Pennaria disticha) BEDOT, 1901, Revue Suisse Zool., tome 9, p. 459; tome 13, 1905, p. 96 (citation of papers to 1850). GOLD-

fucc iSii-i T-Ian^hiirh t\e*r 7.rwil . 11 Hn

FUSS, 1820, Handbuch der Zool., p. 89.
Pennaria symmetric a, CLARKE, 1879, Bull, Mus. Comp. Zool. at Harvard College, vol. 5, p. 240, plate I, figs. 2, 3.

Pennaria , PEEBLES, 1903, Archiv. Entwickelungs-mech., Bd. 14, p. 55, fig. (regeneration).

Sertularia fiennaria, DELLE, CHIAJE, 1841-44, Animali senza vertebra del Regno di Napnh, vol. 5, pp. 5, 17, plate 157, figs. 3,

14, 15; Ibid., 1822, tav. 43.

(tySertuIaria pennaria, LINN^US, 1758, Systema Naturae, edit. 10, p. 813.
Seriolara ftennaria, CAVOLINI, 1785, Mem. Polipi marini, Napoli, p. 134, plate 5.
Globiceps globator, HAECKEL, 1879, Syst. der Medusen, p. 40.
( ?) Medusa, Euphysa globaior, LEITKART, 1856, Archiv. fur Naturgesch., Bd. 22, p. 28, taf. 2, fig. 4.

Fin. i . Pennaria disticha, from nature, by the author, from specimens collected by Dr. Lobianco in the Bay of Naples.
A y hydrocaulus, natural size. B, terminal branches showing law of growth. C, terminal polypite. D and , hydranths
showing completely-annul.ited pedicels.

ANTHOMKDUS.E PENNARIA. I'.")

This hydroid is one of the earliest forms known, yet reliable information from the system-
atic standpoint is still incomplete. One of the best modern descriptions is that <>t Allman,
1872. I believe the American " Penmn-in tinn-lln" to be closely related to P. Jiiliilm.
The only difference appears to be that the terminal ramuli in P. disticha are ringed through-
out, whereas in P. tiarella they are usually ringed only at base and summit, although I have
seen an occasional one ringed throughout in the American hydroid. (See VV. S. Wallace,
1908, Year Book of the Carnegie Institution.)

In the European hydroid, P. Jisticha, the stems become about 150 to 175 mm. high. The
main stem is slightly zigzag and with a uniform growth-curve from base to summit. There are
about forty side branches, regularly alternate. Perisarc annulated at the origins of the branches
and on the ultimate hydranth-bearing ramuli. Hydranths flask-shaped, with a single verticil
of about 10 to 13 basal filiform tentacles each about I to 2 times as long as the body of the
hydranth, and each ending in a blunt, slightly swollen tip. About 20 short, stiff, knobbed
tentacles, irregularly arranged in 3 verticils, arise from the sides of the hydranth above the
basal verticil of tentacles. The medusa-buds are similar to those of Pennaria tiarella and
are without ectodermal ocelli upon their tentacle-bulbs. The medusa? usually wither upon
the hydroid without being set free, but this often occurs also in P. tiarella, especially in the
warm water of Florida.

Weismann, 1883, found that the germ-cells originate in the ectoderm of the inner layer
of the bell-nucleus and do not wander from their place of origin, becoming mature in the
free medusa, or when the medusa-bud is ripe.

Very elaborate studies upon regeneration, regulation, and restitution in injured colonies
were carried out by Cast and Godewski, 1903.

This hydroid is found in the Mediterranean. Pictet, 1893, found it at Amboina, Malay
Archipelago. It occurs at Naples, Italy, from May to November.

Pennaria tiarella McCrady (American).

Plate I, figs. 2-5.
REFERENCES TO THE AMERICAN HYDROID.

G/obiceps tiarella, AOASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344. AVER, 1852, Proc. Boston Soc. Nat. Hist., vol. 4,
p. 193. HAECKEL, 1879, Syst. der Mejusen, p. 39.

Pennarin tiarella, AGASSIZ, A., 1865, North. Amer. Acal.,p. 187, figs. 311-315. NUTTING, 1901, Bull. U. S. Fish Commission
for 1899, vol. 19, pp. 337, 374, figs. 14, 83. HARGITT, 1900, Amer. Naturalist, vol. 34, p. 387, plates 1-4, 36 figs.;
Bull, of the Bureau of Fisheries U. S., for 1904, vol. 24, p. 32, plate 3; 1904, Archiv. fur entwickrlungs-mech. organ-
ism, Bd. 18, p. 453, taf. 24-28; 1902, Amer. Naturalist, vol. 35, p. 31 1, figs. 8, 9; p. 597, fig. 36; 1899, Biol. Bulletin
Woods Hole, vol. i, p. 35-40, 6 figs, (grafting eiperiments); 1900, Science, New series, vol. 12, p. 340; and 1901,
Biol. Bulletin Woods Hole, vol. 2, p. 223 (variations of hydroids and medus.T.) MCCRADY, 1857, Gymn. Charleston
Harbor, p. 51. SMALLWOOD, 1899, American Naturalist, vol. 33, p. 861, 7 figs, (histology). HARTLACB, 1907, Nor-
disches Plankton, Nr. 12, p. 72, figs. 68-70. GOLDFARB, 1906, Journ. Experimental Zool., vol. 3, p. 148 (regeneration).

Pennaria liarclla=(P. svmmetrica, CLARKE), CONGDON, 1907, Proc. American Acad. Arts and Sci., vol. 42, p. 464.

Pennaria, j/>., THACHER, 1903. Biol. Bulletin, vol. 4, p. 96.

I'tnritiria gibbosa, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 278, 344; vol. 3, plate 15, figs. I, 2.

Halcord\Ie tiarella, ALLMAN, i87i,Monog. Tubul. Hydroiils, p. 369.

Kuforyne elegant, LEIDV, 1855, Marine Invert. N. J. and R. I., p. 4, plate 10, figs. 1-5.

THE AMERICAN PENNARIA.

Medusa. The bell is about 2 mm. in height, and is ellipsoidal, being higher than it is
broad. Bell-walls very thin, flexible, and much distorted by the remarkably large ova, which
are situated within the ectoderm of the manubrium and often fill the entire cavity of the bell.
There are 4 small, rudimentary tentacle-bulbs, without ocelli, I at the base of each radial-
canal. Radial-canals straight and quite broad. Velum wide and powerful. Manubrium
of male more slender than that of female, the latter being usually greatly distended with 4 or
5 large ova. Several ova are often set free into the water before the medusa is liberated from
the hydroid stock. The medusa is commonly mature at the time of its liberation, and it is
probable that it does not usually survive for more than a few hours, though Dr. A. Agassiz
has maintained them alive for several weeks. The entoderm of the manubrium is rose-pink,
and there are a number of deep-pink blotches in the entoderm of each radial-canal. The
entoderm of the tentacle-bulbs is pearly-white, as are also the eggs within the manubrium.
These colors vary considerably in hue and may be almost white.

2fi

MEDTS.E (>K THE WOULD.

Hydroid. The hydroid is very abundant upon eel-grass, seaweeds, stones, or wharves,
below low tide. The stems arise from a ramifying stolon. They attain a height of 100 to 125
mm. The main stems branch alternately, the longest and oldest side branches being found
nearest the base of the stock. The side branches give rise to simple or slightly branched

ramuli from their upper sides. These ultimate
ramuli are annulated at base with 5 to 6 rings, and
the main stem and primary branches are annulated
beyond each branch. The stems are covered with
a chitinous, horny perisarc. The polyp-heads are
flask-shaped, and the mouth is situated at the ex-
tremity of a long, slender, conical throat-tube.
There are 2 or 3 verticils of oral tentacles, each
row being composed of 5 to 7 short, stiff tentacles,
each terminating in a knob-shaped cluster of nem-
atocysts. Besides these there is a single circlet of
12 to 16 long, flexible tentacles near the proximal
base of the polypite. The medusa-buds develop
upon the sides of the polypites between the oral
and basal rows of tentacles. There are usually not
more than 2 or 3 buds upon each polypite at the
same time. The entoderm of this hydroid is white
to rose-colored, the ectoderm silvery and translu-
cent, and the perisarc horny-yellow to black.

This hydroid ranges from the West Indies and
Bermudas to the coast of Maine. Pentiaria gibbosa
L. Agassiz, of the Florida coast and West Indies,
is probably identical with P. tiarella, but the
medusae of the southern form wither on the stem
as do those of P. Jisticha. Indeed Agassiz's figure
in 1862, shows the ultimate pedicels of his P. gibbosa, from Key West, Florida, ringed
throughout as in the Mediterranean Pennaria disticha.

Hargitt, 1900, has made a study of the life-history and development of P. tiarella. Early
in the summer the hydroids are found growing on rock-weed, piles of docks, etc.; while late
in the summer they take to eel-grass. The early summer brood is not so bright in color, and
the medusae mature more slowly than in the late brood. The individual hydranths of the
late brood are, however, smaller than those of the early summer brood. The medusae dis-
charge their ova soon after liberation, and then die rapidly. The ova are 0.4 to 0.5 mm. in
diameter and are heavily laden with yolk. They are creamy-white to orange in color. The
cleavage is found by Hargitt to be subject to much individual variability, so that an extra-
ordinarily irregular, loosely connected mass of cells is formed, resembling the condition
described by Rittenhouse, 1907, in Turritopsis nutricula. No matter how irregularly shaped
the embryo may be, it results finally in the formation of a spherical solid morula, and this
soon changes into a pear-shaped, ciliated, planula larva. In 5 to 7 days after the beginning
of development, the larvae settle down and then develop into small monogastric hydroids
covered with ringed perisarc, and provided with whorls of tentacles as in the adult. Hargitt
finds that the removal of small parts of the segmenting eggs does not alter the future history
of development. The eggs may also be divided at the first or second cleavage and will still
develop into normal larvae. Moreover, Hargitt finds that single eggs sometimes separate
normally into two masses, each of which gives rise to a normal embryo. The irregularities
in cleavage observed by Hargitt have been seen also by Miss Cora J. Beckwith. She finds
that the segmentation is mytotic, not amitotic; the nuclear division constantly outnumbering
the cytoplasmic so that a syncytium is formed.

Hargitt, 1901, states that hydroid stocks of Pennaria which grow upon eel-grass in shallow
water are more pinnatified in their branching, and more highly colored than are those which
grow upon stones, etc., in deep water. He also finds that the medusae of the deep-water
hydroids are less active than are those developed upon the hydroids which grow in shallow

Flo. 2. Southern variety of Pennaria tiarella. From
life, by the author, Tortugas, Florida, May
27, 1908.

Branch of hydroid and enlarged view of a polypite
showing pedicels ringed only at their bases.

ANTHOMEDfS.E I'ENNAHI \. 27

water, and are often retained until they wither upon the stems after having cast out their
genital products.

I have found only the pale form of Pennarin on the Florida Reefs, and its medusae appear
to cast out their genital products before being set free, as is described by Hargitt for the deep-
water forms of Woods Hole, Massachusetts. In Florida, Pomona grows in tuft-like clusters
upon gorgonians. This tropical form is called Pennana gtbbosa by L. Agassiz, but those seen
by me can not be separated from the Pennaria of the coast of New England.

Hargitt, 1899, carried out a series of grafting experiments upon Pennaria and other
hydroids. He finds that pieces of hydroids of the same species may readily be grafted one
upon the other, both in oral or aboral relations, there being little or no evidence of polarity
in the regenerative process. Pieces of opposite sex but of the same species will readily unite
in any manner, but pieces belonging to hydroids of different genera would not unite. Elaborate
studies of this character were also carried out by Gast and Godewski, 1903, on P. Jisticha.

In 1900 and 1901, Hargitt studied the variations of the medusx and hydroids of Pen-
naria. The medusa sometimes displays ectodermal blister-like protrusions on its exumbrella,
and is variable in other respects.

According to H. Miiller, 1907 fZeit. fur wissen. Zool., Bd. 89), the eggs of Pennaria
develop at the expense of the weaker egg-cells of the ovary, which they engulf as food to form
the yolk-granules.

Thacher, 1903, shows that in Pennaria, Campaniilana, and EiiJenJnurn, the hydranths,
when they degenerate, are absorbed not by liquefaction of their protoplasm, or by the with-
drawal of the polyps as a whole; but absorption takes place by the degenerating cells of
both ectoderm and entoderm being turned into the digestive tract of the hydroid.

Goldfarb, 1906, finds that light is absolutely essential for the normal growth, develop-
ment, and regeneration of Pennaria. He finds, however, that this is true in a sense also for
Eudendrium, but not to the same degree, for Eudendrnim ramositm colonies, kept in the
dark until all the influence of their previous illumination has been lost, will not then
regenerate new hydranths unless they be again exposed to light; but the surprisingly short
exposure to light of only 5 seconds will suffice to restart the regenerative process.

Pennaria rosea von Lendenfeld.

Pennaria rosea, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, ser. I, vol. 9, p. 51)4, plate 24, figs. 40, 42.
BALE, 1888, Ibitl., ser. 2, vol. 3, p. 747 7*. australii, BALE, 1884, in Cat. Australian Hydroid Zoophytes, p. 45.

Main axial stems 80 mm. long, with about 20 alternately arranged, pinnate side branches.
4 to 6 hydranths on each branch. Hydrorhiza and main stems intensely black and opaque;
outer half of each stem light-yellow, proximal half intense black. Hvdranths rose-colored,
with 9 to 14 oral, and 7 to 12 filiform, basal tentacles. Medusas are produced on the proximal
hydranths. The medusa bell is slender, oval, 2 mm. high, I mm. wide. 4 large rudimentary
tentacle-bulbs with a minute external ocellus upon each. Manubrium with sperm or ova fills
the entire cavity of subumbrella. Ova discharged after medusa is set free. Color, intense rose.

Coast of New South Wales, Australia. Mature in May.

This species is distinguished from the European and American Pennaria by its tentacu-
lar ocelli.

Pennaria adamsia von Lendenfeld.

Pennaria aaamsia, VON LENDENFF.LD, 1884, Proc. Linnean Soc. New South Wales, vnl. 9, p. 595, plate 25, figs. 45-48; plate 26,
fig- 49-

MeJusir. The male medusae are 1.5 mm. long and only 0.7 mm. wide; while the female
medusas are 1.5 mm. long and as broad as they are high. The male medusas have 4 radially
situated marginal tentacles, about as long as the width of the bell. These tentacles have
well-developed basal bulbs with minute ocelli, and the tentacles move about freely. The
spermatozoa fill the space between the manubrium and the sides of the subumbrella, and are
discharged within an hour after the medusa has been set free. The female medusas are
broad, and the tentacles are mere rudiments without ocelli. The bell-cavity is filled with
ova, which are soon discharged.

28 MEDUS.E OF THE WORLD.

H ydroid. The stems are 60 to 80 mm. long, zigzag, and with 15 to 20 alternate branches.
The longest branches are nearest the base and are about 12 mm. long. There are about 8
hydranths, 1.7 mm. long, on each branch. These hydranths are slender, the proximal ones
only about half as large as the distal primary one on each branch. There are 2 verticils of oral
tentacles, each with 4 tentacles alternating in position with those of the other row. These
oral tentacles are short and each ends in a nematocyst-knob. The uppermost row of 4 oral
tentacles is in the 4 perradii, and the lower row in the 4 interradii. There are 24 long
filiform tentacles in a single row at the broad base of the polypite. These are about as long
as the polypite itself. The perisarc of the hydrorhiza and main stems is opaque and black,
and the side branches are yellow. The hydranths are white.

Coast of New South Wales, Australia. Medusae are produced in March.

This species is distinguished by the minute ectodermal ocelli upon the tentacle-bulbs
of the male medusae, and by the length of its tentacles. There are also but 2 verticils of
oral tentacles.

Pennaria pacifica Clarke.

Pennaria pacifica, CLARKE, 1907, Mem. Museum Comp. Zool. Harvard Coll., vol. 35, No. l, p. 6, plate I, figs. 1-6.

Hydrocaulus 20 to 35 mm. high. Internodes of the main stem without annulations at
their distal ends, and with only I or 2 rings at their proximal ends. Branches alternate.
Peduncles taper slightly to the base where there are 2 or 3 annulations. Hydranths with
12 to 14 filiments, and about 16 capitate tentacles somewhat irregularly arranged.

Pinco Island, Gulf of Panama, tropical Pacific.

Distinguished only by its few annulations at the internodes of the main stem. It is
probably only a local variety of P. tiarella.

Pennaria ? vitrea Agassiz & Mayer.

Pennaria vitrea, AGASSIZ and MAYER, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 32, p. 161, plate I, figs, l, 2.

Bell is 3 mm. in height; walls thick and rigid. There are 4 rudimentary tentacle-bulbs.
Velum not prominent. The 4 radial-canals are straight and narrow. The manubrium in the
female specimen was retracted within the cavity of the bell, but it was flask-shaped in the
male and projected for a short distance beyond the velar opening. These conditions, how-
ever, may be merely different states of contraction and not constant differences; but as we
observed only two individuals, one a male and the other a female, we can not be certain upon
this point. The ova are large and pyriform, and are grouped in 4 interradially arranged
clusters within the manubrium. The mouth-opening is simple, and there are no fimbriations
or appendages.

Prominent circular muscles were observed in the ectoderm ot the cavity of the bell in the
female, but these were not seen in the case of the male. It is possible, however, that they
become apparent only during certain states of contraction. In the female the ova and tentac-
ular bulbs were flesh-colored, and the entoderm of the mouth was green. In the male the
tentacular bulbs were green, the entoderm of the manubrium pink, and the lips green.

Found among the Fiji Islands, off Kimbombo Island, November 25, and ofF Mbatiki
Island, December 5, 1897.

Not having seen the hydroid I am very doubtful concerning the generic identity of this
medusa with Pennaria.

Genus TRICHORHIZA Russell, 1906.

Trichorhiza, RUSSELL, 1906, Proc. Zool. Soc. London, p. 99, plate 5.

GENERIC CHARACTERS.

The medusa is similar to that of Pennaria, but with one tentacle-bulb somewhat better
developed than the 3 others. The hydranth is solitary, with a branched, filiform hydrorhiza.
The perisarc forms a cup into the cavity of which the hydranth may be partially retracted.
There are 2 verticils of tentacles, an oral and a basal; and the medusa-buds arise from the
sides of the hydranth between these 2 verticils. The type-species is Trichorhiza brunnea
Russell, from the Firth of Clyde, Scotland.

ANTHOMEDUS.E TRICHORHIZA, STEENSTRUPIA.

29

Trichorhiza brunnea Russell.

Trichorhiza brunnta, RUSSKLL, 1906, Proc. Zool. Soc. London, p. 99, plate 5, figs. 1-2; Abstract, Pruc. Zool. Soc. London,
No. 26, p. 6, Feb. 13, 1906.

Mature medusa unknown. When newly set free from the hydroid the medusa is pyriform,
with a thin-walled bell, well-developed velum, and 4 rudimentary tentacle-bulbs, one of
which is larger than the others. The manubrium when extended is as long as the depth of
the bell-cavity, and it is cylindrical, narrow, and provided with a simple, circular mouth-
opening which is surrounded by nematocysts. There are 4 simple, straight radial-canals,
but no gonads. The manubrium and tentacle-bulbs are yellow, the former being faint and
the latter golden in color.

This medusa is liberated by a solitary hydroid which was found on June 29, 1905, at a
depth of 17 fathoms in Ethick Bay, Bute, Firth of Clyde, Scotland. The hydrorhiza of the

Trichorhiza was entwined among and
around the tentacles of Coryrnorpha
nutans. The hydroid is 1 1 mm. in total
length, the hydranth itself being only
1.5 mm. long. The hydrorhiza is long,
filiform, and sparingly branched, with
about 6 simple branches which arise
from its lower (aboral) half. The
coenosarc does not appear to extend
over this branched lower half of the
hydrorhiza. Above the upper end of
the hydrorhiza the perisarc extends to
form a cup-like hydrotheca, which
bears 4 transverse grooves. Immedi-
ately below this hydrotheca the peri-
sarc displays several longitudinal lines.
Hydranth conical, 1.5 mm. long and
0.8 mm. wide, with an oral circlet of 7
very short tentacles having knob-
shaped ends. There is also a basal
circlet of 12 filiform tentacles, which
are ringed with nematocysts and are
tapering, and fully as long as the hy-
dranth itself. 8 or 10 medusa-buds
arise at various levels from the sides
of the polypite between the basal and
oral circlets of tentacles.

The perisarc of the hydrorhiza is
straw-colored, and that ot the hydro-
theca chocolate-colored. Body of hy-
dranth pale reddish-brown, tentacles
translucent white.

This hydroid bears some resemblance to Blastothela of Verrill, 1878. Amer. Jour. Sci.,
ser. 3, vol. 1 6, p. 374.

FK.. 3. Trichorhiza brunnea, after Russell, Proc. Zool. Soc., London, 1900.
Hydroid, and enlarged view of recently liberated medusa.

Genus STEENSTRUPIA Forbes, 1846.

Sleenslrufia, FOR BBS, 1846, Annals and Mag. Nat. Hist., vol. 1 8, p. 287. LKI:CKART, 1856, Archiv. fiir Naturgesch., Bd.22, p. 29.
Slrrnstrupia + Euf>h\!a, HAF.CKEL, 1879, Syst. der Mcdusen, pp. 29, 3 i . BROWN t, 1895, Trans. Liverpool Biol. Association,

vol. 9, pp. 247, 248.
Eu/ihysa Sticnitrufia, AI.ASSIZ, L., 1862, Com. Nat. Hist. I'. S., vol. 4, pp. 542, \^\. HAM KM , 1X64, Jena. Zeitsrli. fur

Naturw., Bd. I, pp. 338, 339. -FoRin.s, \X^, Hntisll N,iked-eved Medusa-, pp. 71, 72.
Euph\ia, A(.ASSIZ, A., 1865, North Amer. Acal., p. 1*0. UK. MOW. i'jc>4, Bull. Mus. Comp. Zool. at Harvard Coll., vol. VI.

p. 251. VANHOKFFN, 1891, Zool. An/eiger, lid. 14, p. 443.
Ku/>h\sora, MAAS, 1905, Hydromeduscn der Siboga Kipediiion, p. 6.
Heterosirf>hanus + Stcenslrupia+ Eaf>h\sa, HARTLALB, 1907, Nordischcs Plankton, Nr. 12, pp. 74, 76, 80.

30

MEDUS.E OF THE WORLD.

The type-species ot this genus is Steenftrupia rubra Forbes, 1846, from the Atlantic
coasts of Europe.

GENERIC CHARACTERS.

Codonids with radially symmetrical bell, and with 4 radially situated tentacles, I of
which is well developed while the 3 others are short or rudimentary.

This genus is separated from Hybocodon by its radially symmetrical bell. In Hybocodon
i radial-canal is long, I short, and 2 of medium length; but in Steenstrupia the 4 radial -canals
are all of the same length.

Synopsis of the Characters of MeJuste of the Genus Steenstrupia.

S. rubra Forbes=

S. aurata= Euphy-

S. tetrabrachia=E.

S. bigelowi Maas.

S. australis von

S.flaveola Forbes

sa aurata Forbes,

tetrabrachiaH.B.

Lendenfeld.

= S. galanthus

= E. aurata+E.

Bigelow.

Haeckel =(?).

mediterranea

gracilis Brooks=

Haeckel = (?) E.

( ?) S. cranoides

virgulata A.

Haeckel=S. 1m-

Agassiz.

eata Leuckart.

Shape and size

Bell has well-devel-

Dome-like apex.

Pyriform. 4 high,

Pyriform. 10 high,

Half-egg-shaped,

of bell in mm.

oped apical pro-

Cylindrical sides.

2.5 wide.

5 wide.

2.5 high, 1.7

jection. 5 high,

6 high, 4.5 wide.

wide.

3 wide.

Bell-walls and

apex thick.

Condition of ten-

3 long, narrow ten-

3 small, similar,

3 short tentacles,

3 short tentacles

3 tentacle-bulbs,

tacles. Length

tacle-bulbs with-

rudimentary ten-

each= r long, i

about 2 r long

and i long tenta-

in terms of bell-

out tentacles, all

tacle-bulbs, i

large tentacle

and not ringed.

cle, ringed with

radius (r).

similar, i long

long tentacle.

about 7 T long.

One longtentacle

nematocysts.

tentacle ringed

Length = 2 r+ .

All tentacles

4 T long, with

Long tentacle

with nematocysts.

No ocelli. No

sparsely ringed.

many prominent

about 8 r long.

Length = 7 r.

definite rings of

partial rings of

nematocysts on

nematocysts.

tentacles.

Axial-canal above

Axial canal always

No axial-canal.

No axial-canal.

Present. Small,

No axial-canal.

stomach.

present.

slender.

Length of manu-

1 .5 r. Stomach

1.25 r. No ped-

2. 5 r long. Mounted

1.5 r long. Mouth

1.5 T long. Cylin-

brium in terms

mounted on short

uncle.

on very short ped-

not quite reach-

drical.

of bell-radius

peduncle.

uncle ? Mouth

ing level of velar

(<).

extends slightly

opening.

beyond velar

opening.

Gonads.

Single gonad en-

Single gonad en-

Eggs developed in 4 wide interradial,

Gonad encircles

circles stomach,

circles stomach.

8 adradial, longi-

ectodermal swell-

manubrium.

leaving mouth and

tudinal rows, in

ings on sides of

peduncle free.

ectoderm on sides

mid-region of man-

of stomach.

ubrium. Separated

by 4 narrow, per-

radial spaces.

Color.

Tentacle-bulbs and

Variable. Tenta-

Gonads brownish-

Entoderm of man-

Mouth deep violet.

stomach light-

cle-bulbs may be

yellow. Mouth

ubrium, canals,

4 brown patches

pink to reddish-

yellow, orange,

pink. Tentacle-

and tentacles dull

on gonad. Tenta-

brown. Bell apex

red, or colorless.

bulbs and rings

yellow .

cle-bulbs and long

sometimes faint

Manubrium red-

pink.

tentacle brown

pink .

dish to yellow or

with violet spots.

faint purple.

Where found.

Atlantic coasts of

Mediterranean and

Suvadiva Atoll,

Malay Archipelago.

Harbor of Sydney,

Norway, Ger-

Atlantic coasts of

Maldive Islands,

New South

many, England,

Europe.

Indian Ocean in

Wales, Australia.

Scotland, and Ire-

January.

land. (Mediter-

ranean?) (South-

ern United States,

Atlantic coast?)

Hydroid.

Corymorpha nu-

Corymorpha nana,

Unknown.

This medusa is pro-

Unknown.

tans, Sars, 1835;

Alder, 1857.

bably identical

Hincks, 1868;

with S. tetrabra-

Allman, 1871.

chia. Hydroid

unknown.

ANTHOMEDUS.E STEENSTRUPIA. 31

The hydroid generation is Corymorpha Sars, i$3$ = Halatractus of Allman, 1871.

Haeckel, 1879, restricts Steenstrupia to describe medusae having characters as above,
but with an apical projection upon the bell, and with an axial canal extending upward into
this projection from the stomach. Euphysa he would restrict to include forms which lack an
apical projection and an axial canal. As Vanhoffen, 1891, rightly states, an apical projection
is always lacking in young medusae and is highly variable in its development even in mature
individuals of the same species. The same is true of an axial canal. Moreover, among
species discovered since Haeckel wrote his "System der Medusen," Euphysa tetrtibrachin,
Bigelow, 1904 (Bull. Mus. Comp. Zool. at Harvard Coll., vol. 39, p. 251, plate I, fig. i), has
a well-developed apical projection, and no axial canal, and would therefore have no place in
Haeckel's system.

Maas, 1905, would institute a genus Euphysora to describe Codonidae having I large,
and 3 well-developed but still considerably smaller tentacles. Here again, however, we meet
with a condition of degree only. When, for example, are the 3 small tentacles large enough
to be Euphysara or small enough to be Steenstrupia ? In order to avoid confusion, it appears
best to combine all of these forms under one generic name. Vanhb'ffen, 1891, suggests Euphysa
as the proper generic name to include all of these forms, but Steenstrupia takes precedence
over Euphysa, for it was used by Forbes in 1846, while Euphysa was first used by him in 1848.

Bigelow finds that the eggs of S. tetrabrachia are arranged in 8 adradial longitudinal lines
in the ectoderm of the stomach, and Maas states that in S. bigelou'i the gonads are interradial
and separated by very narrow linear interspaces in the 4 principal radii. These conditions
are interesting, for they foreshadow those characteristic of the Oceanidae where the gonads
are interradial or adradial, and are often quite widely separated in the principal radii.

Hartlaub, 1907, defines Heterostephanus (Allman, 1871) as a Steenstruf>ia-\\ke medusa-bud,
not known to be set free from its Coryrnorpha-\\\ie hydroid. Medusa with a single tentacle,
ringed at its base, and terminating distally in a knob. The only known species is H . annuli-
cornis, Allman, l^l=Hybocodon annulicornis Haeckel. This was first described by M.
Sars, 1859; as Corymorpha ? annulicornis.

Steenstrupia rubra Forbes.

Plate I, fig. 7.

Corymorpha nutans (hvdroid), SARS, M., 1835, Beskriv. og Jagtt., p. 6, plate i, fig. 3; SARS, O., 1877, Fauna Littor. Norveg.,

tome 3, p. 2, taf. 2, figs. 25-28.

Corymorpha nutans (hvdroid), BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 63 (literature to 1850).
Corymorpha nutans (hydroid and medusa), HARTLATB, 1907, Nordisches Plankton, Nr. 12, p. 76, figs. 72-75 (complete li

authors, localities, and best modern description of hydroid and medusa).
Corymorpha nutans, HINCKS, 1868, Hist. British Hvdroid Zoophytes, p. 127, plate 22. AI.LMAN, 1871, Monog. Tubularian

Hydroids, p. 388, plate 19.
Corymorpha nuians = S. galanthus (Haeckel)=.S. faveola, Forbes = S. rubra, Forbes, BROWNE, 1896, Proc. Zool. Soc. London,

pp. 463, 464, plate 16, fig. i (review of literature).
Steenstrupia galanthus, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 148 (literature to 1850). HAECKEL, 1879, Syst. ilrr

Medusen, p. 31. Hybocodon ntttans, Ibid., p. 34. MULLER, 1908, Zeit. fur wissen. Zool., Bd. 89, p. 52, taf. 5, figs. 20-24

(origin and structure of the eggs).
Steenstrupia rubra, HARTLAUB, 1904, Wissen. Meeresuntersuch. Kommiss. Meerc Kiel, Abth. Helgoland, Neue Folgc, Bd. 5,

p. 105, fig. 3.

Steenstrupia rubra (Forbes), BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9, p. 247.

Steenstrupia rubra (medusa)+S. faveola, FORBES, 1848, British Naked-eyed Medusa, pp. 73, 74, plate 13, figs, i, 2.
Steenstrupia cranoides, HAECKEL, 1879, Syst. dcr Medusen, p. 30, taf. 2, figs. 10-14.
Steenstrupia gracilts, BROOKS, 1882, Studies Johns Hopkins Vniv. Biul. Laboratory, vol. 2, p. 144. MAYER, 1900, Bull. Mus.

Comp. Zoo!, at Harvard Coll., vol. 37, p. 29, plate 16, figs. 36, 37.
Steenstrupia lintata, LEUCKART, 1856, Archiv. fur Naturgesch., Bd. 22, p. 29, taf. 2, fig. 6. SPACNOLINI, 1876, Catali-i:" \

Mediterraneo, p. 17, taf. I, figs. 1-4. HAECKEL, 1879, Syst. der Medusen, p. 30. DU PLESSIS, 1888, Recueil /.<.!. s,n re,

tome 4, p. 543.

This medusa is found on the coasts of Europe from Norway southward to the Mediter-
ranean. S. cranoides Haeckel = .V. lineata Leuckart, from the Mediterranean, appears to be
identical with S. rubra. S. gracilis Brooks, of the Atlantic coast of the United States south
of Virginia, is probably identical with S. rubra of Europe, but the hydroid of the American
form remains unknown. I believe that 5. Un<-ntn, S. rrtinoiJfs, and .V. i>r/ii-ilis are identical, and
that they are probably identical with .V. ruhrn Forbes.

For description of i'. rubra of Europe, see synoptic table of species of Steenstrupia.
3

32

MEDUSAE OF THE WORLD.

In the American form the development of nematocyst-rings upon the long tentacle is
subject to great variability. The Mediterranean S. cranoides Haeckel lacks such rings,
whereas they appear to be better developed in northern Atlantic specimens from the coasts
of Europe. The same difference appears to be exhibited by our American specimens, those
from Tortugas, Florida, being unnnged or only slightly ringed, while those from North Car-
olina are often profusely ringed. The apex of the bell often bristles with nematocysts, but
in some medusae it may be smooth.

Browne, 1896 (pi. 16, fig. i), gives a figure of S. rubra derived from specimens found by
him at Valencia Island, off the Irish coast. He shows a narrow conical peduncle above the
stomach, the peduncle being about one-fifth as long as the gastric portion of the manubrium.
No such peduncle has been observed in the American S. " gracilis," when the bell is expanded,
although when it is somewhat contracted the appearance of a well-developed peduncle is
often produced. On the other hand, when the stomach is widely distended with food and the
bell expanded no peduncle may be seen. Haeckel's series of figures (taf. 2, figs. 10-12) will

3>

FIG. 4. Corymorpha nulans, after Hartlaub, in Nordisches Plankton.
(Young hydroids and mature medusa.)

serve to illustrate the formation of a peduncle-like body of gelatinous substance above the
stomach through contraction of the bell. I have frequently seen the same phenomenon in
our American S. " gracilis" = S. cranoides Haeckel. I have also observed this peduncle in
living medusae of S. rubra taken off the coast of Cornwall, England.

Hartlaub, 1907, gives a list of the bibliography and of localities for this species, and his
description of the medusa and the young hydroid are the best yet produced (see fig. 4).

The egg is amoeboid as in Amalth&a. The young hydroid has a single circlet of 4 short,
knobbed, oral tentacles, and another circlet of 5 to 8 simple, flexible, filiform basal tentacles.
H. Miiller, 1908, finds that the full-grown eggs are very few in number, having developed at
the expense of other weaker egg-cells in the ovary, which they devour. The exoplasma is quite
wide and is separated from the germinal vesicle. The ooplasma is a network of delicate fibers
of wide mesh, and the exoplasma and endoplasma are distinct, one from another. The egg
contains numerous pseudo-cells in advanced stages of degeneration and also yolk-granules.

ANTHOMEDUS.E STEENSTRUPIA.

33

The following description of the medusa is derived from a study of specimens found by
the author off the coast of the United States:

The bell is 5 mm. high and surmounted by a slender conical projection about 2 mm.
long. There are 4 tentacles. One of these is about 10 mm. long, and is ringed at irregular
intervals by prominent swellings, between which there are small rings at fairly regular inter-

FIG. 5. Corymorfha nuians, hydroid and medusa, after Allman, in Ray Society, 1871-72.
A' Detail showing manner in which medusa 1 bud off from hydranlh.

34

MEDUSAE OF THE WORLD.

vals. The tentacle which is diametrically opposite to the long tentacle is tapering, and about
0.25 mm. long; while the 2 other tentacles are mere bulbs. The velum is well developed.
There are 4 narrow radial-canals, and a slender ring-canal. A long, slender, sinuous
canal extends from the aboral apex of the stomach upward into the apical projection of the
bell. In mature medusae the manubrium extends a short distance beyond the velar opening.
Ordinarily the mouth is a simple, round opening at the tapering extremity of the manubrium,
but when widely open, as in our figure, it shows 4 thick but not prominent lips. The genital
products are found in the manubrium, and in the female the eggs project from the surface of
the ectoderm. Entoderm of manubrium intense yellow-green and rose-color. Apical canal
intense yellow, often flecked with rose-color. The entoderm of the tentacle-bases is rose-
color and yellow, while the entoderm of the large annular swellings is rose-color. When
young the apical projection is not very high, the tentacles short, and without nematocyst-
nngs, and the manubrium short and tapering; not extending beyond the velar opening as
in the mature medusa.

6.

FIGS. 6 and 7. Sieenslrufia rubra.

6. From life, by the author. Off Mousehole, Cornwall, England, October 23, 1907.
7. A. After Leuckart ("S. lineata"), 1856, Archiv. fur Naturgesch., Bd. 22.
B. After Spagnolini ("S. lineata"*), 1876, Catalogo Acalefi Mediterraneo.

Found at Oregon Inlet, Pamlico Sound, North Carolina, in November, and at Beau-
fort, North Carolina, and Tortugas, Florida, in summer. It is apparently identical with
S. cranoides and S. lineata of the Mediterranean.

Definite rings of nematocysts are not found upon the tentacles of the young medusa, and
are very variable in their development in mature specimens, some being profusely ringed
and some entirely unringed. Haeckel describes only unringed individuals from the Medi-
terranean, his specimens being similar to those found by me at Tortugas, Florida.

I am inclined to believe that this Mediterranean and tropical American medusa will
prove to be identical with, or only a variety of, S. rubra of the Atlantic coasts of western
Europe. Certainly no differences, other than those well within the limits of common vari-
ability, can be detected between the medusae of S. rubra and S. lineata = S. cranoides; but a

ANTHOMEDUS.E STEENSTRUPIA. 35

careful comparative study of the hydroids of these forms must be made before we may safely
assert either that they are identical or separate species.

The hydroid of S. rubra is Corymorpha nutans of Sars, and is common on sandy bottoms,
off the northern coasts of Europe, at moderate depths. The stems of the hydroid are
solitary, and are about 50 to 75 mm. high, and about 4 mm. wide at the widest part. The
whole stem is corrugated by numerous narrow longitudinal bands. The widest part of the
stem is usually at a short distance above the lower end. This lower end is bent sharply at
right angles to the main part of the stem and bears long, hair-like filaments which serve to
anchor the hydroid. There are also blunt, papilla-like processes which arise from the sides
of the stem above the bent portion. The polypite is large and flask-shaped, and has a basal
zone of 30 or more long, tapering tentacles, about as long as the polypite itself. In addi-
tion to these tentacles there are 6 to 7 closely crowded verticils of oral tentacles, which are
much shorter and thinner than the proximal. 15 to 20 branched peduncles arise from the
sides of the polypite close to the bases of the proximal circlet of tentacles, and bear the
medusa-buds. The hydranth is light-red, the stem being paler than the polypite. On the
English coast the medusa-buds are set free during the summer.

Allman gives a good description of the hydroid.

Steenstrupia aurata.

Euphysa aurata, (medusa) FORBES, 1848, British Naked-eyed Meduss, p. 71, plate 13, fig. 3.

Euphysa aurata (medusa)+E. mcditcrranea, HAECKEL, 1879, Syst. der Medusen, p. 32, taf. 2, figs. 8, 9.

Euphysa medilrrranca, DU PLESSIS, 1888, Recueil Zool. Suisse, tome 4, p. 543.

Euphvsa aurata (Forbes), BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9, p. 248; 1896, Proc. Zool. Soc. London, p. 474.

Euphvsa aurata, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 749. BEDOT, 1905, Revue Suisse de Zool., tome 13,

p. 134.
Corymorpha nana, (hydroid) ALDER, 1857, Cat. Zooph. Northumberland and Durham, p. ill, plate 7, figs. 7, 8. HINCKS, 1868,

Hist. British Hydroid Zooph., p. 130, plate 22, fig. 3.

Corymorpha nana, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 81, figs. 76-78 (list of authors and localities).
(?) Euphysa virgulata, AGASSIZ, A., 1865, North Amer. Acal., p. 189, figs. 316-319.

This medusa is found off" the Atlantic coasts of western Europe and in the Mediter-
ranean. Steenstrupia virgulata of Massachusetts Bay is probably identical with S. aurata.
For description of the European form, see synoptic table of characters of the species of Steen-
strupia. The European S. aurata appears to be smaller than the American S. virgulata.

Steenstrupia virgulata = (?) S. aurata Forbes.
Plate i, fig. 6.

(?) Steenstrupia aurata, FORBES, 1848, British Naked-eyed Medusae, p. 71, plate 13, fig. 3.

Euphysa virgulata, AGASSIZ, A., 1865, North Amer. Acal., p. 189, figs. 316-319. HAECKEL, 1879, Syst. der Medusen, p. 33.
NUTTING, 1901, Bull. U.S. Fish Com m., vol. 19^.370. HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 33.
Corymorpha virgu/aia, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 84, fig. 79.

Adult medusa. The bell is pyriform, with a broad, dome-shaped apex. It is 5 to 12 mm.
in height, and 4.5 to 9 mm. in diameter. Surface of exumbrella smooth and without rows of
nematocysts. There are 4 tentacles, one at the base of each radial-canal; 3 of these are mere
rudimentary bulbs, but the fourth is large and conical. Its surface is thickly covered with
nematocyst-cells of large size. There are 4 simple, straight radial-canals, and a slender,
circular vessel. The velum is wide, with an indented, free edge. The manuhrium is cylin-
drical without a peduncle, and extends about half the distance from the inner apex of the
bell-cavity to the level of the velar opening. No apical, axial canal. The mouth is a
simple, round opening without prominent lips. The genital products are contained within
the ectoderm of the manubrium. Manubrium light-yellow. The entoderm of the radial-
canals near the bases of the tentacles is intense pink, and the ectoderm of the tentacles is
milky-white. This species was found by Dr. Alexander Agassiz, in Massachusetts Bay, at
Nahant, and is recorded from Woods Hole, Massachusetts, by Nutting and Har^itt.

The figure here shown is reproduced by his kind permission from Dr. Alexander Agassiz's
drawing of the medusa. There is no difference between this medusa and S. aurata of Europe
except that the American form appears to be larger; but the hydroid of the American medusa
is unknown, and until this is discovered it will be impossible to determine the identity of the
American form.

36

MEDUSAE OF THE WORLD.

Steenstrupia tetrabrachia.

Euphysa telrabrachia, BIGELOW, H. B., 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 251, plate I, fig. I.
(?) Euphysora bigelowi, MAAS, 1905, Craspedoten Medusen der Siboga Expedition, Monog. 10, p. 7, taf. I, figs. 1-3.

Bell 4mm. high, 2. 5 mm. wide. Pyriform,with solid apical projection. One long sparsely-
ringed tentacle about 4 times as long as the bell is high. 3 other, smaller tentacles, each about
one-third as long as the bell-height. These bear each about 3 rings of nematocysts, while the
long tentacle bears about 6 such rings. Velum well developed. Manubrium large, spindle-
shaped, with mouth projecting beyond the velar opening. Gonads on sides of stomach.
Eggs arranged in 8 fairly distinct rows. Bell colorless. Gonads brownish-yellow. Manubrium
pinkish. Tentacle-bulbs and rings on tentacles rose-pink. Suvadiva Atoll, Maldive Islands,
Indian Ocean; in January.

This medusa is probably redescribed by Maas as E. bigelowi.

Steenstrupia bigelowi.

Euphysora bigelowi, MAAS, 1905, Craspedoten Medusen der Siboga Expedition, Monog. 10, p. 7, taf. i, figs. 1-3; 1906, Revue

Suisse de Zool., tome 14, p. 84, pi. 2, figs, i, 2. MULLER, 1908, Zeit. fur wissen. Zool., Bd. 89, p. 59.
(?)Euphysora tetrabrachia, BIGELOW, 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 251, plate i, fig. I.

Fir,. 8. Steenstrupia tetrabrachia, after Bigelow,
in Bull. Museum Comp. Zool. at Harvard
College.

FIG. 9. Steenstrupia bigelowi, after Maas, in
Hydromedusen Siboga Expedition.

Bell 13 mm. high; more than twice as high as it is wide, and with a well-developed
apical projection. Bell-cavity 10 mm. deep. Apical projection 3 mm. high. Side walls thin.
4 tentacles at the bases of the 4 radial-canals. One of these tentacles is longer than the bell-
height and bears about 30 swollen nematocyst-warts. The 3 other tentacles are each one-
third to half as high as the bell and are tapering, without nematocyst-warts, but covered with
diffuse nettling cells. Manubrium spindle-shaped, not extending beyond the velar opening.

ANTHOMEDUS.E STEENSTIU'I'IA, HYBOCODON. 37

In most specimens there is a blindly-ending axial canal extending into the gelatinous substance
of the apical projection, but this is not constantly present. The gonads are developed upon
the sides of the stomach, leaving only the basal and mouth ends of the manubrium free. A
cross-section shows that they are separated by 4 minute, perradial, longitudinal lines. There
are therefore 4 interradial gonads. There are no medusa-buds produced by the medusa.

The entoderm of the manubrium, radial-canals, and tentacle-bulbs is filled with yellow
pigment granules. The nematocyst clusters on all 4 tentacles are red.

Found quite commonly in the Malay Archipelago, at Sulu, Ternate, Damar, Manifa,
Saleyer, and Amboina.

This form differs from the closely allied S. letrabrachia Bigelow, from the Maldive
Islands, in that in the Maldive species there are a few rings of nematocysts upon the ten-
tacles, whereas in S. bigelou'i there are only warts, not inclosed rings. Also there appears to
be no axial canal in S. tetrabrachia, whereas this is usually seen in 5. bigelowt. The size
constitutes a disparity in the two medusae 4 mm. high in S. tetrabrachia and about 13 mm. in
S. bi^elowi. Future studies will probably show that these distinctions are not of specific
value, but merely changes due to growth and variation, and that the two medusae are identical
and should be called S. tetrabrachia. "Euphysa" tentaculata, Linko, 1905 (Zool. Anzeiger,
Bd. 28, p. 214), from Barents Sea has also 3 well-developed tentacles, and is 5 mm. high,
with orange-colored manubrium and oval bell. It may be Hybocodon pcndula.

Steenstrupia australis.
Euphysa ausiralis, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 586, plate 21, fig. 33.

Bell is 2.5 mm. high, 1.7 mm. wide. Half-egg-shaped and symmetrical. No lines of
nematocysts over the exumbrella. One very long retractile tentacle, 2 to 3 times as long as
the bell-height. This tentacle has a large basal bulb, and is covered with rings of nemato-
cysts. The other 3 tentacles are mere basal bulbs terminating in a knob-shaped cluster of
nematocysts. Velum well developed. 4 straight radial-canals. Manubrium arises from the
center of the umbrella cavity, and is cylindrical and about half as long as the bell-height.
The gonad encircles the manubrium. Mouth deep violet. 4 brown patches upon the gonad,
and a few brown spots on manubrium near its base. Tentacle-bulbs and the large tentacle
brown with violet spots. Port Jackson, New South Wales, in May and June. Rare.

Hydroid unknown.

Genus HYBOCODON L. Agassir, 1862.

Htbocodon, AGASSIZ, L., i86z, Cont. Nat. Hist. U. S., vol. 4, p. 243. AGASSIZ, A., 1 865, North Amcr. Acal., p. 193. VANHOFFEN,
1891, Zool. Anzeiger, Bd. 14, p. 443. BROWNE, 1896, Proc. Zool. Soc. London, p. 466. HARTLAI B, 1905, Zoolog. Jahr-
buchern, Suppl. 6, p. 544; 1907, Nordisches Plankton, Nr. 12, p. 96.

HybocoJon + ArnphicoJon, HAECKF.L, 1879, Syst. der Medusen, pp. 33, 35.

Amphirodon, BROWNE, 1901, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 275.

Diplura, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 326.

Corymorpha, AGASSIZ, A., 1865, /feiW., p. 192.

This genus was established in 1862 by L. Agassiz for Hybocodon prohfer, a medusa
which arises by budding from a Corymor^/ja-like hydroid on the New England coast. Accord-
ing to Browne, 1896, this medusa is also found off the northern coast of Europe. It is prob-
able that the same medusa was described by Steenstrup, 1842, from Iceland, as Corymorpha
fritillaria, but the hydroid from which Steenstrup supposed this medusa to be derived is
certainly not HvbocoJon, but may be an Arnalthira or Diplura. Steenstrup does not figure
a basal circlet of tentacles upon the polypites, which have only an oral circlet, and below
this a circlet of medusiform gonophores, each with a 4-sided bell, and 4 equally developed
rudimentary tentacles. It is probable that the hydroids of Hybocodon differ more among
themselves than do the medusae, and no final classification of the medusae can be attempted
until all of the hydroids have been discovered.

GENERIC CHARACTERS.

Codonidae with asymmetrical bell. One of the 4 radial-canals is long. I short, and 2 of
medium length. There are I or more long tentacles at the foot of the long radial-canal;
and 3 small or rudimentary tentacles, I at the foot of each of the 3 other radial-canals. The
hydroid is HybocoJon.

38

MEDUS.E OF THE WORLD.

Haeckel, 1874, instituted the genus Amphicodon to include medusae with 3 rudimentary
tentacles, and a cluster of 2 or more long tentacles at the foot of the longest radial-canal.
Vanhoffen, 1891, Browne, 1896, and Hargitt, 1901, have pointed out, however, that the
young medusae commonly have but I long tentacle, and that others appear, and develop from
the side of the basal bulbs of this original tentacle; thus the genus "Amphicodon" is only
a mature Hybocodon. These secondary tentacles may appear before or after the medusa-
buds begin to develop upon the tentacle-bulbs. According to Hargitt, 1902, 1904, Perkins,
1904, and Linko, 1905, the sexual products of the manubrium become mature while
medusa-buds are still being produced upon the tentacle-bulbs.

Synopsis of the Species of Hybocodon.*

H. prolifer L. Agassiz.

H. pendula Haeckel =
Corymorpha pendula
L. Agassiz.

H. forbesii Mayer.

H. unicus Browne.

Shape and size of bell

Dome-shaped. 2.5 high,

Quite similar to H. pro-

Ellipsoidal, asymmetri-

Bell-shaped. 3 high,

in mm.

2.2 wide, i radial-

lifer. 5 high, 3.5

cal. 2.5 high, 2.1

2 wide.

canal long, 2 inter-

wide.

wide.

mediate length, and i

short. Long and short

are 180 apart.

Number of longitudi-

2 extend upward from

As in H. prolifer.

None.

?

nal lines of nemato-

base of well-developed

cysts on exumbrella

tentacle, and I from

base of each of 3 rudi-

mentary tentacle-bulbs.

Condition of well-de-

i to 3 well-developed ten-

Only i well-developed

Only i well-developed

Only i well-developed

veloped tentacle, or

tacles at base of longest

tentacle. No medusa-

tentacle. No medusa-

tentacle "between 2

cluster of tentacles,

radial-canal. Medusa-

buds.

buds.

rudimentary basal

at base of longest

buds are produced at

bulbs." No medusa-

radial-canal.

bases of these tentacles.

buds.

Often only i well-

developed tentacle pro-

duced.

Condition of 3 rudi-

Mere basal bulbs.

i mere basal bulb. 2

i short conical tentacle.

3 mere basal bulbs.

mentary tentacles.

small tentacles. Small

2 mere basal bulbs.

tentacles 90 and

Short conical tentacle

basal bulb 180 from

1 80 and basal bulbs

large tentacle.

90 from long tentacle.

Gonads.

Developed over ectoderm

On ectodermal sides of

On manubrium. No

?

of manubrium. Actinula

manubnuni. No ac-

actinulx attached to

larvs develop upon sur-

tmulie seen attached

gonad.

face of gonad in ecto-

to gonad.

derm of manubrium.

Color.

Entoderm of tentacle-

Pink granules in tenta-

Entoderm of tentacles

?

bulbs, radial-canals,

cle-bulbs. Entoderm

and manubrium yel-

and stomach orange to

of manubrium pink

low, flecked with red.

deep blood-red.

and lilac.

Where found.

North Atlantic coasts of

Coast of New England,

Bahama Islands, and

Stanley Harbor, Falk-

Europe, Iceland, and

United States.

Florida coast. Com-

land Islands.

America.

mon in spring months,

at surface.

Hydroid .

Hydroid described by L.

Hydroid is Hybocodon

Unknown.

Hydroid may be H.

Agassiz, 1862, as H.

pendula= Corymorpha

chilensis Hartlaub,

prolifer.

pendula L. Agassiz.

1905.

*For description of H. chilensis Hartlaub, H. chrislinf Hartlaub, H. pu/cher Hartlaub, and H. (?) januarii, see text.

Hybocodon prolifer L. Agassiz.
Plate 2, fig. i; plate 3, fig. 3.

(?) Corymorpha fritillaria, STEF.NSTRUP, 1842, Generations-wecksel., p. 20, taf. I, figs. 41-46 (the medusa only may be identical

with //. prolifer; the hydroid appears to be an Amaltheea or Diplura).
( ? ) Stcenstrupia globosa, SARS, 1859, Christiania Vidensk. Selsk. Forhandl., p. 101. KOREN and DAWELSSEN, 1877, in Sars's

Fauna Littoralis Novegiie, tome 3, p. 20, taf. i, figs. 1-6.
(?) Corymorpha annulicornis (young medusa), KOREN and DANIELSSEN, 1877, Ibid., p. 8, taf. i, figs. 7-13.

PLATE 2.

Fig. I. Hybocodon prolifer. Woods Hole, Massachusetts, March 4, 1907.
This drawing shows one of the normal aspects of the medusa,
with the bell somewhat contracted. When the bell is extended the
mouth comes to the level of the velar opening.

Fig. 2. Hybocodon pendula. Newport, Rhode Island, April 23, 1897.

Fig. 3. Hybocodon forbesii. Tortugas, Florida, June 26, 1906.

Fig. 4. Dicodoniurn jeffersoni. Tortugas, Florida.

Fig. 5. Dicodoniurn floriJana. Tortugas, Florida, June 17, 1897.

Drawn from life, by the author.

PLATE 2

ANTHOMEDUSJE HYBOCODON. 39

Hybocodon prolijer (hydroid and medusa), AOASSIZ, L., 1861, Cent. Nat. Hist. U. S., vol. 4, pp. 143, 343, plate 133, figs. 10,

11; plate 25, 19 figs.
Hybocodon proltffr, AGASSIZ, A., 1865, North Amer. Acal., p. 193, figs. 325-328. ALLMAN, 1871, Monog. Tubul. Hydroids,

p. 422. VERRILL, 1873, Invert. Animals, Vineyard Sound, p. 736, plate 38, fig. 282. BOHM, 1878, Jena. Xritschr. fiir

Naturw., Bd. 12, p. 195, taf . 7j figs. 7-9. NUTTING, 1901, Bull. U.S. Fish Commission, vol. 19, p. 341, fig. 76. HARGITT,

1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 33, plate 2, fig. 2; 1901, American Naturalist, vol. 35, p. 580, fig. 39;

Ibid., 1902, vol. 36, p. 552; 1901, Biol. Bulletin Woods Hole, vol. 2, p. 222. PERKINS, 1904, American Naturalist 38,

p. 516 (simultaneous sexual and asexual reproduction).
(?) Hybocodon prolijer, BROWNE, 1896, Proc. Zool. Soc. London, p. 466.
Hybocodon prolifer + H.annulicornis + ? Amphicodon fritillaria+ ? A. giobosus + A.amphipleurus, HAKCKEL, 1879, Syst. JerMcdu-

sen, pp. 33, 35, 36, 37, taf. I, figs. 7-9.

(?) Amphicodon fntillaria, BROWNE, 1895, Trans. Liverpool Biol. Soc., vol. 9.

(?) Amphicodon grtnidum, LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 215 (simultaneous sexual and asexual reproduction).
(?) Hybocodon prolifer, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 752.
Non Hybocodon prolijer, BONNEVIE, 1899, The Norwegian North Atlantic Expedition, 1876-1878, vol. 26, Hvdroida, p. 28, plate

i, fig. 6.
Hybocodon prolijer, HARTLAUB, 1907, Nordisches Plankton, Mr. 12, p. 98, figs. 94-97. MULLER, H., 1908, Zeit. fur wissen.

Zool., Bd. 89, pp. 62, 73 (origin and structure of the eggs).
( ?) Hybocodon gravidum + H. islandicus + H . amphipleurus, HARTLAVB, 1907, Nordisches Plankton, Nr. 12, pp. 104, 106,

figs. 99, 100.

The following description is derived from studies made of medusae from the southern
coast of New England, United States:

Adult medusa. Bell about 2.5 mm. high and 2.2 mm. wide. It is asymmetrical, the side
bearing long tentacles being longer than the other sides; or as Browne aptly describes it,
the margin is not at right angles to the longitudinal axis of the bell, but slopes toward the side
bearing the cluster of long tentacles. The 4 radial-canals are of lengths corresponding to
the sides of the bell. The canal leading to the cluster of long tentacles is the longest; while
the canal diametrically opposite to this is the shortest, the 2 other canals being of inter-
mediate length. There are 3 small, rudimentary tentacle-bulbs, I at the foot of the shortest
canal, and I at the foot of each of the intermediate canals. The cluster of tentacles at the
foot of the long radial-canal has wide, hollow, tapering basal bulbs. The mam shaft of each
tentacle is, however, slender, cylindrical, and contractile, and is annulated at regular intervals
by well-developed clusters of nematocysts. Young medusae commonly have but a single long
tentacle, but as growth proceeds they sometimes develop another and finally a third; and
all 3 grow to be of equal length, and form a conspicuous cluster. A number of medusa-buds
in various stages of development arise from the sides of the hollow base of the one or more
long tentacles, near the bell-margin. These medusa-buds themselves develop a single long
asymmetrical tentacle even before the bud is mature. When ready to be set free they resemble
the parent medusa in that they are sometimes observed to be developing a third generation
of medusae upon their tentacle-bulbs. 5 longitudinal lines of nematocyst-cells extend from
the bases of the tentacles to near the apex of the bell. 3 of these rows arise from the bases
of the 3 rudimentary tentacles, and extend up over the surface of the exumbrella immedi-
ately over the radial-canals. In addition to these there are 2 rows which start from the base
of the well-developed tentacles, and extend upward over the exumbrella surface on both
sides of the long radial-canal. The velum is well developed. The radial-canals are narrow
and straight. The manubrium is a simple tube, which usually extends about two-thirds
the distance from the inner apex of the bell-cavity to the velar opening. The mouth has
4 short lips with their edges surrounded by nematocysts. The mature eggs are found in the
ectoderm of the stomach, and there they develop into actinula larvie before being set free.
The entoderm of the tentacle-bulbs is intense orange. The rows of nematocysts upon the
exumbrella often display an orange tinge, as does also the entoderm of the stomach.

Hydroid. The hydroid was found in Massachusetts Bay. growing in tide-pools where
the water was very pure. Stems about 50 mm. in height. They grow singly, or in small clusters,
and do not branch. Each stem terminates distally in a single large polypite. The stems
are not more than I mm. in diameter at the base, gradually enlarging toward upper end,
and are about 3 mm. in diameter at base of polypite. They are covered with a delicate sheath
of chitinous perisarc, which widens and displays several well-developed annulations near
the base of the polypite. The polypite is flask-shaped with a very broad base; the mouth is
situated at the extremity of a narrow cylindrical neck, which is capable of much distension.
There are 2 oral verticils, each composed of about 1 6 tentacles. The tentacles of the row

40

MEDUS.E OF THE WORLD.

near the mouth are only about half as long as those of the lower circlet. In addition to these
there are about 25 long, tapering, hollow tentacles in a zone surrounding the base of the
polypite. During the breeding season, which occurs from January until May, great numbers
of medusae are developed upon the sides of the polypite immediately above the circlet of
basal tentacles. The budding medusae arise singly from the sides of the polypite and are not
produced in clusters upon peduncles as in Hybocodon christina, H. chilensis, and H. pulcher.
Longitudinal bands of orange pigment extend up the stem of the hydroid. The entoderm

FIG. 10. Hydroid, young, and budding medusse of Hybocodon prolifcr, after L. Agassiz, in Con. Nat. Hist. U. S., vol. 4.

(A.) Mature hydranth. (B.) Oral eltremity of hydranth. (C.) Dissection showing intertentacular zone of medusa-buds.

Remaining figures are of medusa-buds and young medusae.

of the polypite is orange. L. Agassiz has shown that the entoderm of the stem is thrown
into longitudinal ridges which form partial septa projecting into the cavity of the stem. The
cavity, however, is continuous, and the septa do not fuse as in some other Hyboconidae.

This species is found upon the New England coast. According to Browne, 1896, it is
found upon the British, and also on the Irish coast, at Valencia, and off Iceland (Steen-

ANTHOMEDUS.E HYBOCODON. 41

strup) and Norway (Sars). It is apparently widely distributed over the North Atlantic,
along the shores of continents and islands. It is rarely taken far from some coast. We
can not be certain that the American and European forms are identical until the hydroids of
both are discovered.

Hargitt, 1904, states that the egg-cleavage is closely similar to that of Pennaria. Also
Browne, 1895, and Hargitt, 1902, 1904, find that the ova begin their development within
the walls of the manubrium of the medusa and are set free as actinulae. Muiler, 1908, and
Hargitt, 1904, find that the developing embryos within the walls of the manubrium absorb
their fellow ova, as has been observed by Doflein in Tuhularia mesembryanthcmurn, and by
Allen in T. crocea. Developing actinulae and budding medusae are abundant at Woods Hole,
Massachusetts, during the spring months.

Hargitt, 1902, and Perkins, 1904, find that actinula larvae develop upon the manubrium
at the same time that medusa-buds are being set free from the tentacle-bulbs. When set free
the actinulae have 10 tentacles. The mouth and oral zone of tentacles develop only after the
actinula is set free; and appear at the pole which was adjacent to the parent medusa during
the attached period. Linko, 1905, also observed this simultaneous process of development
of medusa-buds and of actinula larvae in his " Amphicodon graviJum," which develops
actinulae with 1 1 tentacles. It is probably identical with H . prolifer. A single specimen was
found in Barents Sea, north Russia.

H. Miiller, 1908, finds that the ova are large and amoeboid. Only about 2 eggs survive
to maturity in the ovary; the others having been devoured by the successful eggs. The
ooplasma is a network, the exoplasma being narrow-meshed and the endoplasma wider.
There are numerous pseudo-cells in degenerate stages, sometimes dividing amitotically.

Hybocodon pendula Haeckel.
Plate i, fig. 2.

Corymorpha nutans (hydroid), STIMPSON, 1853, Marine Invert. Grand Manan, p. 9.

Corymorpha pendula (hydroid), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 276, 343, plate 26, 6gs. 7-17. VERRILL,

1873, Invert. Anim., Vineyard Sound, pp. 510, 736, plate 36, fig. 273.
Corymorpha pendula, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 85, fig. 81. NUTTING, 1 901, Bull. U.S. Fish Commission,

vol. 19, pp. 337, 370, fig. 15. MAY, 1903, American Naturalist, vol. 37, p. 579, n figs, (histology and embryology).
Corymorpha pendula (medusa), AGASSIZ, A., 1865, North Amer. Acal., p. 192, fig. 324.
Hybocodon pendula, HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 34, plate 2, fig. 3.
Hybofodon pcndulus (medusa), HAECKEL, 1879, Syst. der Medusen, p. 34.
Monocautus pendulus, ALLMAN, 1871, Monograph Tubularian Hydroids, p. 397.

Adult medusa. Bell pyriform and about 5 mm. in height. It is relatively higher than
the bell of H. prolifer, and the gelatinous substance at the apex is much thicker. There
are 5 rows of nematocysts upon the exumbrella, as in H. prolifer. The basal bulb of the
well-developed tentacle is much smaller than in H. prolifer, and no medusa-buds have
been observed to arise from it. The well-developed tentacle is 2 to 3 times the length of the
bell-height. Its surface is studded with large, swollen rings of nematocysts, which give it
a heavy appearance. The tentacles at the bases of the 2 intermediate radial-canals are quite
well developed, and this is not the case in H. prolifer. The velum is wide and thin. The
radial-canals are narrow and straight. The manubrium is longer than in H. prolifer, and
extends a short distance beyond the velar opening. There is a small peduncle. The lips
are thickly covered with nematocysts. Pink pigment-granules are found in the entoderm
of the tentacle-bulbs. The entoderm of the manubrium is pink and lilac, and contains also
some pink pigment-granules.

Hydroid. The hydroid (Corymorpha[Hybocodori\ pendula)\s found in depths of IO to IOO
fathoms off the New England coast, with its base buried in the sand. It is 80 to 125 mm. in
height, and 6 mm. in diameter at the widest part. It always grows singly and is never branched.
The mid-region of the stem is very thick and is covered with a canaliculated coenosarc, but
the basal end narrows considerably, as does also the region near the free upper extremity,
which is long, slender, and pendulous. The stem is anchored by a number of root-like,
tubular, fleshy processes. The perisarc exists only as a thin delicate film. The head of the
polypite is large and highly contractile. There is a single verticil of long, hollow tentacles
at base of polypite. The mouth is situated at the extremity of a large flask-shaped proboscis,

42 MEDUSAE OF THE WORLD.

and is surrounded by a couple of rows of numerous irregularly arranged tentacles. These
oral tentacles are highly contractile, and are much smaller than those at the lower base of the
polypite. The medusae are borne upon branched stolon-like diverticula of the side walls of
the polypite, immediately above the zone of basal tentacles. Fully-developed medusae have
not been seen to be set free from the hydroid, but the similarity of the most advanced medusa-
buds observed to the free medusa found in the ocean leaves but little doubt concerning this
point. May, 1903, has studied the histology and embryology, and concludes that the medusa-
buds may at times become free, but usually mature while still attached to the hydranth.
This species has been found from Vineyard Sound to the mouth of the St. Lawrence River.
The medusa appears upon the southern New England coast in April and May, but is not seen
during the summer months, although the hydroid is abundant at this time.

It is possible that Euphysa tentaculata Linko, 1905, is identical with H. pendula (see
Zool. Anzeiger, Bd. 28, p. 214). Linko's medusa is from Barents Sea, north of Russia. I
have referred to this medusa in the description of Steenstrupia bigelowt.

Hybocodon forbesii Mayer.
Plate I, fig. 8; plate 2, fig. 3.

Hybocodon jorbesii, MAVF.R, 1894, Bull. Mus. Comp. Zool. at Harvard College, vol. 25, No. II, p. 236, plate I, fig. l; 1904,
Memoirs Nat. Sci. Museum Brooklyn Institute, vol. i, No. i, p. 8, plate 2, fig. 13.

Medusa. Bell asymmetrical, about 2.5 mm. in height and ellipsoidal in shape, being
slightly higher than broad. The gelatinous substance is of uniform thinness. There is a
single well-developed tentacle situated at the base of the longest radial-canal. A short, conical
tentacle is found at the base of the shortest radial-canal, and two smaller tentacle-bulbs are
situated one at the base of each of the intermediate canals. The well-developed tentacle is
about as long as the bell-diameter. Its base is small, and hardly greater in diameter than
the shaft of the tentacle. Its free extremity is fusiform, and covered with prominent nemato-
cyst-cells. No medusa-buds have ever been observed. The velum is narrow. The 4 radial-
canals are straight and slender and the circular canal is narrow. There are no rows or clusters
of nematocysts upon the exumbrella. Manubrium is spindle-shaped and swollen, and the
mouth is a simple, round opening situated at the extremity of a narrow tubular neck, which
extends beyond the velar opening. The entoderm of the terminal swelling of the large ten-
tacle is yellow streaked with red. Entoderm of manubrium yellow with red flecks.

This species is found in Nassau Harbor, New Providence Island, Bahamas, and at
Tortugas, Florida, in March to May. It is an abundant surface form. I have captured
many hundreds of specimens, but have never found them producing either medusa-buds or
actinula larvae. It is distinguished by its decided yellow and orange color, and the absence
of meridional lines of nettle-cells over the exumbrella.

"Hybocodon unicus."

Amphicodon unicus, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 276.

(?) Hybocodon chilensis (hydroid), HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 545, fig. W.

Bell 3 mm. high, 2 mm. wide, bell-shaped. I solitary tentacle between 2 rudimentary
basal bulbs; 3 perradially situated bulbs without tentacles. Medusa-buds ( ?) Manubrium
cylindrical, nearly as long as the umbrella cavity. Color ( ?) Asymmetry of bell ( ?) One
specimen was found by Vallentin, and briefly mentioned without figures by Browne, from
Stanley Harbor, Falkland Islands. Hybocodon chilensis Hartlaub, from the coast of Chile,
may prove to be the hydroid of this medusa ( ?) It will be impossible to identify the medusa
from the brief mention of it given by Browne, unless, indeed, it be rediscovered in Stanley
Harbor.

Hybocodon chilensis Hartlaub.

Hybocodon chilensis, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 545, fig. W.

(?) Amphicodon unicus (medusa), BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 276.

(?) Steenstrupia occidentalis (medusa), FEWKES, 1889, Bulletin Essex Institute, Salem, vol. 21, No. 7, p. 107, plate 3, fig. I.

Hybocodon occidenlalis, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 545.

As Hartlaub states, this hydroid may be the stock of Amphicodon (Hybocodon) unicus
Browne, from the Falkland Islands.

ANTHOMEDUS.E HYBOCODON. 43

The hydrocaulus is very thick and massive, about 50 mm. long and unbranched save for
the presence of" its roots and stolons. It is covered with a stiff layer of chitin, which is not
expanded at the base of the polypite. Polypite large, with 17 to 20 proximal tentacles, each
about 6 mm. long. There are also about 27 oral tentacles arranged in several rows. Above
the bases of the proximal tentacles there are 8 long, thick, medusa-bearing stolons, which
are thickly covered with numerous clusters of medusa-buds. Each medusa-bud has a single
very large tentacle. The stem of the hydroid is rusty yellow and the polypite light rose-color.
Found at Calbuco, Chile, South America.

This Hybocodon is closely related to, or possibly identical with, the form from Norway
described by Bonnevie under the name Hybocodon prolifer, but it differs from H. prolifer
Agassiz in having large, specialized, medusa-bearing stolons. Hartlaub proposes to call this
Norwegian hydroid Hybocodon christincc.

Fewkes, 1889 (Bull. Essex Inst., and also Amer. Naturalist, vol. 32, p. 597), gives a
brief description of a medusa from the coast of California which he calls by two names, Steen-
strupia occidentahs and S. californica; and which may be derived from Hybocodon chilensis.
This medusa is described as follows: Size (?) Bell ovoid without an apical prominence.
4 (?) 5 ( ?) rows of meridional lasso-cells extend upward from the 4 tentacle-bulbs, over the
exumbrella toward the bell-apex. I long tentacle and 3 rudimentary tentacle-bulbs at the
bases of the 4 radial-canals. The long tentacle is ringed at regular intervals and has a
large pigmented basal bulb from which there arise numerous medusa-buds. Color ( ?)
Velum well developed. 4 straight, narrow radial-canals. Manubrium shorter than the
depth of the bell-cavity. No axial canal. Coast of California, United States.

Hartlaub proposes to call this medusa Hybocodon occidentalis. I find nothing in Fewkes's
description to distinguish it from H. prolifer L. Agassiz, but apparently there is less difference
between the medusas of the various forms of Hybocodon than between their hydroids.

Hybocodon christinae Hartlaub.

Tubular ia prolifer^ Hybocodon prolifer, BONNEVIE, 1899, The Norwegian North Atlantic Eipedition, 1876-79,70!. z6, Hydroida,

p. z8, plate I, fig. 6.
Hybocodon chrislin,t, HARTLAUB, 1905, Zoolog. Jahrbuchern, Suppl. 6, p. 546; 1907, Nordisches Plankton, Nr. iz, p. loz, fig. 98.

The medusa attributed by Bonnevie to this hydroid has a single well-developed tentacle
with a basal cluster of large medusa-buds, each bud bearing a single tentacle. The medusa-
buds resemble H. prolifer, but the hydroid is distinguished by bearing its medusae upon 8
branched peduncles.

Hydrocaulus unramified, tubular, springing from a ramified hydrorhiza; occurrence
solitary; longitudinal striping, no collar; height about 50 mm. The hydranth has 14 proxi-
mal tentacles and 2 distinct circles of (oral) distal tentacles. The oral tentacles are shorter
and more numerous than the proximal. There are 8 blastostyles in a circle about midway
between the oral and basal tentacles, and these bear numerous medusae upon short pedicels.
The medusa-buds have 4 very wide radial-canals, and I highly developed tentacle which
exhibits at its swollen base the bud-rudiments of 4 new medusae even before the first has
become detached. This species is distinguished by its well-developed, branched, medusa-
bearing stolons. Found off Bodo, Norway. It is closely related to H. chilensis Hartlaub, of
the northern coast of Chile.

The medusa-buds in H. christinie appear to be confined to the under side and the sides
of the base of the well-developed tentacle.

Hartlaub, 1907, finds that the medusa becomes 4 mm. high and 3 mm. wide, with thin
bell-walls and an evenly rounded apex. The 4 radial-canals and ring-canal are band-like,
and wider than in other species of Hybocodon.

Hybocodon pulcher Hartlaub.

Auliscus pulcher, SAMUNDSSON, 1899, Vid. Meddcl. Nat. For. Kjobenhavn, Ser. 6, Aarg. i, p. 4Z5, taf. 4, 7 figs.

Hybocodon pulcher, HARTLAUB, 1905, Zool. Jahrbuchern, Suppl. 6, p. 545 ; 1907, Nordisches Plankton, Nr. iz, p. 96, figs. 9Z, 93.

Hydroid 40 to 50 mm. high with an oral circlet of 30 short tentacles, and another circlet
of 24 to 30 long tentacles around widest part of body of the hydranth. The only distinctive
characters of this species are the well-developed medusa-bearing stolons, the symmetrical

44

MEDUSAE OF THE WORLD.

bell, and 2 large principal tentacles of the budding medusa. In H. prolifer, on the other hand,
the medusae are usually set free with but one well-developed tentacle. The bell of H. pulcher
may become asymmetrical in later life ( ?) The hydrorhiza is thin and branching. There is a
flexible collar-like pensarc at the base of the hydranth. Medusas are developed upon stolons
from the hydranth above the circlet of long tentacles. When set free the medusa is 1.5 to 2 mm.
high, of symmetrical form. 5 longitudinal lines of nematocysts extend up the sides of the
bell to the apex. 3 small tentacle-bulbs. 2 equally well-developed tentacles arise side by
side, from the base of one of the radial-canals. Entoderm of manubrium and tentacles red.
Found off Iceland. I believe this form is probably identical with H. prolifer.

Genus MICROCAMPANA Fewkes, 1889.

Microcampana, FEWKES, 1889, Amer. Naturalist, vol. 32, p. 595; Bull. Essex Inst., Salem, vol. 21, No. 7, p. in.

The type-species is Microcampana conica Fewkes, from Santa Cruz Island, off the coast
of California.

GENERIC CHARACTERS.

Anthomedusae with 6 radial-canals, and 6 radially placed marginal tentacles. One of
these tentacles is well developed, and the other 5 are rudimentary.

It is possible that the vaguely described Rhabdoon singulare, of Keferstein und Ehlers,
(1861, Zoolog. Beitrage, p. 86, taf. 13, figs. 6, 7), from Messina, Mediterranean, is a form
of Microcampana, but there are apparently 12 longitudinal lines of nettling cells over the
exumbrella, and it is uncertain whether there are 4 or 6 radial-canals. The bell is 1.5 mm.
high and oval with uniformly thin walls. It is possibly an abnormal medusa of P elella.

Microcampana conica Fewkes.

Microcampana conica, FEWKES, 1889, Bull. Essex Inst., Salem, vol. 21, No. 7, p. in, plate 4,
fig. 8; American Naturalist, vol. 32, p. 595, fig.

Size ( ?) Bell conical with a well-developed, elongate, conical
apex; slightly asymmetrical. Exumbrella smooth, without meridional
rows of nematocysts. 6 marginal tentacles, 60 apart. 5 of these
are rudimentary, but the sixth is club-shaped, and about half as long
as the bell-height. 6 straight, narrow radial-canals and a ring-canal.
Manubrium conical to spindle-shaped, about as long as the depth of
the bell-cavity. There is a long, slender, straight, axial canal above
the stomach. Bell pink, tentacle-bulbs bright-red, manubrium yellow.
Found off Santa Cruz Island, California; under the cliffs of Punta
Diablo.

Genus DICODONIUM Haeckel, 1879, sens. ampl.

Dieodonium+Dinema s HAECKEL, 1879, Syst. der Medusen, pp. 27, 28.

A T orc Dinema, VAN BEN ED EN, 1867, Mem. Acad.Roy.des Sci.Belgique, 10111.36, art. 2, pp. 127, 130.

Dicodonium, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 443.

Sarsiella, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 66.

The type-species of this genus is DicoJonium cornutum Haeckel, 1879, of the Red Sea.
Dinema Van Beneden is a medusa which arises by budding from a Perigotiirnus-\\ke hydroid
and therefore belongs to the Tiannae. We use the term Dicodonium in the sense proposed
by Vanhoffen, 1891.

GENERIC CHARACTERS.

Codonidae with 2 well-developed and 2 rudimentary tentacles. No meridional lines of
nettle-cells upon the exumbrella. An apical projection to the bell and an axial canal pro-
jecting upward from the stomach, may or may not be present.

Some of the so-called "species" of Dicodonium are probably only abnormal specimens
of Sarsia with 2 tentacles instead of the normal 4.

FIG. II. Microcampana
after Fewkes in Amer-
ican Naturalist.

PLATE 3.

Fig. I. Dicodonium Jefferson:. Tortugas, Florida, June 15, 1897.

Fig. 2. Sarsia rnirabilis, young medusa. Nahant, Massachusetts, March

26, 1897.
Fig. 3. Hybocodon prolijer. t\ primary; t", secondary tentacle beginning to

develop; m, young medusa-buds. Agassiz Laboratory, Newport,

Rhode Island, September 17, 1895.
Fig. 4. Sarsia mirabilis, mature male. Nahant, Massachusetts, May 7,

1897.
Fig. 5. Hydroid of Sarsia mirabilis with ripe male medusa-bud attached to

the hydranth. Swallows Cave, Nahant, Massachusetts, May 8,

1897.

Drawn from life, by the author.

PLATE 3

ANTHOMKDUS.E DICODONIUM.

45

Tabular Description of the Medusa of DicoJonium.

D. cornutum
Haeckel.

D. dissonema
Haeckel.

D. ocellatum
= Dinema
ocellatum
Haeckel.

D. floridana
Mayer.

D. jeffersoni
Mayer.

D.adriaticum
Graeffe.

D. dinema =
Sarsiella
dinema
Hartlaub.

Shape and size

With bulging

Bell-shaped to

Bell-shaped

Cylindrical with

Dome-like

Bell-shaped

Half-egg-

of bell in

sides, and

egg-shaped

to hemi-

dome-like

0.75 high,

with a small,

shaped. 3

mm.

pointed

with blunt

spherical. 5

apex. 4

0.5 wide.

conical, api-

high, 2 wide.

apex. 4

conical

high, 4 to 5

high, 3 wide.

cal projec-

high, 4 wide.

a prx.

wide. Ex-

tion. 4 high,

umbrella

3.5 wide.

besprinkled

with tufts of

nettle-cells.

Length of 2

Curled up-

6 r lone,.

2 r lun^. v. irli

2 r long with a

r long. Basal

8 r long. A

2 r + long.

long tenta-

ward. 3 r

Verv large

fairly large

knob-like

bulbs bear

large car-

Basal bulb-

cles m

long. A

basal bulbs

tentacle-

swelling near

ocelli.

mine ocellus

small; with

terms of

row of bns-

and with an

bulbs, each

tip. Extreme

on each ten-

.'i .'Hi.

bell-radius

thngclusters

abaxial ocel-

with an ab-

tip ends in

tacle-bulb.

.

of nettle-

lus.

axial ocellus.

thin lash. No

cells on their

ocelli.

abaxial

sides. Tips

club-shaped.

Basal bulbs

small.

Condition of

Not present.

>

Mere bulbs.

Small, taper-

Mere basal

Men- basal

Absent.

2 rudimen-

ing, rudimen-

bulbs with

bulbs with

tary tenta-

tary.

abaxial

small car-

cle-bulbs.

ocelli.

mini- "1 rill.

Axial canal

Well devel-

Present.

Not present.

Not present.

A small axial

Not present.

above stom-

oped .

canal is

ach.

presrnt.

Shape of man-

Spindle-shap-

Spin. He-shap-

Club-shaped.

Flask-shaped,

Cylindrical,

Short, thick,

Spindle-

ubrium,

ed. 1.5 r

ed. 2 to 3

5 to 7 r

widest above,

with a nar-

and four-

shaped,

and length

long. Mouth

times as

long.

but near mid-

row tubular

sided.

6r long.

in terms of

a simple

wide in mid-

dle.

mouth-end.

bell-radius

round open-

dle as at

ing sur-

either end.

rounded bv

nettle-cells.

Gonail.

Ring-like, swol-

Large swollen

Gonad en-

Encircling

Large gonad

In stomach

Encircles en-

len, encir-

gonad en-

circles stom-

stomach.

encircles

wall.

tire stom-

cling middle

circles stom-

ach, thickest

stomach

ach.

third of

ach.

near mouth.

from base to

manubrium

near mouth.

Color.

Stomach, ten-

)

Bell-margin

Entoderm of

Entoderm of

p

Entoderm

tacles, and

rose-re, 1.

stomach and

stomach

brownish-

nettle-ring

Nematocyst

tentacles yel-

creamy

yellow.

of margin

clusters of

low. Knoh-

pink; of

dark purple-

exumbrella

like ends of

tentacle-

red.

black.

tentacles

bulbs deli-

Ocelli of ten-

flecked with

cate green

tacle-bulbs

red to orange.

or pink.

black.

Ocelli red.

Where found.

Red Sea, Gulf

Coast of Aus-

Trieste, Adria-

Tortugas, Flor-

Tortugas, Fla.

TruMr, Adria-

Normandy,

of Suez.

tralia.

tic Sea.

ida. On sur-

M.iv to

tic Sea.

coast of

face in June

June, com-

France.

to July, rare.

mon.

Mediterran-

ean ?

Remarks.

Development

Development

Development

Development

Development

1 ' 'pment

unknown.

unknown.

unknown.

unkri"u n.

unknown.

unknown.

Is this an

Occasionally

This species

abnormal

rudimentary

is distin-

Sarsia with

tentacles de-

guished by

only 2 ten-

velop so as to

stiff sensory

tacies ?

be almost as

hairs which

long as the

border ocelli

larger pair.

46 MEDUSAE OF THE WORLD.

Dicodonium cornutum Haeckel.

Dicodonium cornutum, HAECKEL, 1879, Syst. der Medusen, p. 27, taf. I, fig. 6.

Haeckel found this species at Tur, near Sinai, in the Red Sea. See tabular description of
medusae of Dicodonium. It has no ectodermal ocelli upon the bulbs of the 2 large tentacles.

Haeckel presents a beautiful figure of the medusa, drawn
from life.

Dicodonium dissonema Haeckel.
Dicodonium dissonema, HAECKEL, 1879, Syst. der Medusen, p. 27.

Haeckel describes this from a preserved specimen from
the coast of Australia. See tabular description of the
medusae of Dicodonium.

"Dicodonium ocellatum."

FIG. 12. Dicodonium cornutum, after Sarsia ocellata, BUSCH, 1851, Beobach. wirbellos. Seeth., p. 16, taf. 2, figs. 1-3.
Haeckel, 1879. Dinema ocellatum, HAECKEL, 1879, Syst. der Medusen, p. 29.

This medusa is described by Busch from Trieste, Adriatic, and it is probably an abnormal
Sarsia with only 2 of its 4 marginal tentacles developed. See tabular description of the medusae
of Dicodonium.

Dicodonium floridana Mayer.
Plate 2, fig. 5.

Bell about 4 mm. high and 3 mm. wide, with thin, uniform, vertical walls and a slight
apical projection. There are 2 equally-developed, diametrically opposed tentacles, each
about three-fourths as long as the bell-height. Near the outer end of each of these tentacles
there is a large, knob-like, swollen region, which terminates in a thin, tapering, nematocyst-
bearing lash. The knob-shaped swelling is hollow and its cavity is connected with the general
gastrovascular system of the medusa by means of a slender tube which extends through
the entodermal core of the tentacle. The basal bulbs of the tentacles are not large, and there
are no ocelli.

In addition to the 2 long tentacles, there are 2 small, tapering, rudimentary tentacle-
bulbs 90 from the large tentacles. The velum is well developed. There are 4 straight,
narrow radial-canals and a simple, narrow circular canal. The manubrium is flask-shaped,
being narrower at its base than at its middle point. The mouth projects slightly beyond the
velar opening, and is a simple, round opening at the extremity of a long, tapering neck. The
gonads are within the wall of the manubrium. The entoderm of the stomach is yellow, and
that of the distal bulbs of the tentacles yellow flecked with orange. The entoderm of the basal
bulbs of the tentacles is also tinged with yellow.

This medusa is occasionally found at Tortugas, Florida, in June. Occasionally a speci-
men is taken in which the normally rudimentary tentacles have developed so as to be nearly
as long as the pair of large tentacles, thus illustrating the imperfect line of separation between
Dicodonium and Sarsia.

Dicodonium jeffersoni Mayer.

Plate 2, fig. 4; plate 3, fig. I.
Dinema jeflersoni, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 30, plate 37, fig. 126.

The bell is dome-shaped, higher than a hemisphere, and about 0.75 mm. high. The
exumbrella surface is sparsely sprinkled with nematocysts. There are 2 short marginal
tentacles, and 2 rudimentary tentacle-bulbs. The tentacles are radially situated, and are
covered with numerous small wart-like clusters of nematocysts. There are 4 ectodermal
ocelli, I upon the outer side of each of the 4 tentacle-bulbs. The velum is well developed.
There are 4 straight, narrow radial-canals and a narrow circular vessel. The manubrium
is about as long as the depth of the bell-cavity. It is simple, cylindrical, and tube-like, and
the mouth is a round opening at the extremity of a short, cylindrical neck. A simple canal

ANTHOMEDUS^E DICODONIUM, SARSIA. 47

projects upward from the stomach into the gelatinous substance of the apex of the bell.
This is probably only the remnant ot the connection between the medusa and its hydroid
stock. The gonad is ring-like, and encircles the stomach, leaving the short proboscis free.

The entoderm of the tentacles and tentacle-bulbs is of a delicate green or pink. The
entoderm of the stomach is creamy pink. The ocelli are bright-red, and all other parts are
colorless.

This medusa is quite common at the Tortugas, Florida, in May and early June. Although
small, it appears to be mature, for sperm is often given off from the gonad of the males.

Dicodonium adriaticum Graeffe.
Dicodonium adriaticnm, GRAEFFE, 1884, Arbeit. Zool. Inst. \Vien, Bd. 5, p. 551.

Bell 4 mm. high, 3.5 mm. wide, bell-shaped, with a small, conical, apical projection.
2 long, radially placed tentacles, more than 12 mm. long, with large, thick, basal bulbs, each
with a large carmine ocellus upon the abaxial side of the bulb. 2 tentacle-bulbs at the bases
of the 2 radial-canals 90 away from the large tentacles. These basal bulbs have small red
ocelli. Each of these 4 ocelli are bordered by a ring of stiff sensory hairs. There are 4 small,
interradial tentacle-bulbs without ocelli. Thus there are 2 long tentacles, and 6 rudimentary
tentacle-bulbs. 4 radial-canals. Stomach short, thick, and 4-sided, with the gonads in the
stomach-wall. Mouth simple, with 4 lips. Found at Trieste. Adriatic Sea, in October.

Dicodonium dinema.
Sarsiclla dintma, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 67, fig. 63.

Bell oval, half-egg-shaped, 3 mm. high, 2 mm. wide. Exumbrella thickly besprinkled
with nematocysts. Only 2 tentacles, 1 80 apart. These are longer than the bell-diameter.
They have small basal bulbs, each with a reddish-brown ocellus. No trace of tentacle-bulb
or tentacles 90 apart from the well-developed tentacles. Manubrium about 2 times as long
as the bell-height. Spindle-shaped and encircled throughout by the gonad. Manubrium,
tentacles, and 4 radial-canals brownish-yellow. Found off the coast of Norway and in the
Mediterranean ( ?) Is this an abnormal young Sarsia with only 2 tentacles ? Hydroid
unknown. Medusa rare.

Genus SARSIA Lesson, 1843.

Sarsia, LESSON, 1843, Hist. Zooph. Acal., p. 333. AGASSIZ, L., 1849, Mem. Amer. Acad., New Scries, vol. 4, p. 224; 1862, Cont.
Nat. Hist. U. S., vol. 4, p. 211. WAGNER, 1885, Wirbellosen des Weissen Meeres, p. 76. VANHOFFEN, 1891, Zool.
Anzeiger, Bd. 14, p. 442. HARTLAUB, 1897, Hydromedusen Helgolands, p. 454; 1907, Nordisches Plankton, Nr. 12,
p. 7. VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 582. CHUN, 1895, Bibliothcca Zoologica,
Bd. i, Heft. 19, p. 4. GOETTE, 1904, Zool. Anzeiger, Jahrg. 27, p. 473.

Coryne + Syndictyon, AGASSIZ, A., 1865, North Amer. Acal., p. 175.

Coryne (hydroid), CALKINS, 1899, Proc. Boston Soc. Nat. Hist., vol. 28, p. 336.

Codonium + Sarsia + Svndictvon, HAECKEL, 1879, Syst. der Medusen, pp. 13, 16, 20.

Syncoryne, WEISMANN, 1883, Entsteh. Sexualzellen Hydromedusen, pp. 56, 216.

Syncoryne (hydroid), HARTLAUB, 1905, Zool. Jahrbuchern, Suppl. 6, p. 524. NUTTING, 1901, Proc. Washington Acad. Sci.,
vol. 3, p. 165.

Syncoryne (medusa), HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, pp. 29, 30.

Syncoryne, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 119 (citation of all references to 1850).

SynJiclon, AGASSIZ, A., 1862, in L.Agassiz's Cont. Nat. Hist. U. S., vol. 4, p. 340. MAYER, 1904, Mem. Nat. Sci. Museum Brook-
lyn Institute, vol. i, p. 7.

Stenyo, DUJARDIN, 1845, Annales Sci. Nat., ser. 3, tome 4, p. 257.

The type-species of this genus is Sarsia tubulosa of the northern coasts of Europe. This
medusa was first described by Lesson, 1843. The hydroid form was first described by
Gartner, 1774, in Pallas's Eleunch. Zooph., under the name of Coryne. Ehrenberg, Sars,
and Allman introduced the name Syncoryne. Staurtdia producta also gives rise to a medusa
which can not be distinguished from Sarsia.

GENERIC CHARACTERS.

Codonidae with 4 long, simple, equally developed tentacles, I at the foot of each radial-
canal. The manubrium is tubular and surrounded by a ring-like gonad. There is an
ectodermal ocellus upon the outer side of each tentacle-bulb. There are no meridional
nematocyst-tracts upon the exumbrella.

48 MEDUSA OF THE WORLD.

An apical projection of the bell may or may not be present; and there may or may not be
an axial canal extending upward from the stomach into this projection. The hydroid is
Syncoryne or Staundia.

We use the name Sarsia in the sense defined by Vanhoffen, 1891 (Zool. Anzeiger, p. 442).

In 1862, 1865, A. Agassiz described, under the generic name Syndictyon, a Sarsia having
reticulate nematocyst-cells upon its exumbrella, and clusters of such cells upon its tentacles.
These are, however, only characters of immaturity and largely disappear in the full-grown
medusa, which is a true Sarsia in all respects. In 1879, Haeckel formed the genus Codonium
to include medusae resembling Sarsia but distinguished by the possession of an apex upon
the bell into which a blindly-ending axial canal extends from the stomach. A bell-apex
and axial canal are characters which are acquired during growth in varying degrees by
almost all species of Sarsia, and are therefore not of generic value.

Two European and one American species of Sarsia produce medusae by asexual budding
from the tentacle-bulbs or from the walls of the manubrium. Chun, 1895 (Bibliotheca Zoo-
logica, Heft 19, fig. 2), showed that both ectoderm and entoderm of the manubrium take
part in the formation of these proliferating medusae; the entoderm of the manubrium forming
the entoderm of the daughter medusa, and the same being true of the ectoderm. Sarsia
prolifera Forbes, described and beautifully figured by Haeckel, 1879, under the name Codonium
codonophorum, produces medusa-buds upon its tentacle-bulbs.

The majority of Sarsia medusae are probably produced asexually by hydroids of the
genus Syncoryne, but at least one medusa identical with Sarsia is derived from the hydroid
called Staundia Dujardin, 1843. Such medusae may conveniently be placed in a subgenus
Staundwsarsia. A medusa which appears to be closely related to Sarsia is produced by the
remarkable parasitic hydroid Hydrichth vs.

The generic name Syncoryne was restricted by Allman, 1871-1872, to designate the
hydroid which produces the medusa Sarsia. The name Syncoryna was first proposed by
Ehrenberg, who applied it to hydroids now known as Clava, Coryne, etc., and in this old
sense it does not apply exclusively to the hydroid of Sarsia. By general consent, Allman's
name has been accepted in this restricted sense, Calkins, 1899, being almost alone in main-
taining that the generic name of the hydroid should be Coryne.

The commonly accepted arrangement is to retain the old name Coryne to include hydroids
in which the reproductive elements are produced in fixed sporosacs growing upon the
hydranth, while Syncoryne applies to like hydroids which, however, produce free medusae.

Weismann, 1883, found that the germ-cells of both sexes of Syncoryne sarsii originate
in the ectoderm of the budding medusa, and do not wander from their place of origin, but
become mature in the free medusa. Goette, 1904, finds, however, that in Sarsia the sperm
originates and remains in the peripheral ectoderm of the manubrium of the medusa, but the
p gg-cells, contrary to Weismann's contention, originate in the entoderm of the medusa-bud
while it is still attached to the hydroid, but afterwards they migrate into the ectoderm of
the manubrium, where they mature.

Many of the species of Sarsia display considerable individual variability, the colors of
the manubrium and tentacle-bulbs ranging from green or yellow to red. An apical pro-
jection and an axial vessel above the stomach may or may not be developed, and the length
of the manubrium at maturity is subject to much variability. Moreover, the hydroids may
form densely or sparingly branched colonies in accordance with environmental conditions,
and as is well known in S. mirabihs, free medusae are produced in early spring, whereas late
in the season the medusae mature while still attached to the hydroids. Much contusion has
been introduced into the synonymy of the genus, and different stages of the same medusa
have occasionally received different specific names.

Sarsia " nodosa" Busch, 1851, appears to be the young of some European Sarsia. The
exumbrella of very young medusae of Sarsia are usually besprinkled with nettling cells and
their tentacles bear prominent nematocysts, and Busch's medusa displays both of these
characters. (See Hartlaub, 1907, Nordisches Plankton, Nr. 12, p. 68, fig. 64.)

ANTIKIMKDrs.K SAKSIA.

49

Tubular Description of the Medusa of Sarsia.

S.tubulosa Lesson,

i8 43 *.

S. mirabilis L. Agassiz
49-t

S. eximia = Syncor-
yne eximia
Hincks, 1868.

S.radiata von
Lrndenfeld.

S. conica= Codon-
ium conicum

Haeckd.

Shape and size of

Egg-shaped. Half-

As in S. tubulosa, 7 to

Oval with quite

Semi-ovate. 3

Barrel-shaped with

bell in mm.

ellipsoidal. loto

10 high, 4 to 8 wide.

thick walls. 3

high, 2. 5 wide. long p<

12 high, 6 to 8

high. 2 wide.

apex. 12 high, 4

wide. With or

wide.

without apical

projection.

Character of 4 ten-

Basal bulbs small;

As in S. tubulosa.

Basal bulbs large,

As in S. eximia

7 r long, with small

tacles. Length

each with abaxial

with abaxial ocel-

basal bulbs.

in terms of boll-

ocellus. Tentacles

lus on each. Ten-

radius (r).

about 4 to 6 r

tacles about 4 r

long, and without

long-

prominent clusters

of nettle-cells.

Shape and length

Narrow, cylindrical

Narrow tubular base.

Cylindrical; wide

As in S. eximia,

Spindle-shaped,

of manubnum

at its base. Mid-

Mid-region swollen and

above, with nar-

i .5 r long or

only 1.5 to 2 r

in terms of bell-

dle part cylindri-

cylindrical. Line of

row, short, tubular

less.

long.

radius (r).

cal, swollen by

demarcation between

neck above mouth

gonad. Stomach

narrow, cylindrical,

Length about 2 r

small, spindle-

basal, and wide middle

or less.

shaped. Mouth at

part very sharp.

end of short coni-

Stomach swollen and

cal neck, as in

spindle-shaped and

S. mirabilis.

near outer end of tin-

Length 4 r.

manubrium. Beyond

stomach there is a

short, tapering, narrow

neck. Mouth is a

round opening. Total

length 3 to 6 r. Average

4 r.

Gonads.

Ring-like and de-

Thick ring encircling

Encircles manu-

As in S. eximia.

Encircles stomach,

veloped over

manubrium. Begin-

brium from base

leaving both ends

nearly the whole

ning abruptly at a short

to mouth end,

free.

length of the

distance from base,

leaving only short

manubnum, leav-

and extending to upper

neck above the

ing a short dis-

part of stomach, leav-

mouth free. No

tance at both ends

ing both ends of man-

medusa-buds.

free. No medusa-

ubnum free.

buds.

Color.

Entoderm of stom-

Entoderm of stomach

Entoderm of stom-

Entoderm of

?

ach green, bluish

usually green, some-

ach yellowibh-

manubrium

lilac, or red-

times green and red,

red to reddish-

and tentacle-

Tentacle-bulbs

or red. Tentacle-bulbs

brown or green.

bulbs deep

green, blue, or red.

green or red. Ocelli

Ocelli black in

brown.

Ocelli black.

black.

young and ml m

mature medusa.

TentacJe-bulbs

red, or brownish-

red.

\Vhere found.

North Atlantic

From coast of New Jrr-

Coast of England,

New South

Indian (Kc.m.

coasts of Europe

scv northward to Arctic

S( ntland, Helgo-

W.ilrs, Aus-

to Arctic Ocean.

Ocean. Coasts of Nor-

land, Nnnvav,

tralia.

wav. northern Russia,

Juneau, Alaska,

Greenland, Alaska to

North Pacific

northern Chile along

(Nutting).

Pacific coast to South

America.

Hydroid.

Syncoryne sarsia

Svnotrvne "miraluli "

Svncoryne eximia,

Sviu i>rvnr r.uli-

1'iikrmwn.

Loven. Hartlaub,

L. Agassi/,. S. reticu-

Allman.

aba.

1905, describes

lata is a closely related

(this hvdroid ?)

variety (see text).

from Terra del

Fuego, Antarctic.

*Description based on accounts by European writers.

j-Description based on original observations of medusa: found off southern coast of New England, United States.

50

MEDUS/E OF THE WORLD.

A number of Syncoryne hydroids have been described which probably produce Sarsia
medusae, but are not known so to do. Among these may be mentioned Syncoryne crassa
Pictet, 1893 (Revue Suisse Zool., tome I, p. 8), a small hydroid only 2 mm. high and rela-
tively thicker and shorter than S. pulchella of Allman, 1871 (Monog. Tubularian Hydroids,
p. 279, plate 6, fig. 3). Pictet's hydroid comes from Amboina, Malay Archipelago. It has
30 to 40 short, knobbed tentacles, and the medusa-buds arise singly between the tentacles.
The hydrorhiza is net-like, and the hydroid is pale orange in color.

Hartlaub, 1907, has made a masterly study of the genus Sarsia, and shows that the
species fall conveniently into two groups: the Eximia group, with short manubrium occupied
entirely by the gonad, which extends from its base to near the mouth, and the Tubulosa group,

Tabular Description of the Medusa of Sarsia. Continued.

S. prolifera Forbes
= S. codono-
phora Haeckel.
(See text.)

S. gemmifera Forb.
= S. siphono-
phora Haeckel.

S.hargitti = S.pro-
ducta Hargitt.

S. flammea Hart-
laub, 1907.

Sarsia (Stauridio-
sarsia)producta =
Stauridiaproducta
Wright, Hartlaub.

Shape and size of

Bell-shaped, with

Bell ellipsoidal.

Pyriform. 1.5 high,

Oval. 12 high, 7

Three-fourths-egg-

bell in mm.

or without apical

8 high, 6 wide.

i wide.

wide.

shaped. 10 high,

projection. 2.5

7 wide.

to 8 high, 3 to 8

wide.

Character of 4 ten-

Basal bulbs very

Well-developed

Well-developed

Tentacle tips knob-

Large basal bulbs

tacles. Length

large and wide,

basal bulbs with

basal bulbs with

like. Shafts cover-

with abaxial ocelli.

in terms of bell-

with black or red

abaxial ocelli.

abaxial ocelli.

ed with broken

Tentacles about 3

radius (r).

ocelli. Tentacles

Tentacles4r long.

Tentacles ir long.

(partial) rings.

to 4 r long.

hollow, tapering.

No ocelli.

2 to 6 r long;

with clusters of

medusa-buds

upon their bases.

Shape and length

Spindle-shaped with

Very long, tubular,

Spindle-shaped at

Conical spindle-

Cylindrical, about

of manubrium

narrow neck and

with spindle-

both ends, narrow-

shaped, only two-

3 r long.

in terms of bell-

simple, round

shaped stomach

er and tubular in

thirds as long as

radius (r).

mouth-opening.

near outer end.

middle. 2 to 3 r

the depth of the

About i .5 r long.

Mouth at extrem-

long. A circlet of

bell-cavity.

ity of narrow

medusiform

neck. Manubrium

gonads( = medusa-

8 r long.

buds ?) near prox-

imal end of man-

ubrium.

Gonads.

Encircling stomach.

Gonad ( ?)

Gonad ( ?)

Ring-like, encirc-

Gonad at base of

Medusae pro-

Medusae produc-

Medusiform buds

ling the manu-

stomach. No me-

duced asexually

ed asexually by

on stomach in cir-

brium from its

dusa-buds.

upon the tentacle-

budding from

clet near base.

base to near

bulbs.

sides of manu-

mouth.

brium. There may

be 10 to 22 or

more of these

budding medusae

upon manubrium

at one and same

time. Law of bud-

ding discussed in

text.

Color.

Entoderm of tenta-

Entoderm of stom-

Basal part of manu-

Entoderm fiery red

Stomach brownish,

cles and stomach

ach, tentacles, and

brium orange.

or orange.

gonads yellowish-

yellow to sage-

radial-canals or-

Distal end blue-

white, tentacle-

green. Mouth or-

ange-yellow to or-

green. Tentacle-

bulbs red. Ocelli

ange. Ocelli

ange-red. Ocelli

bulbs orange,

black.

brown. Tentacle-

black.

edged with green.

bulbs contain red

Ocelli black.

entodermal pig-

ment.

Where found.

English Channel to

Atlantic coasts of

No Man's Land,

Arctic Ocean.

Coast of Great

Mediterranean.

Europe, from

near Woods Hole,

Britain to Helgo-

Norway south-

Massachusetts,

land.

ward to Canary

United States. A

Islands.

single specimen

was found.

Hydroid.

Unknown.

Unknown.

Unknown.

Stauridia producta.

ANTHOMEDUS.E SARSIA.

51

with long, tubular manubrium with the gonad confined to a short length near its free outer
end. He states that the Eximia group are represented by such forms as Sarsia eximia,
brachygaster, flarnmea, barentst, prolijera, angulata, and apiculata. The Tubulosa group are
represented by S. tubulosa, Jensa, decipiens, litorea, pulchella, frutescens, mirabilts, reticulata,
princeps, rosaria, etc.

The genus Plotocnide Wagner, 1885, is defined by Hartlaub, 1907, as a Sarsia-Kks
medusa with nettle-cells upon the exumbrella. The gonad surrounds the manubrium from
the base downward. No ocelli. Hydroid unknown. It appears to me that unless it be
proven that the hydroid is different from Syncoryne, this genus should be merged with Sarsia.

Weismann, 1881 (Zool. Anzeiger, Bd. 4, p. 61), shows that the circulation of fluids
within the gastrovascular cavity of Coryne pusilla is aided by the rhythmical contraction of
the walls of the gonophore. The systole and diastole are not always of equal duration, but
each ranges from 60 to 75 seconds. Thus the circulation in certain hydroids may be aided
by periodic peristaltic contractions as well as by the movement of cilia.

Annandale, 1907, Journal and Proc. Asiatic Society of Bengal, vol. 3, finds Syncoryne
filarnentata, sp. nov., developing free medusae and growing in brackish pools of one-third
the salinity of sea water at Port Canning, Lower Bengal. The mature medusa is unknown.

Tabular Description of the Medusa of Sarsia. Continued.

S. rosaria= Coryne
rosaria A. Agassiz,
1865.

S. minima von
Lendenfeld.

S. brachygaster
Gronberg,
1898.

S. angulata
Mayer.

S. gracilis
Browne, 1902.

S. princeps
Haeckel, 1879.

Shape and

Oval, with small

Ovate. 3 high,

Three-fourths- Half-egg-shap-

Cylindrical,

Conical with thin

size of bell

apical projection.

2.5 wide.

egg-shaped, ed. 3 High,

thick-walled.

walls. 281040

in mm.

15 to 30 high, 10

widest above 2.8 wide. 4-

5 high, 3 wide.

high, 15 to 30

to 15 wide.

the middle. 15 sided in con-

wide.

to 18 high, 8 traction.

to 10 wide.

Character of

Basal bulbs large;

Tentacles about

Basal bulbs

Basal bulbs

Ocelli on basal

Basal bulbs

4 tentacles.

each with abaxial

2.5 r long.

large with a large, with well-

bulbs. Each

elongate, con-

Length in

ocellus. Tentacles

small abaxial developed ab-

tentacle 3 r

ical, and Banked

terms of

3 to 4 r long.

ocellus. Ten-

axial ocellus.

long, termina-

by pad-like clus-

bell-radius

Basal bulbs

tacles 3 to 4

Tentacles z r

ting in a ni-nia-

ters of nemato-

(r).

flanked by pads

rlong.

long-

tocyst-knob.

cysts. Each bulb

of nettle-cells.

bears small ab-

axial ocellus.

Tentacles 8 r

long.

Shape and

Spindle-shaped,

As in S. rosaria.

As in S. eiimia.

Shaped as in S.

Two-thirds as

Tubular, cylin-

length of

only 1.510 2 r long.

rosaria. r to

long as the

drical, with a

manubrium

one-twelfth r

depth of the

round mouth-

in terms of

long.

bell-cavity.

opening 4 r

bell-radius

long.

(r).

Gonads.

Encircles stomach,

As in S. rosaria.

As in S. eximia.

As in S. rosaria.

CO

As in S. rosaria.

leaving both ends

free. No medusa-

buds.

Color.

Entoderm of stom-

Entoderm

Entoderm of

Entoderm of

(0

Entoderm of ten-

ach ranges from

brown.

stomach and stomach and

tacle-bulbs and

yellow to pink or

tentacle-bulbs tentacle-bulbs

manubrium pur-

reddish-brown to

orange-red. robin-egg

ple. Ocelli

purple. Tentacle-

Ocelli black. blue; never

black.

bulbs yellowish-

red. Ocelli

brown to red.

deep-brown.

Where found.

Pacific coast of

Port Jackson,

Spitzbergen and Bahamas to

Stanlev Harbor,

Greenland and

North America,

New South

Greenland, in Tortugas,

Falkland Is-

Spitzbergen, in

southern Califor-

Wales, Aus-

summer. Florida. Win-

lands.

summer.

nia to Puget

tralia.

ter to mid-

Sound .

summer.

Hydroid.

Syncoryne rosaria

Syncoryne min-

Unknown.

Unknown.

(?

Unknown.

A. Agassiz and

ima.

Syncoryne sar-

Fewkes.

sii Hartlaub,

1905(0 1=

this medusa a

young Slab-

beria ?

52

MEDUS.E OF THE WORLD.

Sarsia tubulosa Lesson.

Syncorvna sarsii (hydroid), LOVEN, 1835, K. Vet. Acad. Handl. for Ar., p. 275, plate 8, figs. 7-10.

Oceania tubulosa (medusa), SARS, M., 1835, Beskriv og Jagtt., p. 25, plate 5, fig. 1 1 . Also: Syncorine sarsii, 1846, Fauna littor.
Noveg., part. 2, p. 2, tab. i, figs. 1-6.

Syncoryne sarsii, LOVEN, 1837, Archiv. fur Naturgesch., Jahrg. 3, p. 321, taf. 6, fig. 25. WEISMANN, 1883, Entsteh. Sexuabellen
bei Hydromedusen, Jena, pp. 56, 216. GARSTANG, 1894, Journal Oxford Club, vol. 2, No. 30, p. 7. CITRON, 1902,
Archiv. Naturges., Jahrg. 68, pp. I, 26, taf. i, 2 (sensory cells of tentacles). HINCKS, 1868, British Hydroid Zoophytes,
p. 52, plate 7, fig. 3. ALLMAN, 1872, Monog. Tubularian Hydroids, p. 275.

Syncoryne sarsi and Sarsia tubulosa, BEDOT, 1905, Revue Suisse de Zool., tome 13, pp. 120, 147 (all literature 1835-1850).

Sarsia tubulosa, LESSON, 1843, Hist. Zooph. Acalephes, p. 333. SCHI'LZE, 1873, Ueber den Bau von Syncoryne Sarsii, p. 14,
taf. 3. ROMANES, 1885, Jellyfish, Star-fish, and Sea Urchins, etc., International Scientific Series, vol. 49 (reactions to
stimuli). LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 212. HARTLALIB, 1907, Nordisches Plankton, Nr. 12, p. 19, figs. 10-16.
BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 9; 1895, Proc. and Trans. Liverpool Biol. Soc., vol. 9, p. 246.

Sarsia tubulosa, SARS, i$$$=Sarsia macrorhyncha, BUSCH, 1851; BROWNE, E. T., 1905, Proc. Royal Soc. Edinburgh, vol. 25,
p. 758.

Syncoryne gravata, HINCKS, 1868, Hist. British Hydroid Zooph., p. 53.

Sarsia tubuhsa+S. macrorhyncha, HAECKEL, 1879, Syst. der Medusen, pp. 16, 19.

Sarsia tubulosa + S. pulchella, FORBES, 1848, British Naked-eyed Medusa-, pp. 55, 57, plate 6, figs. ^, 3.

Corynt pusilla, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 340.

(!) Syncoryne Sarsii, HARTLAUB, 1905, Zoolog. Jahrbiichern, Suppl. 6, p. 525, fig. F (hydroid from Terra del Fuego).

Sarsia macrorhyncha, BUSCH, 1851, Beobach. wirbell. Seeth., p. lo, taf. 3, figs. 7-10; taf. 4, figs. I, 2.

FIG. 13. "Sarsia litorea," after Hartlaub, in Nordisches Plankton.

FIG. 14. "Sarsia decipiens," after Hartlaub, in Nordisches Plankton.

FIG. 15. "Syncoryne gravata," after Hincks, in British Hydroid Zoophytes probably hydroid of Sarsia

T^l- ' . _J ' r 1 _T^l__ TT 1' t_ . - _1 ' ^1_ - ! L..^ :^ J' _

mirabilis

This medusa is found off the English coast early in the spring, but it disappears before
August. It is evidently an Arctic species, is abundant off the northern coasts of Europe,
and is found off Iceland. It is very closely related to, if not identical with, the American
S. mirabilis, but may possibly be distinguished by its more slender and higher bell, its very
long manubrium, and its somewhat shorter tentacles. All of these characters are, however,
very variable in Sarsia mirabilis, and I have become convinced that the American and
European forms are at most only varieties, one of the other. For details see the tabular
description of medusae of Sarsia.

Romanes, 1885, carried out many interesting physiological experiments upon this
species, and showed that its ocelli are organs for the perception of light. The medusa is
sensitive only to rays between the red and violet, and is strongly attracted by the light. The
smallest part of the bell-margin is capable of initiating and maintaining the rhythm of the
bell, but if the margin be entirely removed all pulsations of the bell instantly cease, while
the cut-off margin continues to pulsate. Stimulation of the subumbrella of the bell causes
contraction of the manubrium (proboscis), and indeed the bell, when deprived of its margin,
still responds by contractions to all sorts of stimuli, chemical, thermal, electrical, or mechan-
ical, although sustained rhythmical pulsation is never resumed.

The hydroid is Syncoryne sarsii, which is common in shallow water along the coasts from
England to Norway. The polypites are spindle-shaped and elongate and have about 12 to

PLATE 4.

Fig. i. Sarsta mirabilts. Abnormal medusa with a tentacle arising from
the side of the manubrium. Nahant, Massachusetts, May 8,
1897. Metschnikoff, 1870, observed a similar abnormality in
Slabbena catenata.

Fig. 2. Median section of a young medusa of Sarsia mirabilis. The ocellus
on the right-hand tentacle is shown as if depigmented in order to
illustrate its structure. The eye on the left-hand tentacle-bulb is
shown normally pigmented.

Figs. 3 and 4. Sarsia mirabilis var. reticulata, young medusa. Figure 4,
one of the nematocyst-cells from the tentacles of figure 3. Agas-
siz Laboratory, Newport, Rhode Island, June 13, 1895.

Drawn from nature, by the author.

ANTHOMEDUS.E SARSIA. 53

16 short tentacles which arise at irregular intervals from their sides. Each of these tentacles
ends in a knob-like cluster of nematocysts. The 2 or 3 medusa-buds arise from the sides of
the polypite between the tentacles. The stems of the hydroid are quite smooth, sparingly
branched, and about 12 to 15 mm. high. The stems are translucent, slightly horny in color, and
the polypites are light-red.

I can not determine any well-defined distinctions between S. sarsii and S. mirabtlis,
excepting that in S. sarsn the medusa-buds appear always to arise from near the middle of
the sides of the polypite between the tentacles, whereas in S. mirabilts the medusa-buds arise
from near the base of the polypite at or below the level of the lowest zone of tentacles.

Hartlaub has given excellent figures and descriptions of a number of medusae which are
closely related to Sarsia tubiilosa, if not mere varieties of the latter. These are S. pattersoru
Haddon, S. frutescens Allman, 5. decipiens Hartlaub, S. litorca Hartlaub, and S. pulchella
Forbes. (See Hartlaub, 1907, Nordisches Plankton, Nr. 12, pp. 29, 30, 32, 36; figs. 20-226,
23, 24, 28, 29.) I hesitate to quote these as distinct species, for I have observed the same
or nearly the same variations among individuals in swarms of S. mirabilis at Nahant, Woods
Hole, and Newport on our coast. A statistical study, or better still, a study of the respective
hydroids, is required before we can hope to determine these so-called "species" with certainty.

Garstang, 1894, observes that the hydroid of S. tubulosa gives rise to dimorphic medusae,
as does S. mirabilis on the coast of New England, where early in the spring the medusae
are set free, whereas in May they mature while still attached to the hydroid.

Sarsia tubulosa Lesson, variety Sarsia mirabilis L. Agassiz.

Plate 3, figs. 2, 4, and 5; plate 4, figs. 1 and 2.
LITERATURE RELATING TO THE AMERICAN VARIETY or SARSIA TUBULOSA.

Sarsia mirabilis, AGASSIZ, L., 1849, Mem. Amer. Acad., New Ser., vol. 4, p. 224, plates 4, 5. STIMPSON, 1853, Marine Invert.
Grand Manan, p. u. HAECKEL, 1879, Syst. der Medusen, p. 17. FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard
Coll., vol. 8, p. 141, plate 3, figs. 1 1, 12. LEVINSEN, 1893, Vid. Meddel. Nat. For. Kjobenhavn (5), Bd. 4, p. 143. BIR-
ULA, 1896, Annuaire Musee Zool. Imp. Sci. St. Petersbourg, tome I, p. 332. HARTLAUB, 1907, Nordisches Plankton,
Nr. 12, p. 37, figs. 30-40. LINKO, 1900, Mem. Acad. Sci. St. Petersbourg, ser. 8, vol. 10, No. 4, p. u, taf. I, figs. 1-12
(structure of ocelli); 1905, Zool. Anzeiger, Bd. 28, p. 212; 1900, Traveaul Soc. Jmperiale des Nat. St. Petersbourg, torn.
29, p. 151.

Sarsia mirabilis (medusa), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 21 1-217; vol. 3, Ibid., plate 18, figs. 15-250.

Sarsia glafialis, MORCH, 1857, Beskriv af Gronland, p. 95.

Corync mirabilis (hydroid), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 185-21 1, plates 17-19; vol. 3, Jbid., figs. 1-16,
plate 17; figs. 1-14, plate 18; figs. 1-27, plate 19.

Corvnt mirabilis, AGASSIZ, A., 1865, North Amer. Acal., p. 175, figs. 282-285. CALKINS, 1899, Proc. Boston Soc. Nat. Hist.,
vol. 28, p. 336.

Oceania lubulosa, GOULD, 1841, Invert, of Mass., p. 348.

Syncorvnt mirabilis, ALLMAN, 1871, Monog. Tubul. Hydroids, p. 278. HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol.
24, p. 30, plate v, fig. i; 1901, Amer. Naturalist, vol. 35, p. 578, fig. 33; 1903, Science, vol. 16, p. 344 (variations).
HARTLAUB, 1905, Zool. Jahrbuchern, p. 526; Zool. Jahrb., Syst. Abth., 1901, Bd. 14, p. 356. TORREY, 1902, Univ.
California Publications, vol. I, p. 31. NUTTING, 1901, Bull. U. S. Fish Commission, vol. 19, pp. 328, 372, figs. 3, 81.

( ?) Syndictyon angulatum (young medusa), MURBACH and SHEARER, 1903, Proc. Zool. Soc. London, vol. 2, p. 168.

(})Sarsi barfnisii (young red-colored medusa), LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 214 (north of Russia).

( ?) Euph\sa tentaculata (abnormal medusa with only 3 well-developed tentacles), LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 214
(Barents Sea, North Russia).

Syncoryne densa, HARTLAUB, 1897, Wissen. Meeresuntersuch. Komm. Meere. Kiel, Helgoland, Neue Folge, Bd. 2, p. 452, taf.
166, figs. 4, u ; taf. i6r, figs. 7, 8 (hydroid forms dense tufts). Also: Sarsia sp., HARTLAUB, 1896, Verhandl. Deutsch.
Zool.Gesell., Leipzig, Bd.6, Vers., p. 182 (medusa; with branched manubria); 1907, Nordisches Plankton, Nr. 12, p. 26,
figs. 17-19, 22a (hydroid and medusa).

The following description is derived from a study of medusae and hydroids obtained
off the southern coast of New England, United States.

Adult medusa. Bell is half-egg-shaped, about 7 mm. in height and 4 mm. in diameter.
There is no apical projection, and the gelatinous substance is not very thick at the aboral
pole and becomes successively thinner near the margin. There are 4 long, highly contractile
tentacles, one at the base of each .radial-canal. The surface of each tentacle is covered with
prominent nematocyst-cells, which are clustered especially at the outer end of the tentacle.
Each tentacle arises from a well-developed basal bulb which contains a single ectodermal
ocellus upon the outer nerve-ring on the abaxial side of the tentacle-bulb. According to
Linko, 1900, the ocellus is composed of a cup-shaped invagmation of densely pigmented
ectodermal cells between which there are spindle-shaped bipolar nerve-cells. The ento-

54

MEDUSAE OF THE WORLD.

dermal core of the tentacle is hollow, and its lumen is continuous with the gastrovascular
system of the medusa. The ectodermal cells of the tentacle-bulb are very thick and are
probably nervous in function. The velum is well developed, being wide and thin. There
are 4 straight, slender radial-canals, and a simple, narrow, circular tube. A short, blindly-
ending tube extends upward from the base of the stomach into the gelatinous substance of the
bell. The entodermal cells of this small projection are several layers thick. The manubnum
is long and extends far beyond the velar opening. Its proximal part is slender and tubular,
but in its outer parts it is much distended by the genital products and consists of a long,
wide, cylindrical tube. The line of demarcation between the narrow and the wide part of
the manubrium is very sharp. The mouth is a simple opening at the extremity of a short
flask-shaped proboscis, and the lips are studded with nematocysts. The mature genital pro-
ducts are found in the ectoderm of the distal part of the manubrium. The entoderm of
the manubrium is usually green, but occasionally it is red. The entoderm of the tentacle-
bulbs is either red or green, and in some individuals the entoderm of the bulb is red while
the ectoderm is green.

FIG. 16. "Syncoryne pulchella."

FIG. 17. "Syncoryne frutescens."

The above figures are after Allman, in Ray Society, 1871-72.

Hydroid and young medusa. The hydroid stock is Syncoryne mirabilis. The stems
are attached by a creeping stolon. They are about 15 millimeters in height and branch
profusely. The main stems and also the side branches terminate each in a single polypite.
The stems are incased in an unannulated chitinous perisarc, which terminates sharply at
the bases of the polypites. Each polypite is fusiform and has about 12 to 18 tentacles which
arise in 3 or more indefinite whorls from the sides of the polypite. These tentacles are not
long, but are quite contractile. Each terminates in a knob-shaped cluster of nematocyst-
cells. The mouth of the polypite is a simple round opening situated at the extremity of a
conical proboscis. Medusa-buds are developed upon the sides of the polypites immediately
below the tentacles, near the lower base of the polypite. Each polypite bears I to 4
medusa-buds in various stages of development. In Massachusetts Bay the breeding season
begins early in March and lasts until the end of May. During March the medusa-buds

ANTHOMEDUS.E SARSIA.

55

develop 4 long tentacles, and are set free in an immature state, but during the last half of
the breeding season they fail to develop tentacles or give rise to mere short lashes upon their
basal bulbs; and they become sexually mature while attached to the hydroid, the manubrium
of each bud being greatly distended with the genital products. This observation was first
made by L. Agassiz, 1862 (pp. 189, 203), and has been confirmed by us in hydroid stocks
obtained in Swallow's Cave, Nahant, Massachusetts. Plate 3, fig. 5, is derived from one
of these sexually mature medusa-buds found upon a hydroid on May 8, 1897. It will be
observed that the manubrium of the bud is distended with sperm, while the tentacles are not
developed. In this connection it is interesting to observe that Pennaria and Podocoryne
carnca sometimes give rise to medusae which are sexually mature at their time of liberation,
while in other stocks of the same species the medusae are set free in an immature condition.
Garstang, 1894, observes the same phenomenon in the European S. sarsii.

This hydroid of our Sarsia is very abundant, from March until May, in Massachusetts
Bay, where it appears to grow equally well both in pure sea-water and in the brackish mouths
of rivers. The medusae appear in great numbers on the southern coast of New England

FIG. 18. Hydroid and medusa of "Sarsia Jensa," after Hartlaub, in Nordisches Plankton.

between February and April. They become rare during May, and are not seen during the
summer months. The hydroid extends northward to the Greenland coast, but has not been
recorded from Beaufort, North Carolina, or farther south. Linko and Birula, 1896, found
it in the White Sea, and Linko, 1905, records it from the eastern parts of Barents Sea between
Kanin and Kolgujew Islands. Calkins, Torrey, and Hartlaub have found this medusa
along the Pacific coast of America as far south as Chile. I believe Syncoryne densa Hartlaub
from Helgoland to be an environmental form of S mirabilis.

We have observed an abnormal medusa of Sarsia mirabilis in which a single well-
developed tentacle arose from the side of the manubrium at the point of juncture of the long
tubular basal region and the gemmiferous part of the manubrium. (See plate 4, fig. i). This
abnormal tentacle was studded with clusters of nematocyst-cells. It lacked a basal bulb
and had no ocellus. Asexual budding of medusa- from the walls of the manubrium is not
known in Sarsia mirabilis. Medusae of Sarsia with branched manubria are described by
Hartlaub, 1896, 1907.

Professor Hartlaub finds that in Sarsiti mirabilis the stomach is confined to the distal
end of the manubrium and the gonad is confined to the mid-region of the manubrium above
the stomach. Both the proximal and distal ends of the manubrium lack the gonad. On the
other hand, in S. brachygaster and S. cximia there is no differentiated stomach-region, and
the gonad may extend over the whole, or nearly the whole, length of the rnanubrium.

56

MEDUSAE OF THE WORLD.

The chief and possibly only well-marked point of difference between the forms S. mira-
bilis and S. tubulosa is that in the hydroid of S, mirabilis the medusa-buds arise from near
the base of the polypite, whereas in the hydroid of 5. tubulosa they arise from points higher
up on the sides of the polypite, between the tentacles.

20.

FIG. 19. Sarsia eximia, after Hartlaub, in Nordischcs Plankton.

FIG. 20. Syncoryne eximia, after Allman, in Ray Society, 1871-72. Hydroid'and young medusa.

PLATE 5.

Fig. I. Sarsia angulata. Nassau Harbor, New Providence Island, Bahamas,
July, 1903.

Fig. 2. Corynitis agassizii. Charleston Harbor, South Carolina, September,
1897.

Fig. 3. Ectopleura minerva. Tortugas, Florida.

Fig. 4. Ectopleura dumortieri, young medusa. Agassiz Laboratory, New-
port, Rhode Island, June, 1893.

Fig. 5. Ectopleura dumortieri, mature male. Agassiz Laboratory, Rhode
Island, July, 1896.

Fig. 6. Sarsia mirabilis var. reticulata. Nahant, Massachusetts, March 25,
1897.

Drawn from life, by the author.

PLATE 5

AXTHI i.MKDTS.E SARSIA. 57

Sarsia mirabilis var. reticulata.
Plate 4, figs. 3 and 4; plate 5, fig. 6.

Syndictyon reticulatum, A. AGASSIZ in L. AGASSIZ'S, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 340. AGASSIZ, A., 1865, North Amer.

Acal., p. 177, figs. 290-300.

Syncor\ne reticulata, ALLMAN, 1871, Monog. Tubul. Hydroids, p. 283.
Syndictvon reticulatum, HAECKF.L, 1879, Syst. der Medusen, p. 21.

( ?) Syndictyon raiculalum, MAAS, 1893, Ergeb. der Plankton-Expedition, Bd. 2, K. c., p. 67.
(tySarsia turrreula, MrCRADV, 1857, Gymn. Charleston Harbor, p. 36, plate 8, figs. 6-8.
S\ncor\ne reticulata, HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 30.
Sarsia reliculata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 45, figs. 41-43.
(?) Sarsia pulchella, AI.I.MAN, 1871, see Hartlaub, 1907, Nordisches Plankton, Nr. 12, p. 34, fig. 27 (abnormal twin medusa:,

fig. on p. 109). SPAGNOLINI, 1876, Catalogo Acalefi Mediterranco, p. 18, tav. 2, figs. i. 2.

Adult medusa. Bell ellipsoidal in shape, being about 4 mm. in height and 3.5 mm. in
diameter. No apical projection. Gelatinous substance quite thick at the aboral pole, but
thin at the bell-margin. There are 4 long, highly contractile tentacles, I at the base of each
radial-canal. Surfaces of these tentacles covered with prominent nematocyst capsules. Basal
bulbs of tentacles well developed and each one contains an ectodermal ocellus upon its outer
side. Velum wide and thin. There are 4 straight, narrow radial-canals, and a slender circular
vessel. Manubrium short and club-shaped, and does not extend far beyond the velar opening.
Mouth a simple, round opening. Genital products developed along the greater part of the
length of the manubrium. The entoderm of the manubrium and tentacle-bulbs is brick-red.

Hydroid and young medusa. Smaller than Coryne mirnliilis, being not more than 3 mm.
in height. Stems slender and hardly ever branch, excepting in old specimens, which some-
times give rise to a single branch near the base of the stem. Polypites large and club-shaped,
and having several whorls composed of 8 to 10 short tentacles. The medusae develop
among the tentacles near the proximal base of the polypite. When set free the young medusa
is remarkably large, being about 1.5 mm. in diameter. The bell is covered \\irh reticulated
clusters of nematocyst-cells (plate 4, figs. 3, 4) which are especially numerous near the bell-
margin above the circular canal. Some of these nematocyst-cells are large and round, while
others are narrow and long. The tentacles are thickly covered with helically arranged clusters
of nematocyst-cells. These cells (plate 4, fig. 4) are ellipsoidal in shape and are mounted
upon a short basal pedicel. Each cell gives rise to a long, sharp-pointed, sensitive hair. The
nematocyst thread lies coiled in a helix within the cavity of the cell. As the medusa becomes
mature the reticulated nematocysts disappear from the surface of the exumbrella and the
nematocysts upon the tentacles become less prominent. This medusa is found upon the Ne\\
England coast from April until June. It is distinguished from Sm-sm mirabilis only by its
nematocyst-covered tentacles and exumbrella and its small hydroid. It appears also to be
constantly brick-red, while 5. mirabilis is highly variable in color. It is often impossible
to distinguish mature medusae of S. reticulata from those of .V. mirabilis. It is possible that
the S. pulchella of Spagnolini, 1876, from Naples, Italy, is identical with .S'. r, n, uLita.

Sarsia eximia Boehm.

Coryne eximia, ALLMAN, 1859, Annals and Mag. Nat. Hist., ser. 3, vol. 4, p. 141; 1864, Ibid., vol. 13, p. 357.

Syncoryne eximia, HINCKS, 1868, Hist. British Hydroid Zooph., p. co, plate 9, fig. 2. ALLMAN, 1871, Monog. Tubul. Hydroids,

p. 282, plate 5.
Sarsia eximia, BOEHM, 1878, Jena. Zeitschrift fur Naturw., Bd. 12, p. 191, taf. 6, figs. 7-26; taf. 7, figs. 1-6. (In part), HAEC-

KEL, 1879, Syst. der Mcdusen, p. 17. HARTLAI P II, 1894, \Vissen. Meercsuntcrsuch. Komini^. Mo-rc. Kid, lldi;ii]and,

Ser. 2, Bd. i, p. 187.

Svncorvne eximia (hvdroid). NUTTINC;, 1901, Proc. Washington Acail. Sa., vol. 3, p. 166, plate 14, figs. 3, 4.
Sarsia eximia, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 21;, p. 756 (hydroid and medusa). BROCB, 1905, Bergcns

Museums Aarbog, No. 11, p. 4.

Sarsia tximia=S. bretonica, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 8, figs, i, 2a, 26 (full list of recent literature).
Syncorync eximia, BROWNE, 1907, Journal Marine Biol. Association, vol. 8, p. 37 (growth of the hydroid).

This form is found off the coasts of Great Britain, Helgoland, Shetland Islands, Norway,
and Juneau, Alaska (Nutting). For details of the medusa, see tabular description of the
medusas of Sarsia.

The hydroid is about 30 mm. high, forming a bush-like cluster of profusely branched
stems; the branches are short and simple and arise very irregularly from the main stems,
and are usually faintly ringed at their points of origin. The main stem is usually unringed,

58

MEDUSJE OF THE WORLD.

except at its base, and is quite smooth. The polypites are very elongate, spindle-shaped,
and have about 1 6 to 24 short, knobbed tentacles arranged in 4 to 6 somewhat irregular
verticils. The medusa-buds arise singly upon short peduncles near the bases of the tentacles
of the lower verticils. The entoderm is red to reddish-brown and the stems are yellow.

Browne found that confinement in an aquarium under somewhat unnatural conditions
caused the hydroid to grow rapidly and to form stolons, these being developed from branches

which touched the glass sides of the aquarium.
He also discovered that the medusa becomes
mature in from J to 10 days after being set free
from the hydroid. Later, in 1907, Browne found
that one of these hydroids placed in a glass tube
with a constant current of water passing through
it grew in length from 14 to 77 mm. in the course
of 9 days, and developed branches having a total
length of 500 mm. The hydroid was fed upon
copepods.

Sarsia radiata von Lendenfeld.

Sarsia radiata, VON LENDENFELD, 1884, Zool. Anzeiger, Jahrg. 7,
p. 584; 1884, Proc. Linnean Soc. New South Wales, vol.
9, pp. 583, 635; plate 20, figs. 31, 32; plate 30, figs. 1-4.

Medusa. Bell semiovate, slightly higher
than broad, 3 mm. high, 2.5 mm. wide. 4 ten-
tacles, each about 1.5 times as long as bell-height,
and with large bulbs about half as wide as the
manubrium. Ocelli (?) Velum wide. 4 straight
radial-canals. Manubrium cylindrical, half as
long as the bell-height. The gonad incases the
sides of the manubrium from the inner apex of
the bell-cavity to near the mouth. No medusa-
buds. Entoderm of manubrium and tentacle-
bulbs deep brown. Other parts colorless.

H \droid. The hydrocauli arise from a
creeping hydrorhiza which anastomoses in a
very open network. The perisarc terminates
with an oblique elliptical margin at the base
of each hydranth, and the hydranth is provided
with a muscle at this point which enables it to
bend downward and "shut up" as if it were the
blade of a penknife. The hydranths are spindle-
shaped, narrow, and elongate; and are, includ-
ing their hydrocauli, 3 to 5 mm. high. They have 6 to 8 verticils, each of 4 tentacles, situated
in 4 meridional lines, 90 apart. These tentacles are all knobbed at their ends. The hydranths
which produce medusae are shorter than the sterile polypites. The medusae bud out from the
lower half of the polypite between the tentacles. The entoderm is intensely brown in color,
and the perisarc is bright brownish-yellow. Other parts colorless. Found on the coast of
New South Wales, Australia. The medusae are produced in April and May.

Sarsia conica.
Codonium eonicum, HAECKEI., 1880, Syst. der Medusen, p. 634.

Bell barrel-shaped with conical apex one-third as long as the sides of bell. 12 mm. high,
4 mm. wide. 4 tentacles longer than bell-height, and with small oval basal bulbs. The
manubrium is half as long as the depth of the bell-cavity. The stomach is subspherical and
swollen by the encircling gonad. The mouth is at the end of a short, cylindrical throat-tube
which is free of gonads. Color ( ?) There is a long axial canal above the stomach. Indian
Ocean. Briefly described, without figures, by Haeckel.

FIG. 21. Sarsia eximia, from life, by the author. Mouse-
hole, Cornwall, England, Nov. 14, 1907.

A\THi>MKI>rs.K--SAi;-.I \. .V.I

Sarsia rosaria Haeckel.

Coryne rosaria, AGASSIZ, L., 1862, Cent. Xat. Hist. U. S., vol. 4, p. 340. A- \ i/. \.. i Sf>;, Virrh Arm-r. Aca!., p. 176, fig 289.

Sarsia rosaria, HAECKEL, 1879, Syst. Jer Mi .i'i CDj p. 18.

Santa rosaria= Syncoryne rosaria, FEWKES, iSSij. Am. r. Vituralist, vol. 23, p. 597, plate 25, fig. 7; tcit-figs. 8, 9 (hydroid?).

Syncoryne occidentals, FEWKKS, 1889, Bull. Essei Inst., Salem, vol. 21, No. 7, p. 99, plate 3, figs. 2, 3.

( ?) Syn diclyon angulatum, Mt RUAI H and SIIK^RV.R, 1903, Proc. Zoo]. Soc. London, vol. 2, p. 168.

Codonium apiculum, MURBACH and SHEARER, 190}, Proc. Zool. Soc. London, vol. 2, p. 165, plate 17, fig. i; plate 22, figs. 4, 5;

1902, Annals and Ma^. Nat. Hist., scr. 7, vol. 9, p. 72.
Sarsia apicula+S. rosaria, HARTLAI'B, 1907, Nordischcs Plankton, Nr. 12, pp. 17, 50, figs. 9, 45.

Bell 15 to 30 mm. high, 10 to 15 mm. wide, with fairly thick walls and small apical
projection. 4 equally developed, radially placed tentacles 1 .5 to 2 times as long as bell-height.
The basal bulbs of these tentacles are large and are flanked on either side by a large nemato-
cyst-pad. Each tentacle-bulb bears an abaxial ocellus. There are 4 slender, straight-edged
radial-canals and a narrow ring-canal. The velum is well developed. Manubrium short and
spindle-shaped, and mouth about at the level of the velar opening. There is a short axial
canal above the stomach. The gonad encircles the stomach, leaving both ends tree. N<>
medusa-buds. The colors are quite variable as in other species ot Sarsia. The tentacle-
bulbs range from yellow through red to brownish-red, and the stomach is yellow, pink, or
reddish-violet to purple.

This is the most abundant Sarsia along the Pacific coast of the United States. It occurs
in great swarms in San Francisco Harbor in spring; and in Victoria Harbor, Puget Sound,
in July.

The hydroid is Syncoryne rosaria found by A. Agassiz and Fewkes in shallow water
attached to piles of wharves. Each tentacle terminates in a knob, and the hydroid is a true
Syncoryne.

Sarsia minima von Lendenfeld.
Sarsia minima, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, pp. 584, 915, plate 21, figs. 34, 35.

Bell of medusa 3 mm. high and 2.5 mm. wide with "a long manubnum like the northern
Sarsia;." It is therefore readily distinguished from Sarsia radiata, which has a short manu-
brium. S. minima has a spindle-shaped, nearly cylindrical manubnum which extends tor
about half its length beyond the velar opening. The 4 marginal tentacles are somewhat
longer than the bell-height and are covered with rings of nematocysts. The entoderm ot
the stomach is pale brown, other parts colorless.

Hydroid. The stems are 2 to 3 mm. high and arise from a creeping, slightly branched,
non-anastomosing hydrorhiza. The perisarc which invests the hydrorhiza and hydrocauli is
irregularly annulated or wavy throughout, and terminates at the bases of the hydranths in
a transverse margin. The hydranths are slender, spindle-shaped, 0.6 to 0.8 mm. long, and
with 8 to 12 irregularly scattered tentacles, all of which are knobbed at their ends. When
they produce buds they become stouter, and are so thickly covered by the budding medusae
"that nothing of their bodies remains visible." The perisarc is reddish-brown.

This hydroid is found at Port Jackson, New South Wales, Australia, overgrowing Obclla
geniculata, on buoys and submerged ropes. The medusae are produced in April and May.
Von Lendenfeld did not obtain any mature medusae.

Sarsia brachygaster Grbnberg.

Sarsia brachygaster, GRONBERO, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 1 1, p. 459, taf. 27, figs. 3, 4. HARTLAI n, 1907, N'or-
disches Plankton, Nr. 12, p. n, fig. 3.

Bell 15 to 18 mm. high and 8 to 10 mm. wide and three-fourths-egg-shaped, the greatest
breadth being above the middle. 4 radially situated tentacles, each being about twice as
long as the bell-height. Basal bulbs of these tentacles well developed, and a single MI\
small ocellus upon the outer side of each bulb. Manubrium cylindrical, and two-thirds
as long as height of bell-cavity. Mouth situated at extremity of a shoit cylindrical neck.
The gonad is tubular and surrounds the stomach. No medusa-buds. The manubrium,
gonads, tentacle-bulbs, and tentacles are orange-red. The ocelli are black.

60

MEDUS.E OF THE WORLD.

Gronberg found this species at Spitzbergen in summer, and he also identified it among
a collection of medusae from Jakobshavn, Greenland, where it appears to be rarer than at
Spitzbergen.

Sarsia angulata Hartlaub.
Plate 5, fig. i; plate 6, fig. 3.

Syndictyon angulatum, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College., vol. 37, p. 5, figs. 6-8, plate 3; 1904,

Memoirs Nat. Sci. Brooklyn Institute Museum, vol. I, No. I, p. 7, plate I, fig. 6.
N on Syriilict\on tiHgtihitum, MURBACH and SHEARER, 1903, Proc. Zool. Soc. London, vol. 2, p. 168.
Sarsia angulata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 16.

Bell i mm. high; half-egg-shaped, with moderately thick walls. Becomes almost square
in cross-section when contracted. There are 4 slender tentacles with fairly thick spindle-
shaped ends. These tentacles are each about as long as bell-height and their distal halves are
tapering and are armed with nematocysts. The basal bulbs of the tentacles are not very large,
and each one bears an ocellus formed by a cup-like invagmation of ectodermal cells. The velum
is large, and the radial-canals and circular vessel are of fine caliber. The manubrium is
spindle-shaped with a narrow tubular o?sophagus and without an aboral projection. It is
about two-thirds as long as the height of the bell-cavity. The gonad encircles it, extending
from the base to near the mouth, leaving the throat-tube free. The entoderm of the tentacle-
bulbs and manubrium is robin-egg blue, while the ocelli are deep-brown, almost black. All

other parts are hyaline. This medusa is abundant
in the Tongue of the Ocean, Bahama Islands, in June
and July, and was found at Turks Island in January.
It is rare at Tortugas, Florida.

Sarsia gracilis Browne.

Sarsia gracilis, BROWNE, 1902, Annals Mag. Nat. Hist., ser. 7, vol. 9, p. 275.
(? ?) S\ncoryne sarsii (hydroid), HARTLAUB, 1905, Zoolog. Jahrbiichern,
Suppl. 6, p. 525, fig. F.

Bell 5 mm. high, 3 mm. wide; cylindrical, with
moderately thick walls and quadrangular margin.
4 tentacles, about as long as the bell-height, and end-
ing each in a large knob containing nematocysts. An
ocellus on the basal bulb of each tentacle. Manu-
brium about two-thirds as long as the depth of the
umbrella cavity. Color ( ?) Gonads ( ?) Found at
Stanley Harbor, Falkland Islands, by Vallentin, and
briefly described without figures by Browne. The
hydroid, Syncoryne sarsn, described by Hartlaub from
southern Terra del Fuego, may be the stock ot this
medusa.

The medusa may be a young Slabberia ( ?)

Sarsia princeps Haeckel.

Codonium princeps, HAECKEL, 1879, Syst.derMedusen,p.l3,taf.l,figs. 1,2.

Sarsia princtps, HAECKEL, 1879, Ibid., p. 655. LINKO, 1905, Zool. An-
zeiger, Bd. 28, p. 212. BROWNE, i9O3,Bergens Museums Aarbog,
No. 4, p. 8, plate i, fig. i; plate 3, fig. 4. HARTLAUB, 1907, Nor-
disches Plankton, Nr. 12, p. 47, fig. 44.

Codonium princeps, LEVINSEN, 1893, Vid. Meddel. Nat. For. Kjobenhavn
(5), Bd. 4, p. 143. GRONBERG, 1898, Zoolog. Jahrb., Abth. Syst.,
Bd. n, p. 458, taf. zj, figs, i, 2.

Bell thin-walled and conical, about 25 to 40 mm.
high, and 15 mm. wide. There is a short, conical,
apical projection. There are 4 tentacles with long
conical basal bulbs. The shafts of these tentacles

1-iG. 22. barsia princeps, alter Hartlaub, in

Nordisches Plankton. are very contractile, three or four times as long as

PLATE 6.

Figs, i and i'. Hydroid of Ectopleura Jumortieri. Dredged from a depth

of 5 fathoms in Newport Harbor, Rhode Island, August, 1896.

Figure I, enlarged view; figure i', natural size.
Fig. 2. Medusa of Ectopleura dumortteri. Agassiz Laboratory, Newport,

Rhode Island, July, 1892.

Fig. 3. Sarsia angulata. Turks Islands, Bahamas, January 2O, 1893.
Fig. 4. Protiara formosa, male. Nassau Harbor, Bahamas, June, 1903.
Fig. 5. Protiara formosa, female. Tortugas, Florida, June IO, 1897.
Fig. 6. Protiara formosa, young medusa. Turks Islands, Bahamas, January

20, 1893.
Fig. 7. Zanclea gemmosa. Manubrium of the medusa shown in figure 5,

plate 7.

Drawn from life, by the author.

PLATE 6

ANTHOMEDUS.fi SARSIA.

61

the bell-height, and covered with alternately arranged, wart-like clusters of nematocysts. A
small ocellus is situated on the outer surface of each tentacle-bulb near the bell-margin. There
are also 2 globular swellings, one on either side of each tentacle-bulb adjacent to bell-margin.
The velum is narrow. There are 4 narrow, ragged-edged radial-canals and a slender circular
canal. A short axial canal extends upward from the stomach-cavity into the gelatinous
substance of the apical projection of the bell. The manubrium is long and cylindrical, and
extends for about one-third of its length beyond the velar opening. The mouth is a simple
opening without prominent lips. A single, short, tubular gonad is developed upon the sides
of the manubrium. The manubrium, gonads, and tentacle-bulbs are purple. The ocelli
are black.

This species is found off the coasts of Greenland and Spitzbergen, where it appears to
be common. Gronberg, 1898, found it to be abundant at Spitzbergen, and Linko, 1905,
found it to be common in Barents Sea, north of Lapland, Russia. It is the largest known
Sarsia. Gronberg's description is based upon the study of living medusae. Hartlaub also
gives an excellent figure of the medusa, which we reproduce.

FIG. 23. Sarsia prolifera, from life, by the author. Mouse-
hole, Cornwall, England, Nov. 8, 1907.
FIG. 24. Sarsia codonophora, after Haeckcl, 1879.

Sarsia prolifera Forbes.

Sarsia proliftra, FORBES, 1848, British Naked-eyed Mi-ilus.r, p. 59, plate 7, fig. 3. BUSCH, 1851, Beobacht. wirbeilos. Seeth.,
p. i, taf. i, figs. 1-6. BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 147 (all literature to 1850). HARTLAUB, 1907,
Nordisches Plankton, Nr. 12, p. 15, figs. 7, 8.

Syncorync prolifera, ALLMAN, 1871, Monog. Tubularian Hydroids, p. 83, fig. 38.

Codonium codonophorum+ Sarsia prolifer, HAECKEL, 1879, Syst. der Medusen, pp. 14, 18, taf. I, fig. 3.

Sarsia codonophora, HAECKEL, 1880, Ibid., p. 655.

62 MEDUS.'E OF THE WORLD.

In 1848 Forbes described this medusa from Penzance Bay, southern England, where
it was abundant in August, 1846, and in 1879, and where I found it in November, 1907.
Haeckel describes that which may prove to be the same species from the Mediterranean. The
points of difference between the two medusae will be brought out in the following description.

The bell is bell-shaped; in Forbes's medusa (fig. 23) 3 mm. wide and 4 high; in Haeckel's,
(fig. 24), 8 mm. wide and 8 mm. high. Forbes's medusa lacks an apical projection, while
Haeckel's has a short, conical, pointed apex. Also, in Forbes's medusa there was no axial
canal above the stomach while in Haeckel's there was a narrow, axial vessel. Moreover,
in Haeckel's medusa the margin of the bell flares outward, while in Forbes's specimen this is
not the case.

It is well known to students of the Anthomedusae that apical projections and axial canals
are exceedingly variable, and may be absent or present in individuals of the same species.
The outward flaring of the bell may be brought about through contraction. Haeckel's medusa
appears to be merely a large, highly-colored specimen of S. prohfera Forbes.

The 4 marginal tentacles are I to 3 times as long as the bell-height. Their basal bulbs
are large and tapering and about as wide as they are long. The tentacles taper gradually
from base to shaft and are hollow. Clusters of I to 3 medusa-buds at a time are developed
upon the tentacle-bulbs immediately below the bell-margin. When set free these medusas
are already producing another asexual generation of medusas upon their tentacle-bulbs.
The 4 radial-canals and ring-canal are straight and very narrow. The manubnum is spindle-
shaped and nearly as long as the bell-height in Haeckel's, but not half so long in Forbes's
medusa. The mouth is at the end of a narrow neck and is a simple, round opening. The
stomach is encircled by the gonad, leaving the base and the throat-tube free.

Entoderm of tentacles and manubnum yellow to orange or sage-green. Each tentacle-
bulb bears a prominent ectodermal brown-colored ocellus.

If it be true that Forbes's and Haeckel's forms are identical the medusa must range from
southern England to the Mediterranean. It appears to be rare and its hydroid is unknown.

Allman, 1871 (p. 83, fig. 38). gives a figure of a medusa which closely resembles Forbes's
Sarsta prohjera.

I found numerous specimens of this medusa off Mousehole, Mounts Bay, Cornwall,
England, early in November, 1907. They accord well with the descriptions of Forbes
and of Busch. The entoderm of the manubrium and tentacle-bulbs was sage-green; the
tips of the tentacles light reddish-brown, and occasionally there was some reddish-brown
pigment in the central entoderm of the stomach. The gonad encircled the stomach, leaving both
ends of the manubrium free.

Sarsia gemmifera Forbes.

Sarsia gemmifera, FORBES, 1848, British Naked-eyed Medusae, p. 57, plate 7, fig. 2. CHUN, 1895, Bibliotheca Zoologica, Heft
19, pp. 4, 7, taf. I, figs. 5, 6 (law of development of medusa-buds). BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25,
p. 757. BROCH, 1905, Bergens Museums Aarbog, No. 1 1, p. 4.

Codonium gemrniferum+ Sarsia sipfionophora, HAECKEL, 1879, Syst. der Metlusen, pp. 15, 20, taf. I, fig. 4.

Codonium gcmmifcrum, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 132 (citation of literature to 1850).

Purena gemrtiifera, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 58, figs. 54-58.

This medusa is tound off the Atlantic coasts ot Europe from Norway southward.
Haeckel's Sarsia "siphonopliora" from the Canary Islands is probably another name for the
same species.

When young the medusa produces medusa-buds upon its manubrium. These arise in
a spiral line, one following the other, down the sides of the manubrium. The oldest of the
primary medusa-buds is nearest the base (uppermost) and the youngest nearest the mouth
of the manubrium.

These primary medusa-buds are attached to the sides of the manubnum by short pedicels,
and secondary and tertiary medusa-buds arise from the sides of these pedicels. Thus when
the oldest (uppermost) original medusa-bud is set free, the secondary bud upon its pedicel
remains attached to the manubrium and continues to develop, and when it in turn is set free
the tertiary bud completes its development; this process takes place with each and every
one of the series of buds.

ANTHOMEDUS.E SARSIA.

63

This successive freeing of the three series of buds produces various appearances at different
periods of time, and in this manner we may explain the condition observed by Haeckel, 1879,
in Sarsia " siphonop/iora," where the buds appeared in two series, the oldest and largest of
the upper series being at about the middle ot the manubnum, while the oldest ot the lower
series was near the base of the stomach. Forbes, 1848 ( plate 7, fig. 2 e), shows an intermediate
stage in the budding process, and an early stage is shown by Chun, 1895 (fig. 25 A). The
law of succession in development of the medusa-buds is clearly demonstrated by Chun.

Both ectoderm and entoderm of the
manubrium take an equal share in the
formation of the budding medusae.

For details of the character of the
medusa, see tabular description of the
medusae of Sarsia.

Haeckel failed to observe the sec-
ondary and tertiary medusa-buds upon
the pedicels of the budding medusae in
his Sarsia siplionophora, but this is
probably due to an oversight, tor in all
other respects his medusa appears to be
identical with S. gemmiftra P'orbes.

The gonad develops at the distal
end of the manubrium after the budding
process has ceased. Possibly there may
be 2 or more ring-like gonads ? (See
Hartlaub, 1907.) If this be the case
the medusa may be identical with
Dipuretia fertilis Metschnikoff, 1871.

Sarsia hargitti.

Corynt producta, HARGITT, 1902, American Natural-
ist, vol. 36, p. 550, fig. 3.

Syncorynf produtla, HAROITT, 1904, Bulletin Bureau
of Fisheries U. S., vol. 24, p. 30, plate I, fig. I.

A medusa called Sarsia producta is
produced by budding from the hydroid
of StauriJia producta Wright; and as
this name takes precedence over that
of Hargitt's medusa, it is necessary to
rename the American species.

Bell i .5 mm. high, and I mm. wide.
Apex dome-like and rounded, and sides
slightly compressed. Bell-cavity only
about half as deep as the height of the
bell. There are 4 long, equally devel-
oped tentacles with large basal bulbs,
each with a large, ectodermal ocellus.
Velum well developed. The 4 radial-
canals are narrow, straight, and smooth-
edged. The manubrium projects far
beyond the veJar opening, but is cap-
able of great contraction. Its terminal
part is bulb-like, while the basal portion is spindle-shaped and bears a whorl of medusi-
1'orin gonads. The stomach gives rise to a blunt, dome-shaped aboral projection. The
mouth is a simple, round opening. The basal part of the manubrium is orange, and its
free end blue-green. The tentacle-bulbs are orange, edged with delicate green. The ocelli
are black. A single specimen was found by Hargitt at Woods Hole, Massachusetts, on August

FIG. 25. Sariia gcmmifera, showing law of building. After Chun, in

Bibliotheca Zoologica, Heft 19, taf. I, 1895.
A, early; B, later stages of the process of budding.

64

MEDUSAE OF THE WORLD.

10, 1901. It is distinguished from all other American species of Sarsia by its medusiform
gonads borne upon the manubrium (text-fig. 26). It is not known whether medusae are set
free from the manubrium of the parent medusa.

Sarsia flammea Hartlaub.

Sarsia eximia (in part), HAECKEL, 1879, Syst. der Medusen, p. 17.

Sarsia fammea, LINKO, 1905, Zool. Anzeiger, Bd. 28, p. 212. HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 12, figs. 4-6.
(literature); 1903, Zool. Centralblatt, p. 22.

Bell high oval, 12 mm. high, 7 mm. wide, walls quite uniform and of moderate thickness.
4 tentacles, with well-developed, simple basal bulbs without ocelli. Tentacle tips slightly
enlarged, club-like. Nematocysts over distal halves of tentacles arranged in prominent, broken
rings. 4 slender, straight radial-canals. Manubrium thick, conical, spindle-shaped; only
two-thirds as long as the depth of the bell-cavity. No axial-canal above the stomach. Gonad
ring-like, encircling the manubrium from base to near the mouth. Stomach and tentacle-
bulbs light fiery-red or orange.

From the Arctic Ocean.

FIG. 26. Sarsia hargitii, after Hargitt, in Bull. TJ. S. Bureau of Fisheries.

FIG. 27. Sarsia flammea, after Hartlaub, in Nordisches Plankton. Showing arrangement of nemato-
cyst clusters on tentacles.

Separated from Sarsia eximia and S. brachygaster by its lack of ocelli, and from S.
brachygaster also by its smaller size. The best description is that of Hartlaub, 1907. Hydroid
unknown. It is possible that this may be the medusa of the parasitic hydroid called
Hydnchthys mirus Fewkes.

Genus SARSIA, Subgenus STAURIDIOSARSIA nov. subgen.
Stauridium, HARTLAUB, 1895, Zeit. fiir wissen. Zool., Bd. 61, p. 142; 1907, Nordisches Plankton, Nr. 12, p. 52.

CHARACTERS OF THE SUBGENUS.

Medusa similar to Sarsia, but the hydroid is Stauridia, not Syncoryne.

The name Stauridia should not be applied to these medusae, for it was first used by
Dujardin, 1843, to describe a hydroid which gives rise to a Cladonema medusa. It may be
well to apply the new generic name Stauridiosarsia to hydroids which resemble Stauridia,
but produce Sarsia-\\ke medusae.

ANTHOMKDUS.E SARSIA.

65

Sarsia (Stauridiosarsia) producta.

A^on Stauridir, DUJARDIN, 1843, Ann. Sci. Xaturelles, scr. 2, torn. 20, p. 370.

Stauridia producta, WRIGHT, 1858, Edinburgh New Philos. Journ., ser. 2, p. 283, plate 7, figs. 6-8.

Stauridium productum, HINCKS, 1868, British Hydroid Zoophytes, p. 68, plate 12, figs, i, I a. HARTLAUB, 1895, Zcit. fur

wissen. Zool., Bd. 61, p. 142, taf. 7, figs. 1-19; taf. 8, figs. 1-4; taf. 9, figs. I, 2, 6; 1907, Nordisches Plankton, Mr.

12, p. 53, figs. 47-50.

The medusa is a Sarsia, but the hydroid is similar to Stauridia, and differs from Syn-
corync, the hydroid of Sarsia, in that there is a basal circlet of simple, knobless tentacles;
whereas all of the tentacles of Syncoryne terminate in knobs. The tentacles of Stauridia

28.

FIG. 28. Sarsia producta, after Hartlaub, in Nnrdisches Planktn.

FIG. 29. Medusa; of "Sarsia producla," natural size; after Hartlaub, in \ordischt-s Plankton.

FIG. 30. "Stauridium productum," after Hincks, in British Hydroid Zoophyte-;. Hydroid and young medusa.

are of two sorts the simple, short, stiff, tapering, knobless basal circlet, and above them
several circlets of knobbed tentacles which arise from the sides ot the hydranth. In Syncoryne,
on the other hand, we find only knobbed tentacles. Thus two distinct genera of hydroids
produce one and the same genus of medusa. Moreover, Dujardin, 1843, discovered that the
hydroid which he called "StauriJie" produces the medusa Cladonema rnJititutu.

As the name Stauridia is preoccupied by Dujardin to designate the hydroid of Cladonema
it can not be applied to the medusa of Stauridia producta. I therefore call the medusa of
Stauridia producta by the name Sarsia producta, thus indicating its relationships when in
the reproductive stage.

The medusa Sarsia producta, which is produced by the hydroid Stauridia producta, may
be described as follows:

Bell 10 mm. high, 7 mm. wide, three-fourths-egg-shaped, with thick, gelatinous walls.
4 equally developed, radially placed tentacles; 1.5 times as long as the bell-height. Tentacle-

66

MEDUSA OF THE WORLD.

bulbs large, each with an abaxial, ectodermal ocellus. Manubrium cylindrical, with about
one-third of its length extending beyond the velar opening. A short, conical axial canal above
the stomach. Gonad extends from the base to the distal end of the manubnum. No medusa-
buds. Stomach brownish, gonads yellowish-white, tentacle-bulbs and axial canal red.
Helgoland and British coasts.

The hydroid has a thin-branching hydrorhiza, from which a number of club-shaped
hydranths arise singly. These hydranths are each about 2 mm. long, with a proximal circlet
of 4 to 6 short knobless tentacles, and above 2 or 3 more or less irregular circlets of large
knobbed tentacles, usually with 4 tentacles in each circlet. The medusa-buds are produced
from the sides of the hydranth above the basal circlet of tentacles. The hydranths are red.
The hydroid and medusa are described in detail by Hartlaub, 1895.

The English and Helgoland forms differ considerably and they may be regarded as
varieties one of the other (see Hartlaub, 1895, p. 157). The hydrorhiza of the English form
anastomoses, while that of the Helgoland hydroid branches sparingly and does not anasto-
mose. In the English form the
knobbed tentacles are in whorls of
4 each, whereas in the Helgoland
hydroid there areotten 5, occasionally
6, tentacles in each whorl. In the
English form the medusae are said
to arise from the bases of the ten-
tacles, while in the Helgoland form
they arise from the sides of the
polypite between the knobless basal
circlet and the lowest circlet of
knobbed tentacles. Finally the Hel-
goland medusa has a largeaxial canal
above the stomach, and this appears
to be absent in the English medusa.

Genus HYDRICHTHYS Fewkes, 1888.

H\drichthys, FEWKES, 1888, Bull. Mus. Comp.
Zool. at Harvard Coll., vol. 13, p. 224;
1888, Annals and Mag. Nat. Hist., ser. 6,
vol. i, p. 364. HARGITT, 1904, Bull. U. S.
Bureau of Fisheries, vol. 24, p. 32.

( ?) Plotocnide, HARTLAUB, 1907, Nordisches Plank-
ton, Nr. 12, p. 68.

GENERIC CHARACTERS.

The medusa resembles Sarsia,
but is without ocelli upon the ten-
tacle-bulbs. The mature medusa has
not been determined. The hydroid is
firmly attached to the side of a fish
(Sertola zonata), and is probably
parasitic, and degenerate in many
respects. All of our knowledge of
this remarkable form is derived from
Fewkes, 1888. It differs widely from
all other known forms of Tubulanan
hydroids.

In default of knowledge of the
mature medusa we must remain in
doubt concerning its true place in
our classification. It is possible that
Sarsia flamrnea Hartlaub may be the mature stage of this medusa.

\

FIG. 31. Young medusas of Hydrichthys rnirus, from Fewkes, in Bull.
Museum Comp. Zool. at Harvard College.

ANTHdMKDUS.E HYDRICHTHYS.

67

Hydrichthys mirus Fewkes.

Hydrichthys mirus, FEWKES, 1888, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 13, No. 7, pp. 124-232, plates 4, 5, 7 figs.;
1888, Ann. and Mag. Nat. Hist., ser. 6, vol. i, p. 364; 1888, Proc. Boston Soc. Nat. Hist., vol. 23, p. 391. HARGITT,
1904, Bulletin Bureau of Fisheries U. S., vol. 24, p. 32.

Medusa. The adult medusa is undetermined, but may possibly be Sarsia flammea. The
most advanced medusae reared by Fewkes had the following characters: Bell rounded,
without apical projection, and in form somewhat fuller than a hemisphere. Size ( ?) Outer
surface of bell besprinkled with nematocysts. Bell-walls of moderate thickness, becoming
thinner near the margin. There are 4 long, equally developed, radially situated tentacles,
with long, tapering basal bulbs which lack ocelli. The 4 radial-canals are broad and straight
and there is a simple, narrow, circular vessel. Manubrium cylindrical and about half as long
as the depth of the bell-cavity. Mouth surrounded by 4 simple lips ( :). The entoderm of
the manubrium is yellow and orange, while that of the tentacle-bulbs is orange. Other parts
of the medusa are colorless.

Fin. 32. Hydroid of ffyjriehthyi mirus, from Fewkes,
in Bull, \fuseurn Comp. Zool. at Harvard College.

Fish bearing hydroid, and a magnified part of hydroid
itself.

I'oinig medusa. When first detached from the hydroid the medusa has but two diametri-
cally opposite tentacles, and is more active in its movements than it is when it acquires four
tentacles. This may, however, have been due to the injurious effects of confinement.

Hydroid. The hydroid was found by Fewkes firmly attached to the side of a small fish,
Seriola zonata. The hydroid colony arises from a basal network of tubes which forms a
plate-like hydrorhiza embedded under the scales of the side of the fish, upon which it is probably
a parasite. This basal network gives rise to tubular gonosomes and also to filiform, flask-
shaped bodies, which recall the spiral zooids of Hydractinia, excepting that they appear to
have a terminal mouth-opening.

The medusa-bearing gonosomes are conical and gradually taper from base to summit,
where there may be a terminal opening. Irregularly distributed side'branches arise from the
main axis of the gonosome, and most of these give rise to medusa-buds. These side branches
are usually simple, although occasionally they branch. They are hollow and in communi-
cation with the entodermal cavity of the main stem.

Clusters of medusa-buds in various stages of development arise from the ends of the
branches. When set free these medusae have but 2 tentacles, but later 2 more tentacles develop.

68

MEDUSA OF THE WORLD.

All parts of the hydroid stock are highly contractile and the gonosomes and zooids are
without chitinous sheaths.

The hydroid is colorless, but the medusa-buds are reddish and orange in color. The
terminal parts of the flask-shaped zooids are also pigmented with reddish-orange.

A single stock of this hydroid was found by Fewkes in Narragansett Bay, Rhode Island,
attached to a fish, Seriola zonata, in August, 1887. The fish was swimming near the surface
and was apparently but little injured by the presence of the parasite.

It is interesting to observe that Alcock, 1892 (Annals and Mag. Nat. Hist., ser. 6, vol. 10,
p. 207, I fig.), has discovered a hydroid of the genus Stylactis which is commensal upon a
fish, Minous inermis. This hydroid, however, produces no medusae.

Genus EUCODONIUM Hartlaub.

Eucodonium, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 71.

The type species is Eucodonium brownei Hartlaub = Dipurena sp. Browne, from the
British Coast.

GENERIC CHARACTERS.

Codonidae with 4 equally developed, radially placed tentacles, each of which terminates
in a distal knob. Stomach mounted upon a gelatinous peduncle. Hydroid unknown.

This genus is distinguished from Sarsia and Slabberia by the gelatinous peduncle upon

which the stomach is placed. It is distinguished from
Dysmorphosa by the terminal knobs of its tentacles and
by its ring-like gonad.

Eucodonium brownei Hartlaub.

Dipurena sp., BROWNE, 1896, Proc. Zool. Soc. London, p. 473, plate 16, fig. 2.
Eucodonium brawnei, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 71, fig. 67.

Bell pyriform, thin-walled, I mm. wide. 4 radially
placed, slender tentacles with small basal bulbs. Ten-
tacles twice as long as the bell-height and with scattered
nematocysts along their lengths, and each with a large,
swollen, nematocyst-beanng knob at outer end of tentacle.
No ocelli. Velum wide. 4 straight, narrow radial-canals.
Stomach short, mounted upon a conical, gelatinous

from Europe, after Hincks in Brit- peduncle. Mouth a simple, round opening at about the

level of the velum. Medusa-buds arise from the sides of

the stomach. The tentacle-bulbs are blackish, stomach and terminal knobs of the ten-
tacles, dull dark-brown. Found at Plymouth, England, in September. Gonads( ?) This
is a young medusa, but we can not refer it to any known mature form.

Genus ECTOPLEURA L. Agassiz, 1862 (sens, amend.).

Tubularia dumortierl, VAN BENEDEN, 1844, Mem. Acad. Belgique, tome 17, p. 50, plate 2.

Eclopleura, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 343. AGASSIZ, A., 1865, North Amer. Acal., p. 191. HAZCKEL,
1879, Syst. der Medusen, p. 21. FEWKES, 1882, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 9, p. 295; 1883, Bull.
Mus. Comp. Zool. at Harvard Coll., vol. n, p. 85, plate I, fig. II. ALLMAN, 1871, Monog. Tubul. Hydroids, p. 423.
HINCKS, 1868, British Hydroid Zooph., vol. i, p. 123. VANHOFFEN, 1891, Zool. Anzeiger, p. 443. HARTLAUB, 1894,
Wissen. Meeresuntersuch. Komm. Deutsch. Meere Kiel, Abth. Helgoland (l), Bd.l, p. 188; 1907, Nordisches Plankton,
Nr. 12, p. 93. BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 748.

This genus was founded by L. Agassiz, 1862, the type species being E. dumortieri which
was first described by Van Beneden, 1844, from the coast of Belgium under the name Tubu-
laria dumortien.

GENERIC CHARACTERS.

Codonidae with 2 or 4 simple, unbranched tentacles situated at the bases of 2, or of all,
of the 4 radial-canals. 8 longitudinal rows of nematocysts extend from the 4 tentacle-bulbs
over the surface of the exumbrella to the apex of the bell. The manubnum is short and blunt,
and the mouth is a simple, round opening. The hydroid is Ectopleura, and is closely
related to Tubularia.

FIG. 33. Eucodonium brownei, after Browne,

in Proc. Zool. Soc. London.
FIG. 34. Hydroid of Ectopleura dumortien

ANTHOMEDUS.E ECTO PLEURA. 69

The discovery of Ectopleura pacifica and E. minerva makes it necessary to amend the
generic definition to include medusae with 2 as well as with 4 tentacles.
Ectopleura is distinguished from Zaticlea by its unbranched tentacles.

Ectopleura dumortieri L. Agassiz.

Plate 5, figs. 4, 5; plate 6, figs. I, i', 2.

LITERATURE RELATING TO THE EUROPEAN MEDUSA.

Tubularia dumortieri, VAN BENEDKN, 1844, Mem. Acad. Belgique, tome 17, p. 50, plate I. JOHNSTON, 1847, Hist. British

Zoophytes, p. 50, plate 7, figs, i, 2.

Ectopleura dumortieri, AC;ASSIZ, L. 1861, Cont. Nat. Hist. U. S., vol. 4, p. 342.
Eclofleura dumortieri, HINCKS, 1868, Hist. British Hydroid Zoophytes, p. 124.
Ectopleura dumortieri, ALLMAN, 1871-72, Monog. Tubularian Hydroid s, p. 424. BOHM, 1879, Jena. Zcitschrift fiir Naturw.,

Bd. 12, p. 198, taf. 7, figs. 10-13. HAECKEL, 1879, Syst. der Medusen, p. 22.
Non Ectopleura dumortieri, GRAEFFE, 1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 354.
Ectoflfura dumortieri, HARTLAUB, 1894, Wissen. Meeresuntersuch. Komm. Meere Kiel, Helgoland (i), Bd. I, p. 188. BROWNE,

1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 748.

Ectopkura dumortieri, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 94, figs. 90, 91.
Tubularia dumortieri, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 126 (citation of literature, 1844-48).

LITERATURE RELATING TO THE AMERICAN MEDUSA.

Ectopleura ochracea, A. AGASSIZ, in L. Agassiz's 1862, Cont. Nat. Hist. U. S., vol. 4, p. 343. AGASSIZ, A., 1865, North Amer.
Acal., p. 191, figs. 320-323. HAECKEL, 1879, Syst. der Medusen, p. 22. FEWKES, 1882, Bull. Mus. Comp. Zool. at Har-
vard College., vol. 9, p. 295, plate I, figs. 15, 16, 35, 36. NUTTING, 1901, Bull.U. S. Fish Commission, vol. 19, p. 373, fig.
82. HARGITT, 1904, Bulletin Bureau of Fisheries U.S., vol. 24^.32, plate 2, fig. I. AGASSIZ, A., and WOODWORTH, 1896,
Bull. Museum Comp. Zool. at Harvard College., vol. 30, p. 149, plate 6, fig. 6 (photograph of the medusa).

Ectopleura, sp., HARGITT, 1908, Biol. Bulletin, vol. 14, p. 106, figs. 8-1 1.

The finding of the hydroid of our American Ectopleura "ochracea" leads me to believe
that there are no specific differences between the European and American forms, and that
both should be called Ectopleura dumortieri, this being the older name. Hartlaub, 1907,
however, believes that E. dumortieri is smaller and has somewhat thicker bell-walls than
E. ochracea, but our American medusa varies considerably in these respects, as do also speci-
mens which I captured in the English channel in IQO".

The following description is based upon a study of large numbers of specimens obtained
at Newport, Rhode Island, United States.

Adult medusa. The bell is pyriform and is about 3 mm. high and 2.5 mm. wide. The
gelatinous substance is very thick and is especially so at the aboral pole. 8 longitudinal lines
of nematocyst-cells extend over the surface of the exumbrella from the basal bulbs of the
4 tentacles to the bell-apex, where they meet in a point. There are 4 equally developed,
short tentacles with large basal bulbs. The outer surface of these tentacles is thickly covered
with nematocyst-cells, and their distal ends are coiled in a close helix. Velum narrow. There
are 4 narrow, straight radial-canals and a simple, narrow, circular tube. The manubrium
is fusiform and extends not more than two-thirds the distance from the inner apex of the
bell-cavity to the level of the velar opening. There is no peduncle, and the mouth is a simple
round opening, the edge of which is armed with clusters of nematocysts. A short, blunt,
axial canal is often seen projecting upward from the stomach into the gelatinous substance
of the apex of the bell. The gonads are developed in the ectoderm encircling the stomach.
In brilliant specimens the tentacle-bulbs are yellow with a red core.

The middle region of the entoderm of the manubrium is lilac-colored, the upper part
delicate yellow, and the lower end red or pink, but very often the medusa is dull purple or
dull brownish-purple.

Hydroid. The hydroid shown in plate 6, figs. I, I'was found in August, in Newport
Harbor, Rhode Island, at a depth of about 10 fathoms. It is a Tubularian belonging to
the family Hyboconidas Allman, and is apparently identical with 'Tubularia (Ectopleura)
dumortieri of Van Beneden, 1844.

The hydroid is about 25 mm. in length, and was found clinging to the stems of Obelia.

The stems of the Ectopleura are isolated. Most of them are simple and unbranched, but

occasionally they give rise each to a single side branch near the lower end. The lower end of the

Ectopleura is coiled around the stem of the Obelia in a close helix. The stem of the Ectopleura

6

70 MEDUSAE OF THE WORLD.

is covered by a delicate investment of perisarc which displays a number of small annulations
near the upper end. The stem terminates in a single, large polypite, which is broad and
flask-shaped. The mouth is situated at the extremity of a tubular proboscis, and is sur-
rounded by a single verticil of about 16 short, flexible tentacles. In addition to these there is
another verticil of about 24 long, fleshy tentacles at the wide base of the polypite. These long
tentacles are not very contractile. They taper gradually from base to end. The tentacles
at the base of the polypite are about three times as long as are those surrounding the mouth.
The medusa-buds are borne in numbers on short, branching peduncles which arise from the
sides of the polypite in a zone immediately above the basal tentacles. When set free each
medusa has 4 short tentacles, and the 8 longitudinal rows of nematocysts upon the exumbrella
are well developed. The ectoderm of the stem of the hydroid is a delicate, fleshy-yellow and
the entodermal core is flesh-colored pink. The entoderm of the polypite is fleshy-pink and
yellow.

This medusa is very common throughout the summer in Narragansett and Buzzard's
Bays, and on the southern coast of New England east of New Haven, Connecticut. It has
not been taken north of Cape Cod. It has been recorded from Beaufort, North Carolina, and
I found it in Winyah Bay, South Carolina, but it has not been taken either at Charleston,
South Carolina, or at the Tortugas, Florida. It is found in the English Channel and off the
North Sea coasts of England, Scotland, Holland, and Germany. As in America, it appears to
be abundant only in a few localities. I found several specimens of this medusa in the English
Channel in October, 1907, and they appear to be identical in all respects with medusx from
the southern coast of New England, in America. Hargitt has recently discovered the hydroid
at Woods Hole, Massachusetts.

Ectopleura minerva Mayer.
Plate 5, fig. 3.

Ectopleura, sp., FEWKES, 1883, Bull. Mus. Comp. Zool. at Harvard College, vol. II, p. 85, plate I, fig. n.

Ectopleura minerva, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 38, p. 31, plate 16, fig. 38; plate 37, fig. 125.

( ?) Ectopleura dumortieri, GRAEFFE, 1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 354.

This medusa has 2 well-developed and 2 rudimentary tentacles, instead of 4 equally
developed tentacles as in the northern species of Ectopleura.

Bell 2.5 mm. high and pear-shaped, with a well-developed apical projection. Bell-walls
of moderate thickness. 8 rows of nematocyst-cells extend from the 4 tentacle-bulbs to the
apex of the bell. There are 2 well-developed tentacles and 2 small tentacle-bulbs. There
are 6 to 9 separate wart-like swellings upon the outer side of each of the 2 large tentacles.
These swellings are crowded with nematocysts. Basal bulbs of tentacles small and without
ocelli. There are 4 straight, narrow radial-canals and a slender, circular vessel. The velum
is well-developed. Manubrium pear-shaped and about two-thirds as long as the depth of the
bell-cavity. It is wider near the middle than at either end. The mouth is a simple, round
opening. A simple, short, conical style-canal extends upward into the apical projection of
the bell.

The entoderm of the manubnum and tentacles is a delicate purple, while the supporting
lamella of the bell is of a decided green. There are a large number of brilliant yellow spots
in the entoderm of the radial-canals and tentacle-bulbs.

This form is rare at Tortugas, Florida. It was found by Fewkes at the Bermudas. A
similar medusa was described by GraefFe, 1884, from Trieste, Mediterranean, under the name
Ectopleura dumortieri.

Ectopleura minerva may prove to be identical with E. pacifica Thornely, from the tropical
Pacific.

Ectopleura pacifica Thornely.

Eclopleura pacifca, THORNELY, 1900, Zool. Results, A. Willey, Part 4, p. 452, plate 44, figs, i, la, Cambridge, England.

Hydroid. Stems simple, unringed, diminishing in width toward the base, and 20 mm.
high, rising from a creeping stolon which connects the colony. Some of the stems give rise
to stolons near their bases. The polypite is abruptly marked off from the supporting stalk.

ANTHOMEDUS.E ECTOPLEURA, CORYNITIS. 71

There are two verticils of tentacles of 18 to 20 each. The basal tentacles are very long, and
the oral ones are short, and with knob-shaped ( ?) ends. The medusa-buds are borne in
clusters of from 7 to 9 on branched peduncles, which arise from the body of the polypite
between the two verticils of tentacles. When about to be set free each medusa-bud has 2
well-developed tentacles, thus resembling E. minerva of Tortugas, Florida. E. pacifica is
common in Blanche Bay, New Britain, South Pacific. It is attached to floats, fish-baskets,
etc. Color ( ?) This hydroid may be identical with E. minerva of the tropical Atlantic coast
of North America.

Genus CORYNITIS McCrady, 1857.

Corynitis, McCxADY, 1857, Gymn. Charleston Harbor, Proc. Elliott Soc. Charleston, p. 29. AGASSIZ, L., i86z, Cont. Nat.

Hist. U. S., vol. 4, p. 340. AGASSIZ, A., 1865, North Amer. Acal., p. 1 83 . ALIMAN, 1871, Monog. Tubularian Hydroids,

p. 286.

Coryneles, HAECKEL, 1879, Syst. der Medusen, p. 48.
PlolocniJc, WAGNER, 1885, Wirbelloscn Weissen Meeres, p. 74.
Corynttcs, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 442.

iV<m Corynitis, MURBACH, 1899, Quart. Journ. Mic. Sci., New Series, vol. 42, p. 354 (this is Zanclta}.
(')Modeeria, Protiara (young medusa), HAECKEL, 1879, Syst. der Medusen, p. 47.
(?) Tiaricodon, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 276.
Non Corynitis, NUTTING, 1901, Bull. U. S. Fish Commission for 1899, p. 372 (this is Zanclta).
Nan Corynitis, HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 42 (this is Zanclea).
Syncorync linvillei (hydroid), HARGITT, 1904, Biological Bulletin, Woods Hole, vol. 7, p. 251, i fig.; Ibid., 1908, vol. 14, p.

101, fig. I.

The type species of this genus is Corynitis agassizii described by McCrady from Charles-
ton Harbor, South Carolina.

GENERIC CHARACTERS.

Codonidae in which the manubnum is cruciform in cross-section. The 4 tentacle-bulbs
have large, abaxial, ectodermal ocelli. The tentacles are unbranched and terminate each in
a knob-like cluster of nematocysts, and the shaft of each tentacle is besprinkled with wart-
like clusters of nematocysts. There are 8 irregular, longitudinal rows of nematocysts upon the
exumbrella, 4 of these being radial and 4 interradial in position. The hydroid is S\ncor\n, .

McCrady, 1857, and L. Agassiz, 1862, described a hydroid which they ascribed to
Corynitis agassizii, but which is certainly very close to, if not identical with, the hydroid
generation of Zanclea. Hargitt, on the other hand, has discovered that Corynitis agassizii
arises from a Syncoryne-\\ke hydroid which he called S. lin-ctllt-i.

In the medusae of Ectopleura, Zanclea, and Corynitis there are longitudinal patches of
nematocysts upon the exumbrella. In Corynitis agassizii the tentacles terminate each in a
knob-like club. In Zanclea dichotoma they terminate in a similar manner, but in addition
they give rise to side branches all arising from the aboral side of the tentacles, and each termi-
nating in a knob-like cluster of nematocysts. In Corynitis, on the other hand, there are
merely scattered wart-like clusters of nematocysts over the shaft of each tentacle. It is possible
that Zanclea arose from some Coryrntis-\]ke medusa; certainly Zancleopsis dichotoma is inter-
mediate in many characters between Corynitis agassizii and Zanclea gemmosa.

However doubtful the relationship may be between Corynitis and the ClaJonemince,
there can be no doubt of its close relationship to Slabberia and Sarsia. The young medusa of
Corynitis can not be distinguished from the young of Slabberia.

Murbach and Nutting are mistaken in their Corynitis medusa, which is clearly Gcmmaria
(Zanclea). The medusa of Corynitis is so clearly described and figured by McCrady as to be
unmistakable. I have frequently taken it in Charleston Harbor, South Carolina, where
McCrady discovered it, and his figure and description enable one instantly to recognize it.
It lacks the feathered tentacles of Gemmarta (Zanclea).

Hargitt, 1904, at first followed Murbach in this erroneous identification of Corynitis,
but his final discovery of the Syncoryni--\\ke hydroid of Corynitis enabled him not only to
correct the mistake, but to establish the close relationship between Corynitis and other Sarsia-
like medusae.

72

MEDUSAE OF THE WORLD.

Corynitis agassizii McCrady.
Plate 5, fig. z.

Corynitis agassizii, McCRADY, 1857, Gymn. Charleston Harbor, p. 30, plate 9, figs. 3-8. AGASSIZ, L., i86z, Cont. Nat. Hist.

U. S., vol. 4, p. 340. AGASSIZ, A., 1865, North Amer. Acal., p. 183. ALLMAN, 1871, Monog. Tubul. Hydroids, p. 287.
Corynetts agassizii, HAECKEL, 1879, Syst. der Medusen, p. 49.

( ?) Corvnetcs agassizii, MAAS, 1893, Ergcb. der Plankton Exped., Bd. 2, K. c., p. 67.

Non Halocharis spiralis (hydroid), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 239, plate 20, figs. io-ioc.
Coryne agassizii (hydroid), McCRADY, 1857, Gymn. Charleston Harbor, pp. 29, 32.

Non Corvnitis agassizii, HARGITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 42 (this is Gemmaria=ZancIfa).
Non Corynitis agassizii, NUTTING, 1901, Bull. I'. S. Fish Commission for 1899, vol. 19, p. 372, fig. 79 (this is Gcmmaria=

Zanclca*).

Non Corynitis agassizii, MURBACH, 1899, Quart. Journ. Microscop. Sci., New Ser., vol. 42, p. 354 (this is Gemmaria=Zanclea).
Syncoryne linvillei (hydroid), HARGITT, 1904, Biological Bulletin Woods Hole, vol. 7, p. 251, I fig; 1908, Ibid., p. 101, fig. i.

Adult medusa. Bell miter-shaped and about 2.5 mm. in height. 8 longitudinal rows
of irregularly scattered nematocyst-cells are found upon the sides of the exumbrella immedi-
ately above the circular canal. 4 of these are situated above the tentacles and 4 are inter-
radial in position. The radially situated rows of nematocysts are longer than the interradial.
There are 4 straight, stiff tentacles, which are about three-quarters as long as the bell-diameter.
Each tentacle terminates in a large, knob-shaped swelling, the outer surface of which is thickly
covered with nematocysts. The entodermal core of this swelling is hollow and its lumen

is placed in communication with the gastrovascular system
of the medusa by means of a slender entodermal canal
which extends through the entire length of the tentacle.
Wart-like clusters of nematocysts are scattered over the
shaft of each tentacle. The basal bulbs are large and
hollow, and each one contains a single ectodermal ocellus
upon the outer side. Velum well-developed. There are 4
straight radial-canals and a narrow circular tube. The
large conical apex of the bell is hollow and in direct com-
munication with the stomach of the medusa. The manu-
brium is very large and swollen and fills the greater part of
the bell-cavity. It extends almost to the level of the velar
opening. The mouth is situated at the extremity of a short
oesophagus. It is without prominent lips. The gonads
occupy the greatly swollen sides of the stomach, and their
outer surfaces are smooth. 4 deep, interradial furrows
extend down the sides of the stomach, so that the manu-
bnum is cruciform in cross-section. The entoderm of the
FIG. 35. Hydroid of Corynitis, after Hargitt, basal bulbs of the tentacles is strawberry-red and that

iqo8, Biological Bulletin. ri 'iiiri I n

of the terminal knobs of the tentacles is orange-yellow

flecked with red-colored pigment-granules. The manubrium is dull yellow streaked with
strawberry-red and the entoderm of the radial-canals is faint yellow. The ocelli are dark-
brown or orange.

The hydroid is a Syncoryne, and is described by Hargitt. It grows in tufts which are
sparingly branched and 15 to 30 mm. high. The hydranths are vasiform with cone-shaped
proboscis. There are 15 to 30 tentacles distributed over the proximal third of the body of
the hydranth. These tentacles terminate each in a knob-like free end. The pensarc is plain
or with only a slight annulation. The hydrorhiza is reticulated in a loose network. The
medusa-buds are borne upon the body of the hydranth, usually in small clusters at the bases of
the tentacles, each cluster supported upon a single peduncle, the terminal specimen always
maturing first. The medusae, while still attached to the hydroid, have 2 well-developed, dia-
metrically opposed, club-shaped tentacles and 2 small tentacle-buds. There are no ocelli.
There are 4 radial-canals and the umbrella is bell-shaped. In formalin the hydroid is pale-
yellowish, hydranth somewhat brownish, and gonophores reddish-brown or pink. This
hydroid was found by Dr. Henry R. Linville growing upon rocks and piles under "Mill-
dam Bridge "west of Shelter Island, Long Island Sound, New York. It is described by
Hargitt from preserved material under the name Syncoryne linvillei. When set free the
medusae have only 2 tentacles and 2 small, basal bulbs.

ANTHOMEDUSjE CORYNITIS, SLABBEUIA. 7.1

This medusa is very abundant in Charleston Harbor, South Carolina, during the summer
and early autumn. It is also common in Great Peconic Bay, Long Island Sound, and at
Woods Hole, in September. It has not been taken north of Cape Cod, Massachusetts, or
at the Tortugas, Florida.

Corynitis arcuata Haeckel.
Coryneies arcuala, HAECKEL, 1879, Syst. der Medusen, p. 49.

This variety, or species, is distinguished from Corynitis agassizii by its long, narrow
bell, about twice as high as it is broad, and by its long, thin tentacles. The nematocysts upon
the tentacles are smaller and more numerous than they are upon the tentacles of C. agassizii.
The stomach is small and bell-shaped, with a short oesophagus which does not extend beyond
the velar opening. Haeckel gives the bell-height as 6 and the breadth 10 mm.; but judging
from his description of the medusa these figures must be erroneously stated and should be
reversed.

It is found off the coast of Brazil. Haeckel describes it from a single preserved specimen.

Corynitis (?) coerulea.
Tiaricodon cxruleus, BROWNE, 1902, Annals Mag. Nat. Hist., ser. 7, vol. 9, p. 276.

This medusa is very briefly described by Browne, without figures. The gonads are not
radially separated and there is nothing in Browne's description to indicate that the medusa
belongs to the Tiarinae. Its characters appear to be those of Corynitis.

Umbrella bell-shaped, 25 mm. high, 24 mm. wide, with a rounded summit. The exum-
brella is smooth without nematocysts. 4 fairly stout, radially situated tentacles tapering to a
point. Their basal bulbs are large and cylindrical, a little longer than broad, and with an
ocellus. Velum narrow. The manubrium is placed upon a short, broad peduncle. It is quad-
rangular with 4 perradial folded lips about as wide as the stomach. The mouth extends
nearly to the level of the velar opening. The gonads surround the stomach and extend outward
over the peduncle as 4 perradial lobes. Found at Stanley Harbor, Falkland Islands. The
young medusa resembles a Sarsia. The color is not stated by Browne.

Genus SLABBERIA Forbes, 1846.

Slabbcria, FORBES, 1846, Ann. and Mag. Nat. Hist., vol. 18, p. 286; 1848, British Naked-eyed Medusx, p. 53. FORBES and

GOODSIR, 1853, Tran. Roy. Soc. Edinburgh, vol. 20, p. 311.
Dipurrna, McCRADv, 1857, Proc. Elliott Society, Charleston, p. 135. AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

AGASSIZ, A., 1865, North Amer. Acal., p. 180.

Djpurtna + Bathycodon, HAECKEL, 1879, Syst. der Medusen, pp. 23, 26.
Dipurfna, BROWNE, 1898, Proc. Zool. Soc. London for 1897, p. 816.

Dipurena+Dipurella, HARGITT, 1904, Bulletin Bureau of Fisheries U. S., vol. 24, pp. 30, 31.
Dipurtna, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 442. BROWNE, 1905, Report on Pearl Oyster Fisheries, Gulf of

Manaar, Supplementary Report 27, Roy. Soc. London, p. 133.
S/abbcria (in part), AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.
Slabberia (in part), HAECKEL, 1879, Syst. der Medusen, p. 23.
Purena+ S/abbcria, H\RTLAUB, 1907, Nordisches Plankton, Nr. 12, pp. 55, 62.

In 1846 Forbes described a medusa under the name Slabl>,-ri<i hultfrata. It had 4
radial-canals with a swollen region upon each canal. In other respects it resembled Sarsiti,
excepting that each of the 4 tentacles terminated in a knob-like extremity. Forbes believed the
swelling on the radial-canals to be gonads and established the genus Slabbtria with the
following designation: "Umbrella campanulate; ovaries four, linear, in the course of the
four simple, gastrovascular canals; peduncle proboscidiform, highly extensile, oral orifice cir-
cular; a marginal tentacle springing from an ocellated bulb and terminating in a colored
globular body, placed opposite each of the gastrovascular canals."

Forbes laid especial stress upon the "gonads" upon the radial-canals, and he believed the
manubrium to be simple and tubular, and failed to find collar-like swellings upon it. Haeckel,
1879, p. 241, and Browne, i8g8, p. 816, find, however, that the gonads in Forbes' s medusa are
developed in separated ring-like annuli upon the manubrium and Browne, who sectioned the
supposed "gonads," finds that they contain entodermal cells crowded with small nuclei, and
they are doubtless mere glands similar to those seen upon the radial-canals of many Antho-

74

MEDUSA OF THE WORLD.

Tabular Description of the Medusa of Slabberia.

Slabberia hal-

S.strangulata =

S. strangulata

S.catenata Forbes,

S. ophiogaster=

S . pyramis= Bathy-

terata Forbes,

Dipurena

var. fragilis

1853= Dipurena

Dipurena

codon pyramis

I846. 1

strangulata

Mayer. 3

dolichogasta

ophiogasler

Haeckel, 1879.

McCrady,

Haeckel, 1879,

Haeckel, 1879.

i8 57 . 2

= D. picta Mayer,

= Sarsia stran-

1900= Sarsia

gulata Allman,

clavata Kefer-

1 87 1 . 5

stein, i862. 4

Shape and

Ellipsoidal.

Ellipsoidal to

Ellipsoidal.

Ellipsoidal.

Globular.

Pyramidal.

size of bell

8 high, 6 wide.

hemispherical.

4 high, 3.5

24 high, i. 6 wide.

5 high, 4 wide.

4-sided. 5 high,

in mm.

2 to 4 high,

wide.

5 wide. 4Hnes of

4103.3 wide.

nettle-cells ex-

tended up ex-

umbrella in 4

principal radii.

Character of

Large with an

As in S.halter-

Very small.

As in S. halterata.

Poorly developed,

Ocelli ?

the 4 ten-

abaxial ocellus.

ata.

with abaxial

tacle-bulbs

ocelli.

Length of

2 r

1.5 r, quite

2 r very slender.

2r

2r +

4 r +

each ten-

thick.

tacle in

terms of

bell-ra-

dius (r).

Number and

i large, terminal,

Single knob-like

As in S. strangu-

Proximal halves of

Tapering. Outer

Each tentacle ter-

position of

club-shaped

end and with

lata.

tentacles smooth.

parts thickly

minates in

nettle-

knob, i to 2

nettle-warts

Outer halves

covered with

"suctorial cup."

warts on

rings at base

on shaft of

ringed with 15 to

prominent par-

each ten-

of knob.

each tentacle.

3onematocyst

tial rings and

tacle.

swellings.

spirals contain-

ing nettle-cells.

Character

Narrow, straight,

Narrow, straight,

As in S. strangu-

As in S. strangulata.

As in S. strangu-

Edges of radial-

of the 4

each with swol-

without "glan-

lata.

lata.

canals lined

radial-

len(glandular?) i dular" swell-

with gland-cells.

canals.

enlargement at

ings.

two-thirds

length of canal

above margin.

Length of

4 to 71-.

4 to 7 r.

2 r

4 to 7 r.

8 to 12 T.

4 r+

manubrium

in terms of

bell-radius

Number of

2 or 3. I or 2

2. Small ring

i. As in S.

2 or 3 as in S. hal-

4 to 6 short,

8 separated rings-

ring-like

small genital

near middle,

strangulata.

terata. Preceding

swollen, sepa-

gonads

rings below

and longer

development of

rated by short

which en-

middle, and a

outer ring.

gonads medusa-

intervals.

circle

longer one

buds are borne on

manu-

near outer end

manubrium (see

brium.

of manubrium.

text).

Color.

Entoderm of ten-

Variable shades

Dull-yellow en-

Stomach yellow or

Asin S.catenata.

Stomach canals

tacle-bulbs

of yellow, red,

toderm of

red. Gonads and

Ocelli black.

and tentacles

and tentacle-

and green.

stomach and

tentacles light-red.

light-green.

knobs red-

Ocelli black.

tentacles.

Mouth, tentacle-

Mouth, tenta-

dish-brown;

Ocelli black.

bulbs, and apex of

cle-bulbs and

of manubrium

stomach bright-

terminal "an-

yellowish-

yellow, red, and

chors" darker

brown.

purple. Ocelli

green.

Ocelli black.

deep-purple brown.

Where found.

Southern coasts

Coast of United

Tortugas,

West coast of Scot-

Southern coasts of

Mediterranean,

of England

States from

Maldive Is-

land to Mediter-

England and

Corfu; from

and Ireland.

South Caro-

lands, Indian

ranean.

Ireland to Med-

depth of 20

Rare.

lina to Cape

Ocean, Bige-

iterranean. ( ?)

fathoms, April,

Cod.

low.

Cevlon (Browne).

1877, Haeckel.

'Distinguished by 4 small, isolated (glandular?) bodies upon its 4 radial-canals. Hartlaub, 1907, believes these to be gonads
which become mature only in old medusae.

2 Distinguished by its quite thick tentacles, each ending in a single, knob-like cluster of nematocysts.

3 Separated from S. strangulata only by its long, slender tentacles and uniform dull-yellow color.

4 Hydroid a Syncoryne. Medusa distinguished by numerous total or partial rings of nettling cells which are developed over

outer halves of tentacles. When young medusa-buds are borne on manubrium of free medusa.
5 Is this a fully developed, mature condition of 5. catenata(t).
6 Distinguished by terminal "suctorial cups" on its tentacles and numerous glands (?) which line radial-canals.

ANTHOMEDUS^E SLABBEIIIA. 75

medusae. On the other hand, Browne found the gonads upon the manubrium, arranged in
2 or more isolated ring-like swellings. In 1853 Forbes and Goodsir describe another species
of Slabberia, S. catenata, which lacks glands upon its radial-canals, although Forbes states that
they exist but are "scarcely to be traced."

In 1857 McCrady described a medusa under the name Dipurena strangulata, and in this
the 4 radial-canals are simple, but the manubrium is tubular and encircled by 2 separated,
ring-like gonads.

It is evident that Slabberia of Forbes is similar in all respects to Dipurena, excepting for
the problematical swellings on the radial-canals. It seems best to drop Dipurena and retain
the older name Slabberia. Swellings similar to those on the radial-canals of S. halterata are
seen on the radial-canals ot other Anthomedusae, such as Lymnorea alexandri and Dysmor-
phosa dubta. In L. alexandri the swellings are glandular and they are probably of similar
function in the Slabberia ot Forbes. Hartlaub (1907), however, states that these swellings are
gonads which develop only in old medusae. He does not state, however, that he sectioned the
medusa, and the matter appears to me to be still in doubt.

GENERIC CHARACTERS.

Codonidae in which the manubrium is tubular and is encircled by two or more separated
collar-like gonads. In other respects the medusa resembles Sarsia. The 4 radial-canals are
simple and without gonads. The hydroid is Syncoryne.

In all known species of Slabberia each of the 4 tentacles terminates in one or more knob-
like clusters of nematocysts.

Haeckel, 1879, p. 26, instituted the genus Bathycodon for a medusa resembling Dipurena,
but with short, blindly-ending diverticula upon its radial-canals. We define Slabberia as
equivalent to Dipurena+Bathycodon as defined in Haeckel's System.

Haeckel suppresses the generic name Slabberia on the ground that it was incorrectly
defined by Forbes, but if we are to retain only the correctly defined genera we must at once
drop nearly all of those defined previous to 1850, and even the majority of modern generic
names must vanish! As knowledge increases it appears to me far less confusing to amend and
retain old names rather than to drop them and substitute unfamiliar designations for familiar
objects.

Hartlaub, 1907, would distinguish Purena, with nettle-warts (not rings) upon the tentacles,
whereas Slabberia he would confine to include medusae having tentacles ringed with nema-
tocysts. The distinction between broken rings ("warts") and complete rings appears to me to
be too slight to be of generic value, but is an important specific distinction.

Slabberia halterata Forbes.

Slabberia halterata, FORBES, 1846, Annals, and Mag. Nat. Hist., vol. 18, p. 286; 1848, British Naked-eyed Medusae, p. 53, plate

6, fig. i. AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.
Dipurena halterata, HAECKEL, 1879, Syst. der Medusen, p. 24.
Dipurena halterata, BROWNE, 1896, Proc. Zool. Soc. London, p. 473; 1897, Ibid., p. 816; 1898, plate 49, figs. 2-iA. BEDOT,

1905, Revue Suisse de Zool., tome 13, p. 133 (citation of literature 1846-49).
Slabberia halterata, HART-HUB, 1907, Nordisches Plankton, Nr. 12, p. 64, figs. 61, 62.

Bell ellipsoidal, 8 mm. high, 6 mm. wide, with thick walls. Bell cavity about three-fourths
as deep as the height of the bell. 4 slender tentacles with well-developed basal bulbs, each with
an abaxial, ectodermal ocellus. Each tentacle terminates in a large, elongate, swollen tip with
one or two complete rings of nematocysts above it. 4 radial-canals straight and narrow, and
each exhibits I or 2 small (glandular?) swellings (gonads?) at about one-third the distance
between the inner apex of the bell-cavity and the margin. Hartlaub, 1907, believes these
swellings to be true gonads which become mature only in old medusae.

The stomach-cavity extends upward a short distance above the base of the manubrium
into the gelatinous substance of the apex of the bell.

The manubrium is 4 to 7 times as long as the radius of the bell. It is slender and tubular
and there is a small, spindle-shaped stomach near its distal end. The mouth is at the end of a
short, conical neck and is a simple, round opening.

76 MEDUSAE OF THE WORLD.

2 or 3 ring-like gonads girdle the manubrium. The two upper gonads are short, the
uppermost one being near the middle of the manubrium, and the next lower separated from it
by a short interval. The lowermost (distal) gonad is about twice as long as either of the others,
and it is adjacent to and above the stomach.

The entoderm of the basal bulbs and terminal clubs of the tentacles is reddish-brown and
of the manubrium yellowish-brown. The ocelli are black.

This medusa has been taken at infrequent intervals off the southern coasts of England and
Ireland. The best modern descriptions are given by Browne, 1898, and by Hartlaub, 1907.

It is distinguished by the 4 small swellings on the 4 radial-canals. According to Browne
these are probably glandular, but Hartlaub regards them as gonads which develop later than
the gonads upon the manubrium. Hartlaub, apparently, has not sectioned the medusa, while
Browne bases his conclusions upon a study of sections.

Slabberia strangulata Haeckel.
Plate 7, figs. I, 2, and 3.

Dipurtna strangulata, McCRADY, 1857, Gymn. Charleston Harbor, p. 33, plate 9, figs. I, 2.

Dipurena cervicata, McCRADY, 1857, Gymn. Charleston Harbor, p. 34.

Dipurena conica, A. AGASSIZ in L. AGASSIZ'S, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

Dipurcna cervicata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

Difurena strangulata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 341.

Difurena conica, AGASSIZ, A., 1865, North Amer. Acal., p. 181, figs. 301, 305.

Dipurena strangulata= Slabberia strangulata, HAECKEL, 1879, Syst. der Medusen, p. 23.

Dipurena conica, HAECKEL, 1879, Syst. der Medusen, p. 24.

Dipurena strangulata, FEWKES, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 155, plate 4, fig. 5. BROOKS, 1882, Studies Johns

Hopkins Univ. Biol. Lab., vol. 2, p. 136.

Dipurena conica, VERRILL, 1873, Report Commiss. Fish and Fisheries for 1871-72, p. 735.
Dipurella clavata (immature), HARGITT, 1904, Bulletin of the Bureau of Fisheries, vol. 24, p. 31, plate I, fig. 3.
Dipurena conica, NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 373. HARGITT, 1901, American Naturalist,

vol. 35, p. 578, fig. 34.
Purena strangulata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 55, fign. 51-53.

In 1857 McCrady described those which he believed to be two distinct species of Difurena,
D. strangulata and D. cervicata, from Charleston Harbor, South Carolina. In 1862-65,
A. Agassiz described that which he believed to be another species, from Buzzards' Bay, under
the name of D. conica. A careful study of Dipurena in Charleston Harbor, in September, 1897,
when the medusae were exceedingly abundant, has convinced me that all of these forms are one
and the same species. McCrady found only two individuals of D. strangulata and one of
D. cervicata, and thus A. Agassiz was misled into the assumption that the northern form was a
distinct species. The shape of the bell is, however, very inconstant, and the length of the
manubrium in the Charleston medusae is subject to a wide range of variability.

Adult medusa (plate 7, fig. i). The bell is quite variable in shape, for in some individ-
uals it is hemispherical, while in others it is ellipsoidal and higher than a hemisphere. All
gradations between these two extremes are observed. In a large individual in which the bell
was hemispherical it was 4 mm. in diameter; while another having an ellipsoidal bell was 4 mm.
in height and 3 mm. in width. There are 4 stiff tentacles, one at the base of each radial-
canal. Each tentacle terminates in a single, knob-like swelling which is covered with nema-
tocysts. The entodermal core of this swelling is hollow, and its lumen is placed in communi-
cation with the general gastrovascular system of the medusa by means of a narrow thread-like
canal which extends through the entire length of the tentacle. The basal bulb of each tentacle
is large and there is a single, ectodermal ocellus mounted upon a short pedicel, upon the outer
side of the bulb. The tentacles are all of equal length, and each one is about three-fourths as
long as the bell-diameter. The velum is wide. There are 4 straight, simple radial-canals
and a narrow, circular tube. The gastric cavity projects upward a short distance into the
gelatinous substance of the bell at the point of intersection of the 4 radial-canals. The manu-
brium is always long and increases in length with advancing age. In mature medusae it projects
far beyond the velar opening and varies in length from 5 to 10 mm. It is tubular and highly
contractile, and the mouth is a simple, round opening. Four distinct regions may be distin-
guished upon the manubrium of the mature medusa. First, a long, narrow, tubular region,
extending from the inner apex of the bell-cavity to about the level of the velar opening, then a
short, fusiform, swollen region, then a short, narrow, tubular region similar in size to the

PLATE 7.

Fig. I. Slabberia strangulata, mature male. Charleston Harbor, South

Carolina, September 8, 1897.
Fig. 2. Slabberia strangulata, half-grown medusa. Charleston Harbor,

South Carolina, September 1 1, 1897.
Fig. 3. Slabberia strangulata, young medusa. Charleston Harbor, South

Carolina, September 7, 1897.

Fig. 4. Slabberia strangulata var. fragilts. Tortugas, Florida.
Fig. 5. Zanclea gemmosa, young medusa. Agassiz Laboratory, Newport,

Rhode Island, September 17, 1892.

Drawn from life, by the author.

ANTHOMEDUS^E SLABBERIA. 77

proximal part of the manubrium, and finally at the distal end a long, club-shaped, cylindrical
region, at the extremity of which the mouth is situated. The 2 genital organs are found
within the 2 swollen regions of the manubrium. The color of these medusae is quite variable,
the entoderm of the manubrium and tentacles being either yellow, or yellow and red, or red,
or green and red. In the young medusas the entoderm of the manubrium is usually yellow, and
of the tentacles yellow dotted with red granulations; while in the mature medusa the color
of the entoderm is usually green or yellowish-green.

Young medusa (plate 7, fig. 3). In the youngest medusa observed, the bell is cylindrical
with vertical sides and a slight apical projection. The gelatinous substance is thin. There are
2 diametrically opposed tentacles each of which terminates in a single, large, nematocyst-
covered knob. The other 2 tentacles are as yet undeveloped and are represented by mere
basal bulbs. There are 4 ocelli, one upon each basal bulb. The manubrium is a simple, short,
conical tube and the mouth is a simple, round opening.

When the medusa is I mm. in height, a constriction appears near the proximal end of the
manubrium, which thus becomes divided into an upper and lower swollen region. As develop-
ment proceeds this constriction becomes more marked and finally the manubrium assumes
the adult shape; the upper and lower swellings of the young proboscis corresponding to the two
swollen regions of the manubrium of the adult medusa. Hargitt, 1904, describes the young
medusa under the name of "Dtpurella clavata."

This medusa is exceedingly abundant in August and September in Charleston Harbor,
South Carolina. It has been found by Brooks at Beaufort, North Carolina; and by A.
Agassiz and Verrill in Buzzards' Bay, Massachusetts. It has not been taken north of Cape
Cod, Massachusetts, and has not been observed at the Tortugas, Florida.

Slabberia strangulata var. fragilis.
Plate 7, fig- 4-

Difurena fragilis, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 29, plate 17, fig. 41.
(?) Difurena fragilis, BIGELOW, 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 251.

Bell 4 mm. high and halt-egg-shaped, with moderately thick walls, becoming thinner
near the margin. There are 4 long, slender tentacles, each about as long as the bell-height.
These terminate each in a single knob armed with nematocysts. The tentacle-bulbs upon
the bell-margin are not large, but each one bears a single, black ocellus in the ectoderm of its
outer side. Velum wide. There are 4 straight, narrow radial-canals and a slender, circular
vessel. Manubrium about twice as long as bell is high. It is slender and tubular, with two
separate, swollen regions containing the gonads. The entoderm of the manubrium and of
the basal bulbs of the tentacles is dull yellow, while the entoderm of the terminal knobs of
the tentacle is a faint orange.

This variety is found at Tortugas, Florida, in June. Bigelow, 1904, found a colorless
Dipurcna at Suvadiva Atoll, Maldive Islands, Indian Ocean, which he believes is probably
identical with Dipurena fragilis.

S. fragilis may be distinguished from other American Dipurenae by its \ong,very slender
tentacles, and the dull-yellow color of its entoderm.

Slabberia catenata Forbes and Goodsir.

Plate 8, figs. 8 and 9.

Slabberia catenata, FORBES and GOODSIR, 1853, Trans. Roy. Soc. Edinburgh, vol. 20, p. 311, plate 10, figs. ya-^e.

Difurena dolichogaster, HAECKEL, 1864, Jena. Zeltschrift fur Naturw., Bd. i, p. 337; 1879, Syst. der Medusen, p. 25, taf. 2,

figs. 1-7.

Difurena calenata, HAECKEL, 1880, Syst. der Medusen, p. 655.

Difurena fertilis, METSCHNIKOFF, E. and L., 1870, Verhandl. Gesell. Freunde Nat. Moskau, tome 8, p. 343, taf. 3, figs. 1-5.
Sarsia dolichogaster, SPAGNOLINI, 1876, Catalogo Acalefi Mediterraneo, p. 18, tav. 2, fig. 3 (from Naples, Italy).
Difurena dolichogaster, CHUN, 1895, Bibliotheca Zoologica, Heft 19, p. 6, taf. i, figs. 1-4. *
Difurena catenata, MONTICELLI, 1897, Atti Accad. Sci. Torino, vol. 32, p. 888, I fig. (abnormal specimen from the Gulf of

Cagliari, with a manubrium which gives off 2 side branches, each with mouth and gonad).
( ?) Sarsia clavata, KEFERSTEIN, 1862, Zeitschrift fur wissen. Zool., Bd. 12, p. 27, taf. 2, figs, i, 2.
( ?) Sarsia, sp., ALLMAN, 1871, Monog. Tubul. Hydroids, p. 83, fig. 37.
(?) Sarsia clavata, HAECKEL, 1879, Syst. der Medusen, p. 19.

( ?) Sarsia clavata (Syncoryne clavata), GRAEFFE, 1884, Arbeit. Zool. Inst. \Vien, Bd. 5, p. 351 (first description of the hydroid).
Difurena ficta, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 29, plate 18, figs. 45, 46.
(? Sarsia clavala)^ Slabberia catenata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, pp. 51, 63, figs. 46, 59.

78 MEDUSAE OF THE WORLD.

DESCRIPTION OF AMERICAN SPECIMENS.

Bell high with straight, vertical sides and flatly-rounded top, about 3 mm. high and 1.5
mm. wide. Bell-walls very thick and of a rigid, gelatinous consistency. Bell-cavity about
two-thirds as deep as the total height of the bell itself. There are 4 radially situated, slender
tentacles, all of the same length and not quite as long as the bell-height. Each ot these ten-
tacles terminates in 3 to 5 bulb-shaped, ring-like swellings containing nematocysts. The
basal bulbs of the tentacles are large and swollen, and each of them bears a dark-purple,
ectodermal ocellus upon its outer side. There are 4 straight, narrow radial-canals and a
simple, slender, circular vessel. The velum is quite narrow. The manubrium is a slender
tube about 5 mm. long; and it exhibits two separate, annular, swollen regions marking
the places where the gonads are situated. The mouth is a simple, round opening. The ento-
derm of the manubrium and of the tentacle-bulbs is of a beautiful custard-yellow, while the
entoderm of the terminal series of knobs on the tentacles is of a rich port-wine color.

Several specimens were found in a surface-tow at Tortugas, Florida, early in August,
1898, and in July, 1906.

DESCRIPTION OF EUROPEAN SPECIMENS.

For details of the general characters of the European medusa, see tabular description
of the medusae of Slabberia.

Forbes and Goodsir, 1853, describe this medusa from the western coast of Scotland, at
Tobermory and Loch Laigh.

Keferstein, 1862, and Allman, 1871-72, describe that which I doubtfully believe to be
this medusa from the coasts of France and England. It is the immature condition before
the appearance of gonads, while medusa-buds are still being produced upon its manubrium,
and before the development of nettle-rings upon its tentacles. Chun, 1895, has made a most
careful study of this budding process in specimens from the Mediterranean.

Haeckel, 1879, describes this medusa from the Mediterranean, but did not observe it
in the stage wherein medusa-buds are being produced. An identical medusa is found at
Tortugas, Florida.

Chun, 1895, has made a careful study of the budding process. When the medusa^is
young the manubrium is slender and cylindrical and without gonads. Medusa-buds develop
at this stage. These budding medusae develop in a helical line at regular intervals down
the sides of the manubrium. The oldest of the primary series of medusa-buds is nearest the
base and the youngest nearest the mouth of the manubrium.

These primary medusa-buds are attached by narrow pedicels to the sides of the manu-
brium, and a second series of medusa-buds develop upon the sides of these pedicels, before
the primary medusae are set free. After the primary medusae are set free the secondary medusa-
buds complete their development. Thus the manubrium may present various appearances;
the oldest medusa-buds at any one time may be above, or below, or anywhere upon the manu-
brium. Before they are set free the budding medusas begin to develop a secondary genera-
tion of medusa-buds upon their manubria.

The hydroid of this medusa is described by Graeffe from Trieste, Adriatic Sea. GraefFe
succeeded in rearing the hydroid from the eggs of the medusa. It is a Syncoryne. The poly-
pites are cup-shaped, only 2 to 3 mm. long, and have 4 circlets of 4 tentacles each. The
tentacles are short and stiff, and each one ends in a nematocyst-knob. The short stems of
the hydroid are invested by an irregularly ringed perisarc. The hydrorhiza forms a mass of
profusely branched anastomosing stolons. The medusa-buds arise from the sides of the poly-
pites between the 2 lowest circlets of tentacles. When set free the medusa has a short man-
ubrium and 4 tentacles. The tentacles have terminal nematocyst-knobs and a nettling ring.

The eggs are laid in May and June and the mature hydroid is found in October at Trieste.

Chun is of the opinion that Sarsia clavata is identical with "D. dolic hogaster" ; but
Hartlaub regards S. clavata as being distinct and a true Sarsia. The absence of a series of
rings upon the tentacles of S. clavata inclines me to agree with Hartlaub that it is a Sarsia
and not to be confused with Slabberia catenata. Further studies appear to be required before
any final opinion can be formed upon this case.

ANTHOMEDUS.E SLABBERIA.

79

Slabberia ophiogaster.

Sarsia strangulate, ALLMAN, 1871-72, Monograph Tubularian Hydroids, p. 46, fig. 17.

Dipurena ophiogaster=Tetrapurena ophiogaster, HAUCKEL, 1879, Syst. der Medusen, p. 25.

Dipurena ophiogaster ( ?), BROWNE, 1905, Pearl Oyster Fisheries, Gulf of Manaar, Roy. Soc. London, Suppl. Report 27, p. 133,

plate 2, figs, i, 2 (from Galle Bay, Ceylon).
Purena strangulata, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 55, figs. 51-53.

This medusa is found off the southern coasts of England and Ireland, and an apparently
identical species is described by Browne from Ceylon. Dr. Lobianco kindly permitted me
to study a specimen, which I consider to be of this medusa, from Naples, Italy.

For details, see tabular description of the medusae of Slabberia.

FIG. 36. Slabberia ophiogaster. Drawn by the author, from a specimen collected by Dr. Lobianco in the Bay of Naples.
FIG. 37. "Purena strangulata" (5. ophiogaster), after Hartlaub, in Nordisches Plankton.

In the specimen in Dr. Lobianco's collection from Naples, the bell is 5 mm. high, 4 mm.
wide; oval, with thick walls. The manubrium is 13.5 mm. long and has 2 gonads, one in the
middle of its length and one adjacent to the stomach. The 4 tentacles are each 2.5 mm. long,
and their outer halves taper to a point and are covered with broken, partial rings of nettle-
cells. The proximal halves of the tentacles are smooth and cylindrical. The tentacle-bulbs
are swollen below the bell-margin and have ocelli.

Slabberia pyramis.
Baltivcodon pyramis, HAECKEL, 1879, Syst. der Medusen, p. 26.

This Mediterranean medusa is distinguished from other species of Slabberia by the
terminal "suction cups" upon its tentacles and by the numerous "glandular" organs which
line the sides of the radial-canals. For details see the tabular description of medusae of
Slabberia.

MJ MEDUSAE OF THE WORLD.

Genus MARGELOPSIS Hartlaub.

Nemopsis (hydroid only), MCCRADY, 1857, Gymn. Charleston Harbor, Proc. Elliott Society of Charleston, vol. I, p. 163, plate

10, figs. 4-7.
Marsehftshi HARTLATB, 1897, Hydromedusen Helgolands, Wissen. Meeresuntersuch., Biolog. Anstalt auf Helgoland, Neue

Folge, Bd. 2, p. 48z; Ibid., 1904, Bd. 5, p. 99; 1899, Nachrich. k. Gesell. der Wissenschaften Gottingen, math.-phys.

Klasse, p. 219; Zool. Centralbl., Bd. IO, p. 28; 1907, Nordisches Plankton, Nr. 12, p. 88. BROWNE, 1903, Bergens

Museums Aarbog, No. 4, p. 10. MULLER, 1908, Zeit. fur wissen. Zool., Bd. 89, pp. 43, 73.

GENERIC CHARACTERS.

Anthomedusae with simple, round mouth-opening without oral tentacles, or cruciform
lips. With 4 radially placed clusters of marginal tentacles and 4 radial-canals. The ring-
like gonad encircles the stomach. The hydroid is the pelagic Margelopsis.

The tentacles of the hydroid are confined to definite whorls. In 1857, McCrady described
a free-floating hydroid from Charleston Harbor which has two whorls of tentacles, and between
them a number of medusa-buds arise from the sides of the hydranth. When set free the
medusas bore some resemblance to Nemopsis, and McCrady believed them to be the young
of Nemopsis gibbesn = Nemopsis bachei L. Agassiz. We now know this to have been a
mistake, for Hartlaub, 1899, discovered that actinula larvae develop upon the sides of the
manubrium in Margelopsis haeckehi. Then these actmulae are set free with two whorls of
tentacles, and medusa-buds develop from the sides of the hydranth between the whorls, and
are set free, as in McCrady's hydroid, which is, indeed, a species of Margelopsis.

In 1904 Hartlaub found another free-floating hydranth (Margelopsis stylostoma) which
is apparently detached from Tiarella singularis F. E. Schulze. It is apparent that the Siphon-
ophoras may have been derived from hydroid forms having a similar life-history.

The medusas of this most interesting genus appear to form a connecting link between
the Codonidae and Margelidae of Haeckel. They resemble the Codonidae in their simple
mouth-opening without oral tentacles or prominent lips, and in the single ring-like gonad
surrounding the manubrium. On the other hand, they resemble the Margelidae in having
the marginal tentacles grouped into 4 radially situated clusters. There are, however, no
tentacular ocelli such as are seen in the Margelidas.

The Atlantic species are all very closely related and may prove to be but one. The
chief distinctions are the number of tentacles upon each tentacle-bulb. For example: M. hart-
lanlm Browne has 2; M. haeckeln Hartlaub has 3 to 4; M. gibbesi Hartlaub has 510 6.

The medusa of M. stylostoma, and the hydroid of M. hartlaubii are unknown, and it is
possible that these two may be phases in the life-history of one and the same animal.

The so-called "Perigonimus sulphureus" Chun, 1889 (Sitzungsber. Akad. Wissen. Berlin,
Jahrg. 1889, p. 524), and Steche, 1906 (Zool. Anzeiger, Bd. 31, p. 30, 2 figs.), is probably
not a Perigonimus, but may give off pelagic hydranths. Chun found this hydroid grow-
ing upon a living shell of ffyaliea in the Canary Islands. One portion of the net-like
hydrorhiza gives rise to isolated polypites, and another portion to the medusa-buds. The
hydranths arise directly from the hydrorhiza and are unbranched. They are barrel-shaped,
large, and with an oral circlet of 6 to 9 short, blunt tentacles. One of the hydranths was sharply
constricted by a ring-furrow in the middle of its length as if the oral end were about to be set
free and to become pelagic (Steche). The medusa-buds also arise singly from the hydrorhiza,
each of the medusa-buds being placed upon a short, unbranched peduncle. When set free
the medusas have 4 short tentacles with swollen basal bulbs, and the manubrium does not
project beyond the velar opening. The entoderm of the hydroid and medusa is sulphur-
yellow. The periderm of the hydroid is smooth and has none of the foreign bodies attached
to it such as are seen in Perigonimus. According to Steche the colony feeds upon the eggs
which are set free by the Hyaliea on which it grows.

Margelopsis haeckelii Hartlaub.

Margelopsis hxckelii, HARTLAUB, 1897, Wissen. Meeresuntersuch. Kommis., Meere Kiel, Helgoland, Neue Folge, Bd. 2, p. 482,
taf. :6t, fign. 12-18; 1899, Nachricht. kg!. Gesell. Wissen. math.-phys. Klasse, Gottingen, Jahre 1899, p. 219, 3 fign.;

1907, Nordisches Plankton, Nr. 12, pp. 89, 91, fign. 84-86 (best published description of hydroid and medusa). MULLER,

1908, Zeit. fur wissen. Zool., Bd. 89, pp. 43, 73, taf. 5, fign. 18, 19.

PLATE 8.

Fig. I. Zancleopsls dichotoma. Tortugas, Florida, May 28, 1899.

Figs. 2 and 3. Gemmaria costata ( ?), hydroid and medusa. Tortugas,
Florida, June, 1899. This is probably the hydroid of Zanclea
costata.

Figs. 4 and 5. Zanclea gemmosa, young medusa. Figure 5 shows details of
nematocyst-capsules of the tentacles. Agassiz Laboratory, New-
port, Rhode Island, June 19, 1893.

Fig. 6. Zanclea costata. Tortugas, Florida, May 15, 1906.

Fig. 7. Zanclea costata. Details of nematocyst-knobs of tentacles.

Figs. 8 and 9. Slabberia catenata. Tortugas, Florida, August, 1898.
Figure 9 shows end of one of the tentacles of the medusa shown in
figure 8.

Drawn from life, by the author.

ANTHOMEDUS.E MARGELOI'SIS.

SI

Medusa. Bell pyriform with a flat apex 1.3 times as high as wide. 2 mm. high. Gelat-
inous walls quite thick. 3 or 4 tentacles upon each of the 4 marginal tentacle-bulbs. These
tentacles are ringed with nematocysts. No ocelli. 4 narrow, straight radial-canals and a
wide axial-canal above the stomach. Manubnum wide, spindle-shaped, two-thirds as lout;
as the depth of the bell-cavity. The upper third ot the manubnum has no gonad, but the
lower part is incased in a tubular genital organ which bears planulae. Tentacle-bulbs
brown. Manubrium with dark-brown pigment granules. Found at Helgoland, German
Ocean, in July and early in August.

Hydroid. The planulae which develop upon the manubrium of the medusa grow into
actinula larvae before being set free. The oral pole of the actinula is blunt and is adjacent
to the wall of the stomach of the medusa, while the aboral pole projects outward from the
stomach-wall into the bell-cavity. This aboral pole is conical and terminates in a sucker-
shaped depression. There is a row of 5 to 7 short, somewhat club-shaped tentacles around
the oral pole and two closely-set rows of alternately arranged tentacles near the aboral pole.
Both aboral rows have together about 12 tentacles, and these are somewhat longer than the
5 or 7 oral tentacles. The middle part of the body of the actinula is devoid of tentacles.

FIG. 38. Hydroid and medusa of Margelopsis harckelii, after Hartlaub, in Nor.iisdu-s Plankton.
Medusa is about to set free an actinula larva.

Medusa-buds develop after the actinula has been set free and has become about I mm.
long. These medusa-buds develop in clusters from the sides of the body of the actinula close
to the bases of the lower row of aboral tentacles with which they alternate in position. The
medusa-buds develop 2 or 3 tentacles in each of their 4 marginal clusters while still attached
to the actinula. Hartlaub found large numbers of these interesting hydroids floating in the
water at Helgoland, late in August.

Miiller, 1908, has studied the origin and structure of the ova in the medusa. In common
with HvbocoJon,tYie eggs are large, amoeboid, and few in number when full grown, for the
successful eggs devour the weaker ones in the ovary. The ooplasma is composed of wide
irregular fibers, forming a somewhat narrow-meshed network. The exoplasma is distinct
from the endoplasma and is wider in its meshes.

Margelopsis stylostoma Hartlaub.

Margelofiis slylostoma, HARTLAUB, 1903, Zoolog. Centralblatt, Bd. 10, p. 2S, fig. 2, fig. 4 ( ?)i 1904, Wisscn. Mccrvsuntrrsuch.
Kommis. Deutsch. Meere, Abth. Helgoland, None Folge, Bd. 5, p. 99, fig. I, fig. z (?); I97> Nordisches Plankton, Nr.
i*, p. 91, fig. 87 (88?).

This is a small spindle-shaped, free-floating hydranth, with a sucker-like expansion at
its aboral pole, and with three verticils of tentacles, i. e., 4 short-knobbed oral tentacles, a

7

82

MEDUSA OF THE WORLD.

middle zone of about 10 to 12 long-knobbed tentacles, and a basal (aboral) zone of" about 10
to 12 long-knobbed tentacles. Medusa-buds (?) This form closely resembles Tiarella
singularis F. E. Schulze, 1876, a small, isolated, attached hydroid found at Trieste. T ' . singu-
lans produces medusa at the base of the polypite below the third zone of tentacles.
Margelopsis stylostoma was found at Roscoff, northwest coast of France, in June.

FIG. 39. Hydroid of Margelopsis stylostoma, after Hartlaub, in Nordisches Plankton.
FIG. 40. Margelopsis hartlaubii, after Browne, in Bergens Museums Aarbog, 1903.

Margelopsis hartlaubii Browne.

Margelopsis hartlaubii, BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 10, plate i, fig. 2, plate 3, fig. 3. HARTLAUB, 1907,
Nordisches Plankton, Nr. 12, p. 92, fig. 89.

Bell 2 mm. wide, 2 mm. high. Egg-shaped, with thick walls. 8 marginal tentacles, 2
upon each radially placed marginal bulb. These tentacles are ringed with nematocysts, and
are somewhat shorter than the bell-diameter. No ocelli. 4 narrow radial-canals. Stomach
large, conical, with a broad, flat, quadrangular base, without an axial, apical canal. Mouth
a simple, round opening. Gonad encircling the stomach. Color (in formalin): Gonads,
stomach, and tentacle-bulbs yellowish-brown. Three specimens; coast of Norway, Osterfjord
and Herlfjord, from depths of o to 200 fathoms.

Margelopsis gibbesi Hartlaub.
Plate 9, figs. 4-7.

Ncnwpsis gibbesii, McCfiADY, 1857, Gymn. Charleston Harbor, p. 163, plate 10, figs. 4-7. FRECH, 1898, Lptli.ua pal<eozoica,
Theil i, Bd. I, Lfg. 3, p. 565, Stuttgart (compared with the Graptolites as a floating Tubularian).

Margelopsis gibbesi, HARTLAUB, 1903, Zoolog. Centralbl., Bd. 10, p. 28; 1899, Nachricht. kgl. Gesell. Wissen. math.-phys.
Klasse, Gottingen, Jahre 1899, p. 223, fig. 4.

Bell somewhat higher than a hemisphere, 2.5 mm. high, and walls quite thin and uniform.
There are 4 radially situated clusters of marginal tentacles. Each cluster contains 5 or 6
tentacles which arise from a large common basal bulb. Tentacles taper regularly from base
to tip, but the tip terminates in a knob of nematocysts. There are about 15 rings of nemato-
cysts upon the shaft of each tentacle and the entodermal axis of the tentacle is composed
of chordate cells resembling those of Obeha. There are 4 straight, narrow radial-canals.
Velum well developed. Manubrium wide and flask-shaped, and mouth a simple, round
opening without prominent lips, and without oral tentacles. In the female the ova project
prominently from the surface of the manubrium. The gonad surrounds the stomach on all
sides. The bell of this medusa is highly contractile, often drawing together so as to cause
the mouth to project beyond the velar opening. When the bell is relaxed, however, the manu-
brium extends but little more than half-way from the inner apex of the bell-cavity to the
velar opening.

PLATE 9.

Fig. I. Cladonema perkinsn. Nassau Harbor, Bahamas.

Fig. 2. Cladonema mayen. Moat of Fort Jefferson, Tortugas, Florida,

July, 1905.
Fig. 3. View looking down upon the aboral end of the stomach of ClaJont ma

mayeri, showing the arrangement of the radial-canals.
Fig. 4. Margelopsts gibbesn, male. Beaufort, South Carolina, December

24, 1904.
Fig. 5. Margelopsis gibbesn, male. Oregon Inlet, Pamlico Sound, North

Carolina, November 12, 1904.

Fig. 6. Margelopsis gibbesn, female. Cape Fear, North Carolina, Decem-
ber I, 1904.

Fig. 7. Tentacle of Margelopsis gibbesii.
Figs. 8 and 9. Stomotoca dinerna, young females. Agassiz Laboratory,

Newport, Rhode Island, July, 1892.
Fig. 10. Stomotoca dinema, young male. Agassiz Laboratory, Newport,

Rhode Island, Septemper 15, 1896.

Fig. I, copied from a drawing by Prof. Henry F. Perkins.
Figs. 2 to 10, from life, by the author.

PLATE 9

10

ANTHOMKnrS.E MARGKLOPSI.S, l'KI.A( ;< iin I'll \.

83

The entoderm of the four large tentacle-bulbs is dull-yellow. There are no ocelli. The
entoderm of the manubrium was dull-\'ellow in two medusx, and dull-green in another. The

J O

difference is not a sexual one, however, for one ot the males was yellow while the other \\as
dull-green. The two yellow-colored medusa; were found, one in Oregon Inlet, Pamlico
Sound, North Carolina; and the other at Southport, Cape Fear, North Carolina, in Novi-mln-r.
The green-colored male (plate 9, fig. 4) was found in Beaufort River, at Beaufort, South
Carolina, on December 24, 1903, 12 miles from the ocean, in brackish water.

McCrady, 1857, presents a good figure of that which is probably the free-floating hydroid
of this species. The hydroid is pear-shaped with a hollow cup-like projection from its broad
aboral end. There are 2 whorls of tentacles, a circlet of about 8 tentacles around the broadest
zone near the aboral end and a circlet of about 6 tentacles near the mouth. A number of
medusa-buds develop from the sides of the hydranth in the middle region between the 2
circlets of tentacles. When set free the medusae have 4 short tentacles in each radial cluster.
A single specimen was found by McCrady in Charleston Harbor. The cup-shaped projection
from the aboral end of the hydranth is a feature which recalls the pneumatophore of the
Siphonophorae.

Genus PELAGOHYDRA Dendy.

Pelagohydra, DENDY, 1903, Quart. Journ. Microscop. Sci., vol. 46, p. i. HARTLATB, 1903, Zool. Centralblatt, p. 27.

The type species is Pelagohydra mirabilis, Dendy, from New Zealand.

GENERIC CHARACTERS.

Medusa similar to Margelopsis. Hydroid pelagic and closely resembles that of Marge-
lopsis, but tentacles arise irregularly from the sides ot the hydranth, and are not arranged in

definite circlets as in Margelopsis,

Pelagohydra mirabilis Dendy.

Ptlagohyiira mirabilis, DENDY, 1903, Quart.
Journ. Microscop. Science, vol. 4(1, p. i.
plate 12. HARTLAIB, 1903, Zool. Cen-
tralbl., p. 27, fig. 3.

Hydroid pelagic, with a trans-
lucent globular body 25 mm. wide,
constituting a float and a proboscis
at the end of which the mouth is
situated. The float bears many
long, irregularly arranged tenta-
cles. The proboscis is elongate,
pink in color, and of the shape of
a truncated cone. It has several
irregular whorls ot oral tentacles,
the shortest being nearest the
mouth. Medusae are borne upon
branching stolons which arise from
the sides of the float between the
tentacles. When about to be set
free the medusa is about i mm.
wide, with 4 radial groups of m.i i -
ginal tentacles, each group arising
from a large, pink, basal bulb, and consisting of 5 tentacles, 2 large, .2 small, and i smallest
median tentacle. 4 radial-canals. Manubrium pyriform and without oral tentacles. Nt\\
Zealand.

Family CLADONEMIDJE.

CladonemiJt, GKI;I:NBAUR, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 220. HM.CKI.I.. 1879, Sys,t. ilcr Medusen, p. 98. HART-
LAI-B, 1887, Zool. Anzcigcr, Bd. 10, p. 651; 1907, Nordisches Plankton, Nr. 12, pp. 5, no. GUNTHER, 1903, Mittheil
Zool. Sta. Ncapel, Bd. 16, pp. 56, 57.

FIG. 40*3. Pelagoh\dra mirabilis, after Dcmiy, in Quarterly Journal Micr
scop. Science.

a, entire li\ilr<>i,l, twice natural size.

b, enlarged view, showing medusa-buds developing among tentacles.

84 MEDUSA OF THE WORLD.

This family is of such interest that we devote more than the usual space to its consider-
ation. Haeckel, 1879, classifies it as a family coordinate with the Codonidae, but Hartlaub,
1887, showed that the genus Cladonema had an encircling gonad as in the Codonidae. The
medusa; are, however, highly specialized, both in respect to their anatomy and their mode of
life, and they are undoubtedly derived from some more simply-organized medusae among
the Codonidae. The hydroid of Cladonema is a StanriJia, and is closely related to Syncoryne,
the hydroid of Sarsia.

CHARACTERS OF THE FAMILY CLADONEMID^.

Anthomedusae with feathered or branched marginal tentacles, and with four or more
simple or branched radial-canals. The gonads may be ring-like and encircle the stomach, or
they may be more or less separated so as to be interradial or adradial, or developed in a spec-
ialized brood-pouch above the stomach.

Hartlaub, 1887, and Perkins, 1902, show that in the genus Cladonema the genital products
are developed over the entire gastric portion of the manubrium and the gonads are not con-
fined to restricted meridians, as was believed to be the case by Haeckel. Hartlaub found that in
Cladonema radiatum of the Mediterranean the genital products begin to develop in the ento-
derm. This species exhibits a successive hermaphroditism, although either sperm or ova may
precede. Perkins, however, in his study of Cladonema pcrkinsn of the Bahamas, found that
the genital products were developed in the ectoderm of the manubrium, and he failed to find
sperm or ova in the entoderm.

In Eleutheria, Hartlaub, 1 886, 1887 (Zool. Anzeiger), showed that the genital products
are developed exclusively in the ectodermal lining of a peculiar brood-pouch above the stomach,
and that the medusa is hermaphroditic. The brood-pouch is derived from the ectodermal
layer of the bell-cavity and is not connected with the stomach of the medusa, but communi-
cates with the bell-cavity by means of 6 interradial-canals.

Hartlaub, 1887, divides the Cladonemidae into two groups, one with and the other without
an apical brood-sac above the stomach. As this brood-sac is highly variable in its develop-
ment, some specimens of Eleutheria apparently failing to develop it and, moreover, as it is a
character which develops during the growth of the medusa and is not found in the young
animal, it would seem that Haeckel's classification of the Cladonemidae, based as it is upon
the possession of constant characters, is to be preferred. Moreover, as has been shown by
Giinther, 1903, we can not yet state that the brood-pouch of other genera of Cladonemidae is
similar in structure to that of Eleutheria.

As is well known, Haeckel, 1879 (Sitzungsber. Medicin. Naturwiss. Ges. Jena; see also
Chun, 1880, Ctenophoren des Golfes von Neapel), drew an ingenious comparison between
Ctenaria and the Ctenophorae, and concluded that the Ctenophorae might have been derived
from some form of Anthomedusa. Ctenaria appears to resemble the Ctenophorae in its 2
feathered tentacles, 4 bifurcated radial-canals, and in a peculiar sheath at the base of each
tentacle. However, Hartlaub's discovery that the brood-pouch in the closely-related Eleu-
theria is of ectodermal and not entodermal origin makes it appear that the resemblances
between Ctenaria and the Ctenophorae are of the nature of a mere parallelism and not indic-
ative of a genetic relationship. Haeckel compared the 8 adradial longitudinal lines of
nematocysts upon the exumbrella of Ctenaria with the 8 rows of ciliated combs in the
Ctenophorae. An even more remarkable case of convergence is shown by Kofoid, 1905
(Bull. Mus. Comp. Zool. at Harvard College, vol. 46, p. 163, i plate), in the case of the
Cystoflagellate, CraspeJotella, which is bell-shaped and provided with a well-developed velum,
the walls of which are contractile so that the mode of locomotion is similar to that of a medusa.
There is thus an external similarity of form between this protozoan and a medusa offering an
instance of convergence of a most striking character. Another remarkable instance of conver-
gence is that of the pelagic holothurian Pelagothuria natatrix, which bears a close resemblance
to a medusa (see Mem. Mus. Comp. Zool. at Harvard College, vol. 17, 1894). I believe also
that the bell of the Narcomedusa? is not homologous with that of the Anthomedusae and
Leptomedusae, but is a mere outgrowth from the walls of the actinula larva.

ANTHOMEDUS.E ZANCLEA. 85

The Cladonemidae are close relatives of the Codonidae, this being illustrated most cleaily
by their hydroids, and they may properly be regarded as highly-specialized Codonidae, as has
been maintained by Hartlaub. For purposes of classification, however, I think it will be
well to permit them to remain in a family of Anthomedusae coordinate with the Codonidae, for
there is no single character which they have consistently in common with the Codonidae. For
example, the gonad in ClaJonemamzy be readily likened to that of the Codonidae, for it encircles
the stomach on all sides, but in some species of Zanclea the gonads show a decided tendency
to segregate interradially, and in Elcutheria they are developed in an ectodermal brood-sac
which is not connected with the manubrium.

The most constant character of the Cladonemidae is their forked, or feathered, marginal
tentacles.

The greatest difficulty in any attempt to classify the Cladonemidae among themselves
arises from our lack ot understanding of the nature of the brood-sac or apical cavity in the bell
ot the genera Pteroncma, Ctenarta, and Dendronema; and no classification worthy of serious
consideration can be effected until we know whether this sac is a mere extension of the stomach
or an ectodermal reproductive cavity similar to that studied by Hartlaub in Elcutheria.

FAMILY: CLADONEMIDAE. Subfamily Pteroneminai

Manubrium without oral tentacles. With 4 to 8 simple radial-canals.

Zanclea GEGENRAUR, 1856. With 2 to 4 marginal tentacles at bases of 4 radial-canals. The tentacles give rise to an
abaxial row of simple branches, each terminating in a cluster of ncmatocysts. With meridional rows of nema-
tocysts upon the exumbrella. No ocelli. No brood-pouch above the stomach. The hydroid is Gtinmaria McCrady,
Allman.

Zancleofish HARTLAUB, 1907. Similar to Zanclea, but with tentacular ocelli and without meridional rows of nemato-
cysts.

Pteronema HAECKEL, 1879. With 4 well-developed tentacles similar in structure to those of Zanclea. With 4 simple
radial-canals. With a brood-sac above the stomach. Hydroid unknown.

Eleutheria QTATREFAGES, 1842. With 4 to 6 simple radial-canals, and an equal number of bifurcated tentacles. With
a brood-pouch above the stomach. This brood-pouch is not connected with stomach-cavity, but communicates
\vith bell-cavitv by means of simple interradial pores. Genital products develop exclusively in brood-pouch. Medusa
is hermaphroditic. Hydroid is Clavatella Hincks.

Mnestra KROHN, 1853 (Archiv. fur Naturges., Jahrg. 19, Bd. i, p. 278). Medusa degenerate, sessile upon Pliyllirhoc
in the Mediterranean. With 4 to o small marginal tentacles having nettling capsules along their abaxial sides. A
ring of nematocysts extends around the bell-margin, and 4 linear tracts of nettle-cells extend upward over the
exumbrella, t from each tentacle-bulb. With 4 simple, unbranched radial-canals and ring-canal. No brood-pouch
above stomach. Throat of medusa blocked by a spongy mass of entoderm. Medusa attaches itself by its mouth to
throat of Phyllirhoe, and the larvae of the Mnestra probably develop within the gastric cavity of the Phyllirhoe* It
is elaborately described by Gunther, 1903 (Mitt. Zool. Sta. Neapel, Bd 16, pp. 35-62, plates 2, 3).
FAMILY: CLADONEMID.'E. Subfamily Dendroneminte:

The mouth is surrounded by simple, or branched, oral tentacles. Some or all of the radial-canals bifurcate.

Ctrnaria HAECKEL, 1879. With 2 marginal tentacles, each of which gives rise to a row of nematocyst-bearing filaments.
With 4 bifurcated radial-canals. With simple, unbranched, oral tentacles. With an apical sac above the stomach.
Hydroid unknown.

Clailonema DTJARDIN, 1843. With 4 to 5 bifurcated radial-canals, or 8 to 10 simple radial-canals, or with both simple
and bifurcated radial-canals. With a tentacle at foot of each radial-canal. These tentacles branch dichotomouslv
or complexly, and branches end in suckers or nematocyst-knobs. With simple, unbranched, oral tentacles. No
brood-sac above stomach. Hydroid, Siauridia Dujardin.

Deridroneiiia HAECKEL, 1879. With bifurcated radial-canals, and dichotomously branched tentacles. Branches end in
suckers, or in nematocyst-knobs. Mouth surrounded by 4 groups of dichotomously branched oral tentacles. There
is an apical sac above the stomach.

Genus ZANCLEA Gegenbaur, 1856.

Zanclea, GEGENBAVR, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 229. KEFERSTEIN UNO EHLERS, 1861, Zool. Beitr. in Xt-apcl und

Messina, p. 85. AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344.
Zariflea-t- Gemtntiriii, HAE( KEI., 1879, Syst. der Mcdlisen, pp. IO2, 103.
Zanclea, FEWKES, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 150. HARTLAI/B, 1887, Zoo], Anzeiger, Bd. 10, p. 654; njo^, N'or-

dischcs Plankton, Nr. 12, p. 112. VANHOFFEN, 1891, Zool. Anzeiger, Jahrg. 14, p. 445. <!i NIH>K. i'|o}. Mitt. Zool.

Sta. Neapel, Bd. 16, p. 57.

Gemmaria= Zanclea, BROWNE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 750.
Gemmaria, McCfiADY, 1857, Gymn. Charleston Harbor, p. 49. A<:, i '. \ . iSi^, North Amcr. Acal., p. 184. An \nx, 1872,

Monog. Tubul. Hvdroids, p. 289. BROWNE, 1896, Proc. X.ool. Sot. London, p. 4111.

\'on Gemmaria, FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard I'. .Hi vol. 8, p. 150.

Gemmaria, HARTLAI'B, 1887, Zool. Anzeiger, Bd. 10, p. 6^4. -fJiM'Hm, [903, Mitt. Zool. Sta. Neapel, Bd. if>, p. 57.

Gemmaria (hydroid), HARTLAI R, 190$, Zoologischen Jahrbiich., Supplement 6, p. 527.

Corvnith, Mi r. i:v 11. 1^90., Quart. Journ. M ici., new ser., vol. 42. p. ^4. \t TTING, 1901, Bull. I'. S. Fish Commission

for iScfij, pp. ^29, 372. HARGITT, 1904, Bull. U. S. Bureau uf Fisheries, vol. 24, p. 42.
Halorharis (hydroid), Ai.tssiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 239, plate 20, figs. io-ioc.
Gemmaria+ Zanclea, HARGITT, 1908, Biol. Bulletin, vol. 14, p. 104.

The type species of this genus is Zanclea costata Gegenbaur, 1856, of the Mediterranean.

Mi

MEDUSAE OF THE WORLD.

Tabular Description of the Medusa of Zanclea.

Z. costata Gegenbaur,
1856.

Z. gemmosa McCrady,
1857-

Z. implexa Allman,i864

Z. cladophora = Gem-
maria cladophora A.
Agassiz, 1865.

Shape and size of bell
in mm.

Ellipsoidal. 4 to 5 high,
3 to 4 wide.

Pyriform. 6 high, 4
wide.

Dome-like to globular.
1.5 high, 1.75 wide.

Pyriform. Size (?)

Length of the 4 merid-
ional rows of nettle-
cells above tentacle-
bulbs, in terms of

From tentacle-bulbs to
very near a pex of bell .

One-fourth to one-third
distance from margin
to apex.

0.25 to 0.5

-5

bell-height.

Number and length of
tentacles in terms of
bell-radius (r).

4 tentacles, each 3 to 4 r
long-

4 tentacles, each 3 to 4 r
long.

2 to 4 tentacles, each
about 4 r long.

2 long and 2 short ten-
tacles; long tentacles
4 r, short over one-third
r long.

Character of abaxial
filaments upon ten-
tacles.

Very numerous, slender,
and ending in a spheri-
cal capsule containing
3 to 5 nematocysts.

Very numerous, slender,
and ending in a spheri-
cal capsule containing
3 to 9 nematocysts.

Very numerous, slender
filaments, each ending
in an oval capsule
containing 4 to 8
nematocysts.

Very numerous, slender
filaments, ending in
knob-like clusters of
nematocysts. Num-
ber of nematocysts in
each knob ?

Length and shape of
manubrium.

Not quite half as long as
depth of bell-cavity.
Flask-shaped.

Flask-shaped, 4-sided.
As long as depth of
bell-cavity . Mouth
with 4 small lips.

Urn-shaped. Only half
as long as depth of
bell-cavity. Mounted
upon short peduncle.
Mouth round opening.

Conical. 4 prominent
lips.

Gonads.

4 interradial masses of
eggs on swollen sides of
stomach.

On 4 interradial sides of
stomach. Much
swollen.

On upper sides of
stomach, more or less
separated in 4 chief
radii.

Gonad encircles stom-
ach.

Color.

Gonads red, tentacles
yellowish-brown.

Entoderm of stomach
and tentacle-bulbs
creamy-pink. Ento-
derm of manubrium
delicate green.

Stomach and tentacle-
bulbs brownish-red to
crimson. Entoderm
of tentacles orange.

Tentacles light-brown.
Orange and yellow
spots in tentacle-
bulbs. Color of stom-
ach ( ?)

Where found.

Mediterranean. Common.
Medusa having only 2
tentacles, but in other
respects similar to Z.
costata found at Tor-

West Indies northward
to Woods Hole.

Atlantic coasts of west-
ern Europe to Medi-
terranean.

Coast of Massachusetts,
Nahant, Woods Hole.

tugas, Florida.

Hydroid.

Hydroid ( ?) (see text).

Hydroid: Gemmaria
gemmosa.

Hydroid: Gemmaria
implexa.

Hydroid unknown.

GENERIC CHARACTERS.

Cladonemidae with 4 simple radial-canals, and 2 to 4 radially situated tentacles which give
rise to filiform, knobbed, nematocyst-bearing side branches. The gonad is developed upon the
sides of the stomach. There is no apical brood-pouch above the stomach. There are merid-
ional lines of nettle-cells over the exumbrella. There are no tentacular ocelli. The hydroid
is Gemmaria Allman, 1871.

In 1857, McCrady founded the genus Gemmaria for a Zanclea-\ike medusa with only 2
tentacles. Vanhoffen, 1891, and Browne, 1905, have observed that the 2-tentacled young of
other species of these medusae often becomes 4-tentacled when adult, and according to Browne,
1905, "Gemmaria" implexa may become mature with 2 or with 4 tentacles. It is evident,
therefore, that there are no definite generic distinctions between Zanclea and Gemmaria, the
"Gemmaria" condition being only a developmental or retarded phase of Zanclea.

Murbach and Nutting have erroneously applied the name "Corynitis" to a 2-tentacled
medusa of Zanclea. Apparently these authors had not seen the medusa of Corynitis, which is

ANTHO.MKDCS.E ZANCI.KA.

well figured by McCrady. I have frequently taken Corynitis in Charleston Harbor, South
Carolina, where McCrady discovered it, and it is wholly different from any "Geniniaria" or
Zanclea. (See also Hargitt, 1908.)

The common '/aticli-a ot the southern coast of New England, in America, may become
mature with only 2 tentacles, and with poorly developed lips. On the other hand, a vcrv
similar medusa is occasionally found in the same waters with 4 tentacles and with 4 well-
developed lips. I am inclined to believe that these conditions represent developmental staues.
or starved and well-fed states of one and the same medusa.

Zanclea costata Gegenbaur.
Plate 8, figs, z, 3, 6, and 7.

Zanclea conaia, GEGENBAUR, 1856, Zeit. fur wisscn. Zool., Bd. 8, p. 229, taf. 8, figs. 4-7. HAECKEL, 1879, Syst. dcr Mcdu sen

p. 103.
Hydroid ( ?), MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 35, plate 41, figs. 137, 138.

Gegenbaur found many specimens of this medusa in the Mediterranean. His fully

developed medusae had 4 long, equally developed tentacles. For details see tabular description

of the medusae of Zanclea.

I have found a medusa at Tortugas, Florida, which accords well with 7, ancle a costata,

excepting that it has but 2 tentacles.

Medusa from Tortugas, Florida. Bell 6 mm. high and ellipsoidal, with bluntly-rounded

apex. Without an apical projection. Bell-walls of uniform thickness everywhere. There are

2 long, spindle-shaped, rough-looking tentacles somewhat longer
than the bell-height. These are thickly covered with papillae and
beset on their outer sides with a row of filaments, each of which
ends in a spherical knob, the surface of which is covered with
blunt papillae. Each of these knobs contains a cluster of nemato-
cysts (plate 8). Basal bulbs of tentacles are elongate, spindle-
shaped, and hollow. No traces of tentacle-bulbs 90 away from
the 2 long tentacles. 4 swollen lines ot nematocysts extend upuaul
over the exumbrella from the bases of the 4 radial-canals and
nearly meet near the apex of the bell. Velum well developed. 4
radial-canals, and the circular vessel straight and narrow. Manu-
brium flask-shaped, with 4 simple lips. Entoderm of the manu-
brium and tentacles milky white, There were 4 interradial
gonads. A single specimen was found in a surface-tow at Tor-
tugas, Florida, on May 15, 1906.

This species is distinguished by its ellipsoidal, high, rounded
bell, without a sharp-pointed apex, the 4 very long meridional lines
of nematocysts over the exumbrella, and the spherical nematooyst-
capsules at the ends ot the tentacular filaments.

The Tortugas medusa differs from //uiclea cost<ita,as described
by Gegenbaur, in having but 2 tentacles. It is well known, hu\v-

Flc. 41. Zanclea costata, from the
Mediterranean, after Gegen-
baur, in Zeit. fiir wissen.
Zool., Bd. 8.

ever, that in Zanclea some individuals ot the same species develop 4 while others become
mature with but 2 tentacles.

Gegenbaur states that the female gonads are red (pink '). In the Tortugas specimen the
male gonads were milky-yellowish white.

Hydroid (plate 8, fig. 2). That which is probably the hydroid stock of this species was
tound by Mayer, 1900, at Tortugas, Florida, and erroneously ascribed to '/.anclea gctntnosa;
but Murbach, 1899 (Quart. Journ. Micros. Sci., New Series, vol. 42, p. 354), describes and
figures the hydroid ot '/.anclea genniiosa, and it is now evident that the form described In
Mayer is specifically distinct from Z. gemmosa and is probably Zanclea costata, the medusa
of which occurs at Tortugas.

Hydroid of Zanclea costata (?). The hydrorhiza is creeping, tubular, and net-like, and
gives rise at irregular intervals to short, more or less twisted hydrocauli. Both the hydrorhiza
and hydrocauli are covered with a horny, chitmous pensarc, which is annulatcd in the hydro-

MEDUSAE OF THE WORLD.

caulus. The hydrocaulus is corrugated and opaque in color throughout its length, and in
this respect differs from the European G. nnplcxa Allman, 1871. The fully developed
hydranths are only 1.5 mm. in height. They are elongate, spindle-shaped, and taper gradually
from near the base to the summit, so that the diameter near the mouth is less than that near
the base. The tentacles arise in 5 to 8 well-defined whorls from sides of hydranth, each whorl
consisting of 4 to 6 tentacles. These tentacles are short and taper from base to end, termi-
nating in a large knob containing nematocysts. The entodermal cores of the tentacles are
composed of vacuolated cells. The tentacles are stiff and project rigidly outward from the
sides of the hydranth. 4 to 8 medusa-buds in various stages of development arise from the
sides of the hydranth near the base immediately below the lowest whorl of tentacles.

When set free the young medusa (plate 8, fig. 3) has 2 well-developed, radially situated,
diametrically opposed tentacles, and 2 rudimentary tentacle-bulbs. There are 4 radially
situated, nematocyst-bearing ridges upon the exumbrella, and these extend about half-way
up the sides of the bell. The bell-walls are very thin and flexible. There are 4 straight,
narrow radial-canals, and the manubrium is a short, simple tube with no trace of the gonad.
The mouth is a simple, round opening. Before being set free the tentacles are coiled inward
so as to lie within the bell-cavity. Soon after the medusa has become liberated, however,
the tentacles uncoil and project outward. The deep-lying entoderm of the hydranths is a
delicate creamy-pink, while the more superficial entoderm is translucent and milky in color.
The cells of the superficial entoderm are large and vacuolated. The hydrorhiza and hydro-
caulus is protected by a horny, yellow, amber-colored, chitinous sheath, but this does not
extend over the hydranths. The hydroid was found at Tortugas, Florida, in June, 1899,
growing upon a piece of floating gulfweed (Sargassum).

Zanclea gemmosa McCrady.

Plate 6, fig. 7; plate 7, fig. 5; plate 8, figs. 4 and 5.

Gemmaria gemmosa, McCRADV, 1857, Gymn. Charleston Harbor, p. 49.

Zanclea gemmosa, McCRADV, 1857, Gymn. Charleston Harbor, p. 49, plate 8, figs. 4, 5. AGASSIZ, L., 1862, Cont. Nat. Hist.

U. S., vol. 4, p. 344. HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 121, fign. 116, 117.
Gemmaria gemmosa, AGASSIZ, A., 1865, North Amer. Acal., p. 184, fig. 306. HAECKEL, 1879, Syst. dcr Medusen, p. 104.

FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, No. 8, p. 150, plate i, figs. 8-12. FEWKES, 1884,

American Naturalist, vol. 18, p. 196, fig. 2.
Corynitis agassizii (hydroid), MURBACH, 1899, Quart. Journ. Micr. Sci., vol. 42, p. 354, plate 34, fig. 12 (medusa, figs, n,

I3 H-)

Gemmaria sagiltaria, HAECKEL, 1879, Syst. der Medusen, p. 103, taf. 7, figs. 3, 4 (young medusa).
Zanclea gemmosa (hydroid), WALLACE, 1908, Carnegie Institution Year Book, No. 7.

Adult medusa. Bell 6 mm. in height, and pynform. There is a well-developed, solid,
apical projection. Side walls quite thin. There are 4 long, radially situated tentacles with
large, hollow basal bulbs, which lack ocelli. The shafts of the tentacles are crowded with
large numbers of short filiform branches, which arise only from the dorsal (exumbrella)
side of the tentacles, although the twisting of the tentacle may cause them to appear to arise
from all sides. Each of these filiform branches terminates in a large spear-head-shaped
body which bears a number ot long, delicate, sharp-pointed bristles, and which contains
3 to 9 egg-shaped nematocyst-cells. A club-shaped swelling extends longitudinally from
the base of each tentacle a short distance up the side of the exumbrella. These swellings are
thickly covered with nematocysts. There are 4 straight, slender radial-canals, and a narrow
circular vessel. The velum is wide and delicate. The manubrium is flask-shaped, and cruci-
form in cross-section and extends to the level of the velar-opening. The mouth is a simple,
cruciform opening without prominent lips. The gonad is upon the sides of the stomach.
These sides are greatly swollen and give to the manubrium its flask-shaped appearance.
The entoderm of the manubrium and tentacles is of a delicate creamy-pink and the ectoderm
of the manubrium is of a delicate shade of green.

Young medusa (plate 7, fig. 5; plate 8, figs. 4, 5). In the medusa, I mm. in height,
there are but 2 marginal tentacles. The bell-walls are very thin and the manubrium is a
simple fusiform tube. There is no apical projection to the bell, and no trace of gonads upon
the manubrium, and the mouth is a simple, round opening without cruciform lips.

This species extends from the West Indies to the southern coast of New England. It is
quite common in Newport Harbor late in the summer.

ANTHOMEDUS^E ZANCLKA.

S'.)

The hydroid stock has heen described by Murbach under the name of "Cor\nitis
agiissizii," but is evidently a Gtmnmria. The hydranths are 1.5 to 2 mm. in length and
arise singly from a slender, thread-
like, anastomosing hydrorhiza. The
hydrorhiza is covered with a dc-h-
cate perisarc, but this does not
extend over any part of the hy
dranths. In this respect the species
is very different from Z. costiiin.
The hydranths are also more cylin-
drical and uniform in width through-
out the polypite than in Z. fostatn.
The tentacles are irregularly ar-
ranged and the medusa-buds are
found not only near the base of the
hydranth, but scattered along the
sides. In Zanclea costata they are
confined to a zone immediately under
the lowest circlet of tentacles. The
color of both forms is quite similar,
excepting that the hydrorhiza is deep
pink in the Z. gemmosa and rosin-
yellow intheZ.co.sta/rt. W.S.Wallace
found this hydroid growing upon
floating Sargassum at Tortugas,
Florida, in July, 1908.

The young medusa
here figured was obtained

at Newport, Rhode Is- Fic.42. Z0nr/fagfmmo5a,fromFewkes,inBull.Mus.,Comp.Zool. at Harvard College.

land, and was identified by Dr. Alexander Agassiz as the young of '/.anclca gemmosa.
Z. gemmosa readily devours the medusae of Obeha.

Zanclea iraplexa Allman.

Cor\nt pelAgiea+ Corytif implexa (hydroid and medusa), ALDER, 1857, Catalogue Zoophytes Northumberland, Trans. Tync-

sidc Nat. Field Club, vol. 3, pp. 13, 18 (of the "separate"), plate 7, figs. 1-6.
Zanclea implexa, ALLMAN, 1864, Annals and Mag. Nat. Hist., vol. 13, p. 357. HINCKS, 1868, Hist. British H^lr.n.l /.. ',,)..

p. 59, plate 9, fig. 3. HARTLAUR, 1907, Nordisches Plankton, Nr. 12, p. 116, fign. 106-111 (list of localities, authors,

and best description of the medusa).
Gemmariii implexa, ALLMAN, 1871-72, Monog. Tubularian Hydroids, pp. 223, 290, plate 7, figs. i-io. H \M x n . iX-<>. V. i.

dcr Medusen. p. 105. JICKELI, 1883, Morphol. Jahrbuch, Bd. 8, p. 6n, taf. 26 (hi-riiWv ). CKJ, N-. 4 . Arbeit

Zool. In.st. \Vien, Bd. 5, p. 353. BROWNE, 1896, Proc. Zool. Soc. London, p. 461. HARI.ITT, 1904. Mnth. /.nol. Si.iiinn

Neapel, Bd. 16, p. 574, taf. 22, figs. 27-29 (hydroid and medusa from the Bay of Naples). BKOWNI, 1905, Proc. Roy.

Soc. Edinburgh, vol. 25, p. 750.

This medusa is closely related to /.unclea gemmosa of the Atlantic coast of the I'mted
States, but it is much more swollen than the American form, and differs in other n-spivts.

The bell is globular with fairly thin side walls and a thick, solid, dome-like apex. It
is 1.5 mm. high, 1.75 mm. wide. There are 4 prominent nematoryst tracts extending one-
fourth to half the distance from the tentacle-bulbs up the sides of the bell, and the nettle-
cells in each tract are arranged in two rows.

There are usually 2 long tentacles 1 80 apart alternating with 2 short tentacle-bulbs,
but according to Browne, 1905, the medusa may become mature with 2 or with 4 long ten-
tacles. The long tentacles are about twice as long as the bell-diameter and their upper,
abaxial sides bear each a double row of numerous filaments, which terminate in oval capsules
containing 4 to 8 nematocvsts. The basal bulbs of the tentacles are globular, and lack ocelli.

There is a slight linear (glandular?) swelling occupying the middle third of each of the
4 radial-canals. The stomach is mounted upon a short peduncle and is urn-shaped, with
a simple, round mouth-opening. The gonads surround the stomach and are more or less

90

MEDUS/E OF THE WORLD.

separated in the 4 principal radii. The basal bulbs of the tentacles and the stomach are
brownish-red to bright crimson, and the entoderm of the tentacles is orange.

This medusa is found off the Atlantic coasts of Western Europe, and in the Mediter-
ranean. Graeffe, 1884, states that at Trieste, Adriatic Sea, the medusae bud forth from the
hydroids in the spring, and a second brood appears to develop in summer.

The hydroid is Gemmaria implexa, well described and figured by Allman, 1871-72.
The stems of the hydroid are erect, 6 to 8 mm. high, and arise at intervals from a creeping,

net-like hydrorhiza which is attached to
shells, stones, etc., in shallow water. The
stems are usually single, but occasionally
they branch. The perisarc covers the
hydrorhiza and extends as a transversely
corrugated, opaque tube up the lower
part of each hydranth. Below the
zones of tentacles the perisarc becomes
abruptly smooth, and ends in a slightly
flaring cup. The hydranths are cylin-
drical to club-shaped, with dome-like
peristome. There are 40 to 50 short
tentacles, each ending in a knob of
nematocysts. These tentacles are ar-
ranged in 10 to 13 irregular verticils of
about 4 tentacles each, the tentacles
being about 90 apart. The medusa-
buds are borne upon peduncles which
arise in clusters from the sides of the
hydranth below the middle, but above
the lowermost verticils of tentacles.

When set free the medusa has 2
well-developed tentacles which give
rise to numerous filaments, each ending
in a capsule bearing terminal bristles
and containing several nematocysts.
There are also 2 small tentacle-bulbs
90 apart from the well-developed ten-
tacles. The bell is oval and higher
than a hemisphere and 4 lines of
nematocysts extend up about one-fourth
to one-third of the bell-height, one above
each tentacle-bulb. The 4 radial-canals
are straight and simple, and the manu-
brium is a simple tube with a round
mouth-opening and straight cylindrical

Fie. 43. Hydroid of Zanclea implexa, after Allman, in Ray sides. It lacks a peduncle and is about

Society, 1871-72. See also, Fig. 44 . na lf as l on g as depth of bell-cavity.

The hydranths are white with pale-pink entoderm. The perisarc on the proximal portion
of the hydrocaulus is brown and on the distal portion, colorless. It is closely related to the
hydroid of Zanclea gemmosa.

Zanclea cladophora Hartlaub.

Gemmaria cladophora, AGASSIZ, A., 1865, North Amer. Acal., p. 184, figs. 307-310. HAECKEL, 1879, Syst. der Medusen, p. 104.
NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 371, fig. 78. HARGITT, 1904, Bull. U. S. Bureau of

I-' i .lirrn-s, vol. 24, p. 42.
Zanclea cladophora, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 121, fign. 112-115.

Bell pyriform and about as broad as high. Apical projection solid, rounded, and dome-
like. There are 2 long, diametrically opposed tentacles from the outer sides of which arise
a great number of slender filaments, each terminating in a knob-like body which contains

ANTHOMEDUS.E ZANCLKA. XAM MXH'SIS.

91

nematocysts. These long tentacles are 1.5 times as long as the bell-diameter. There are also
2 small, conical tentacles 90 apart from the long ones. The small tentacles do not give rise
to side-filaments. No ocelli. A ridge of nematocyst-cells extends from the base ot each ten-
tacle about half-way up the side of the exumbrella. Velum broad. There arc 4 broad radial-
canals and a narrow circular tube. The outline of the bell-cavity does not follow that of
the outer surface of the bell, but is widest at the inner apex. Manubrium conical, with a wide-
proximal base and regularly-narrowing sides. It does not extend quite to the level ot the
velar opening. The 4 lips are quite prominent, and the mouth is a rectangular opennm.
The mouth is surrounded by 2 or 3 rows of large nematocyst-cells, similar to those found
upon the sides of the exumbrella. The gonad is developed upon the sides of the stomach,
which it encircles. The tentacles are light-brown in color, and the entoderm at their bases
contains orange pigment. There are bright-yellow pigment-spots at the bases ot the two
rudimentary tentacles. This species was found at Xahant, Massachusetts, in 1862, by Dr.

Alexander Agassiz. Nutting, 1901, obtained
it at Woods Hole, Massachusetts, in August,
1899. Neither of these observers states the
dimensions of the medusa.

The peculiar outline ot the subumbrella
and the constriction above the lips figured
by A. Agassiz may be due to unnatural
contraction.

Many of the characters of this so-called
remain in doubt.

species

Zanclea nitida.

Gtmmaria nitida, HARTLAI B, 1905, Zoolog. Jahrbiichrrn.
Suppl. 6, p. 527, fign. J, H (hydroid).

PolvpitesaboutS mm. long. Club-shaped

rio. 44. Zanclea ini[>lc\a, after Hartlaub, in Norillschcs rlankton. - ' .

to cylindrical, with about 50 short knobbed

tentacles, irregularly distributed. Hypostome short and flatly conical. Each polvpite IK i
i to 3 clusters of small medusa-buds. These are borne upon short branching stems which
arise between the tentacles near the lower, basal part of the polypite. The condition of the
medusa is not described. Found at Juan Fernandez Island, off the Pacific coast of South
America.

Genus ZANCLEOPSIS Hartlaub, 1907.
ZancleopaSf HARTLAUB, 1907, Nordisches Plankton, Nr. 12, pp. 115, 116.

GENERIC CHARACTERS.

Cladonemidae related to Zandca, but with large, stout, lateral branches on the tentacles.
With ectodermal ocelli upon the tentacle-bulbs, and without meridional clusters of nemato-
cysts upon the exumbrella.

The type species, and only known form, is Zancleopsis Jichotorna, from Tortugas, Florida,
described by Mayer under the name Gemimirni Jn-hotvma.

This genus is distinguished from Zatiflra by the large lateral branches upon its tentacles,
and by the absence of nettle-cell tracts upon the exumbrella. Also, it has <j> < Hi, and these
are not known in Y.nnclea.

Zancleopsis dichotoma Hartlaub.
Plate 8, fig. i.

Gtmmaria dichotoma, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 35, plate 17, fig. 40.
Zancleoftsis dichotoma, H^RTLArn, 1907, Nordisches Plankton, Nr. 12, p. 115, fig. 105.

Bell miter-shaped, with thin walls, and solid apical projection. 3 mm. high and about
2.5 mm. wide. There are 2 rudimentary tentacle-bulbs and 2 well-developed, diametrically
opposed tentacles. These long tentacles are of unequal length, one being about as long as

92 MEDUSA OF THE WORLD.

the bell-height, while the other is only about two-thirds this length. Each of these tentacles
terminates in a club-shaped, nematocyst-beanng end. The small tentacle gives rise to 2 to 3
side branches while the large one gives rise to 4 side branches. These side branches arise
from the outer (aboral) side of the tentacle, and each one terminates in a knob-like end con-
taining nematocysts as in the end of the main shaft itself. The youngest and least-developed
side branch is always found nearest the side of the bell, whereas the oldest is nearest the outer
end of the tentacle. The basal bulbs of the tentacles are large and swollen and contain each
an ectodermal ocellus upon the outer (abaxial) side of the bulb. The ectodermal core of each
tentacle and of the side branches is hollow. The nematocyst bulbs of the tentacles are
provided with delicate bristles. The velum is quite narrow. There are 4 straight, simple
radial-canals and a narrow circular vessel. The manubrium is flask-shaped, and cruciform in
cross-section, with 4 small, cruciform, nematocyst-covered lips. The stomach-cavity does
not project upward into the gelatinous apex of the bell. The gonads are developed upon
the interradial sides of the stomach.

The entoderm of the manubrium and the circular and radial-canals is dull ocher-yellow,
while the entodermal cores of the terminal bulbs of the tentacles is of a brighter yellow.

The 4 ocelli (2 on the bulbs of the well-developed tentacles and 2 on the rudimentary
tentacle-bulbs) are dark reddish-brown.

This medusa is occasionally found at Tortugas, Florida, from May until July. It is
taken each year upon the surface, but never in large numbers.

This species is widely separated from other medusae of the genus Gemmana by its well-
developed ectodermal ocelli upon the 4 tentacle-bulbs and by the total absence of nematocyst
tracts upon the exumbrella. The "filaments" arising from the abaxial sides of the 2 long
tentacles are almost as thick as the main shaft of the tentacle itselt, and in comparison with
species of Gemmaria they are greatly reduced in number. This numerical disparity is, how-
ever, counterbalanced by their great size.

Genus PTERONEMA Haeckel, 1879.

Microstoma, preoccupied by CUVIER, 1817, for Fishes.

Microstoma, LESSON, 1829, Voyage autour du monde sur la Coquille, tome 2, Zooph., p. 130.

Pteronema, HAECKEL, 1879, Syst. der Medusen, p. 101. VANHOFFEN, 1891, Zool. Anzeiger, Jahrg. 14, p. 445.

The first medusa to be described which belongs to this genus is Microstoma ambigua
Lesson, from New Guinea.

Haeckel, 1879, describes that which is probably the same medusa, in an expanded state,
under the name Pteronema darwinii. This species is described from the coast of Australia,
but the exact locality is not stated.

GENERIC CHARACTERS.

Similar to Zanclea, but distinguished by having a brood-sac above the stomach, the
nature of which is unknown. If it be similar to that of Eleutlieria it has no connection with
the stomach. Haeckel considers it to be continuous with the stomach, but his studies do
not demonstrate this to be true, for he cut no sections.

There are no meridional tracts of nematocysts upon the exumbrella in Pteronema.

Pteronema darwinii Haeckel.

( ? ?) Microsioma ambigua, LESSON, 1830, Voyage de la Coquille, Zooph., tome 2, p. 130, plate 14, figs. 5, 5'.

(? ?) Microslotna ambiguus, LESSON, 1843, Hist. Zooph. Acal., p. 295.

(? ?) Zanclea ambigua, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 344.

Pteronema darwinii, HAECKEL, 1879, Syst. der Medusen, p. IOI, taf. 7, figs. I, 2.

(? ?) Pieronema ambiguum, BEDOT, 1905, Revue Suisse de Zool., tome 13, p. 144 (literature, 1830-1850).

Haeckel describes his Pteronema darwinii from a single specimen preserved in glycerin.
He states that the preservation was good, but that the tentacles were contracted and are
represented in his figure as being of double the length observed in the preserved specimen.
This medusa was obtained somewhere ofF the coast of Australia.

Lesson, 1830, describes a medusa under the name Microstoma ambigua from Waigion,
New Guinea, which may possibly be identical with P. darwinii. Lesson's description is,

ANTHOMEDUS.E FTERONEMA, ELEUTHEHIA.

93

however, too vague to be of service, and his figure 5' is almost equally hopeless. The spherical
bell is apparently turned inside out, and there are 4 short, feathered, marginal tentacles and
a flaring, conical manubrium, the tentacles and manubrium being bright yellow. His figure
5 is even worse for purposes of identification. I advise the dropping of Lesson's medusa
from future lists, for it will doubtless be impossible to determine it even generically.

In Haeckel's medusa the bell is 6 mm. high, 4 mm. wide, pyriform with a pointed

conical apex. There are no longitudinal lines of nemato-
cysts over the exumbrella. There are 4 radially placed
marginal tentacles, longer than the bell-height, and each
provided with an abaxial row of filiform side branches,
each one of which terminates in a spear-head-shaped
capsule containing 5 to 10 nettle-cells. The basal bulbs
of the tentacles are conical and lack ocelli. There are 4
straight, wide, jagged edged radial-canals and a narrower,
smooth edged circular vessel. The stomach is spindle-
shaped, halt to two-thirds as long as the depth of the
bell-cavity, and provided with 4 simple lips. Gonads ( ?)
on the adradial sides of the stomach.

The distinguishing characteristic of this medusa is
the elongate, spindle-shaped brood-pouch above the
stomach in the gelatinous apex of the bell. Haeckel
observed ripe eggs and gastrulae in this brood-pouch.
The anatomical relationships between this brood-pouch
and the stomach are unknown, and it may have no con-
nection with the stomach, but be ectodermal and con-
nected with the subumbrella epithelium as it is in
Eleutheria, Haeckel's description affords no solution of
this problem. Color ( ?) Coast of Australia. Exact local-
ity unknown. A single specimen described by Haeckel.

Genus ELEUTHERIA Quatrefages, 1842.

Elcufhcriti, (Ji'ATRKFAr.Es, 1842, Compt. rend. Acad. Set., Paris, tome 15, p.
168; 1842, Annal. des Sci. Nat., serie 2, tome 18, p. 270.

Clavatella (hydroid and medusa), HINCKS, 1861, Annals ami Mai:. Nat. Hist.,
scr. 3, vol. 7, p. 73.

Elfutheria, KROHN, 1861, Wiegemann's Archiv. fur Natures., Jalirj;. 2-, Bd.
i, p. 157. CLAPAREDE, 1863, Beobacht. iiber wirbrllose ThiYre, p. 4.
ALLMAN, 1872. Monog. Tubul. Hydroids, p. 384. HAECKEL, 1879,
Syst. der Medusen, p. io<;. HARTI-AVK, 1886, Zool. Anzeiger, Bd. 9,
p. 707; Ibid., 1887, Bd. 10, p. 652; Iliiil., Bd. 12, p. 665, 1889; 1907,
NorJisches Plankton, Nr. 12, p. 126. BROWNE, 1902, Annals and
Mag. Nat. Hist., ser. 7, vol. 9, p. 279. GUNTHKR, 1903, Mitt. Zool.
Sta. Neapel, Bd. 16, p. 57. KRVMBACH, 1907, Zool. Anzeiger, Bd.
3 1, p. 450.

FIG. 45. Ptcronema danvinii, after
Haeckel, 1879.

The type species of this genus is the extremely v.i li-
able Elfutheria dichotoma of the Mediterranean and
Atlantic coasts of Europe. Haeckel, 1879, p. 106, records 12 subspecies of this medusa. Tin-
best description is by Hartlaub, 1886, 1907.

GKNKRIC CHARACTERS.

Cladonemidae with 4 or more simple radial-canals and an equal number <>t biturcated
tentacles. Terminal branches ot tentacles end each in a knob-like. cluster of nemntocysts.
Manubrium a simple, 4 or more sided tube without oral tentacles or prominent lips. Velum
well developed and there is an urticating ridge around the exumbrella side of the hell-margin
below the ring-canal. There is a peculiar brood-pouch above the stomach, but this pouch is
not connected with the gastrovascular cavity of the medusa. The cavity of this brood-pouch is,
however, connected with the bill-cavity by means of simple, interradial openings. The genital
products are developed exclusively in the epithelial lining of this brood-pouch, which is dei i\ i-d

94

MEDUS.E OF THE WORLD.

from the ectoderm of the subumbrella cavity of the bell. The medusa is hermaphroditic; ova
develop in the ventral, and sperm in the dorsal (aboral) wall of the brood-pouch.

The hydroid is Clavatella of Hincks, 1861 (Annals and Mag. Nat. Hist., ser. 3, p. 73,
plates 7, 8; Ibid., 1868, British Hydroids, vol. I, p. 73, plate 12, fig. 2). In Clavatella the
polypites arise singly from a linear stolon. The polypites are small linear, or clavate, with
a zone of 8 slender tentacles which terminate each in a knob of nematocysts. The medusa-buds
are borne only on two opposite sides of the body of the polypite near its base. The hydroid
lives on Ulva in shallow tide pools, and the medusae are produced in summer and autumn.

Hartlaub found that the entoderm of the planula of Eleuthe ria contains numerous nema-
tocyst-cells. Krumbach, 1907, believes that Tnchoplax, F. E. Schulze, 1891 (Abhandl. Akad.
Berlin), is the creeping planula of Eleuthena.

Elfuthcna, Cladonema, and Amphogona are the only genera of hydromedusae known to
be hermaphroditic.

Fit;. 46. Hydroid (Clavatella prolifera) and its medusa (Eleutheria dichotoma), after Allman, in Ray Society, 1871, 1872.
FIG. 47. Clavatella prolifera, after Hincks, in British Hydroid Zoophytes.

Eleutheria dichotoma Quatrefages.

Eleulheria dichotoma, QUATREFAGES, 1841, Compt. rend. Acad. Sci., Paris, tome 15, p. 1 68; 1842, Annales Sci.Nat., Ser. 2, p. 270,

plate 18. KROHN, 1861, Archiv. Naturgesch., Jahrg. 27, p. 157.
Clavatella prolifera (hydroid), HINCKS, 1861, Annals and Mag. Nat. Hist., ser. 3, vol. 7, p. 73, plates 7, 8; 1868, Hist. British

Hydroid Zoophytes, p. 73, plate 12, fig. 2.

Non Eleutheria dichotoma, CLAPAREDE, 1863, Beob. Anat. und Enrwick. wirbelloser Thiere, p. 4, taf. i, fign. 4-10.
Clavatella prolifera (hydroid), ALLMAN, 1872, Monog. Tubul. Hydroids, pp. 31, 212, 384, plate 18.
Eleutheria dichotoma (non claparede), HAECKEL, 1879, Syst. der Medusen, p. 106.
Eleutheria dichotoma, GRAEFFE, 1884, Arbeit, Zool. Inst. Wien, Bd. 5, p. 353. HARTLAUB, 1886, Zool. Anzeiger, Bd. 9, p. 706,

I fig. Also: Ibid., 1887, Bd. 10, p. 652; 1907, Nordisches Plankton, Nr. 12, p. 127, fign. 1 19, 120. BEDOT, 1905, Revue

Suisse de Zool., tome 13, p. 133 (literature cited to 1850). MULLER, 1908, Zeit. fijr wissen. Zool., Bd. 89, pp. 34, 73,

taf. 3, fign. 3-7 (origin and structure of eggs).

This medusa is exceedingly variable, but the normal form may be briefly described as
follows :

Bell irregularly hemispherical, with its lower surface more or less 6-sided. 0.3 to 0.4 mm.
wide. There is a ridge of large nettling warts extending around the margin. There are 6 ten-
tacles at the ends of the 6 short radial-canals, and not irregularly arranged in reference to the
radial-canals as in E. claparedii Hartlaub = E. dichotoma Claparede.

A. \THOM Kill S.K KI.KI TIIKK1A.

95

Each tentacle bifurcates and is twice as long as the bell-diameter. The 2 terminal
branches end one in an adhesive disk, and the other in a large, knob-like cluster of nettle-cells.
These branches are not quite as long as the basal shaft of the tentacle itself. An abaxial
ocellus is found at the base of each tentacle.

The medusa is hermaphroditic, and the sexual products arc developed in an ectodermal
brood-pouch above, but not connected with, the stomach. The cavity ol the brood-pouch is
connected with the bell-cavity by means ot 6 simple, interradial openings which alternate with
the 6 radial-canals. Sperm develops in the aboral, and ova in the oral ( lower) \\all of the
brood-pouch.

Medusa-buds are also produced upon the exumbrella side of the ring-canal, and this
process is usually associated with the sexual reproduction. The terminal suckers of the ten-
tacles are orange or yellowish-brown. Stomach and canals yellowish.

This medusa is found clinging to green sea-weeds oft the Atlantic coasts of Belgium,
England, and France, and is widely distributed in the Mediterranean. Graefte. 1884, found
it to be common upon Ulvu from July to September at Trieste, Adriatic Sea. The budded
medusse become sexually mature in 3 to 4 weeks. The planula larva creeps over the sea-
weeds.

Detailed descriptions are given by Ouatrefages and Allman, and the best modern account
is that of Hartlaub, who discovered the remarkable character of the brood-pouch or ecto-
dermal gonad.

FIG. 48. Eletithfriti difholiwa, after Hartlaub, in Xordischcs Plankton.

The hydroid is Clavatella prolifera Hincks (see definition of the genus Eleutheria).

Eleutheria dichotoma is apparently quite variable. The tentacles and radial-canals, al-
though normally 6, may range from 4 to 8. Haeckel, 1879, p. 106, designates these aberrations
as "subspecies, " and gives specific names to twelve of them, although a number of these
should he credited to Eleutheria claparcJu Hartlaub and its variations.

Hartlaub finds that the ova develop into planula larvae within the brood-sac before being
set tree.

Miiller, 1908, studied the origin and structure of the eggs of this medusa. The eggs are
small, numerous, and oval in outline, not amcrhoid. There is no visible distinction between
exoplasm and endoplasm. The ooplasma is a network of very fine fibers. There are num-
erous small yolk-granules.

Eleutheria claparedii Hartlaub.

Eleuthrria elafaredii, HARTI.AI-K, 1889, Zonl. Anzeiger, Bil. 12, p. 665; 1907, Noriitscln^ Pl.mkt"ii. Nr. 12, p. 129. fii;. 121.
EIrullifria dichotoma, CLAPAREDF., 1863, Bcobacht. Anat. und Entwicklungsges, \virhrllosrr Thicrr, p. 4. t.if. I, figs. 4-10.
SPAGNOLINI, 1876, Catalogo Acalefi Mediterraneo, p. 24, tav. 4, fig. 2.

Bell 0.4 to 0.5 mm. wide, irregular in shape, usually more or less hemispherical (fig. 41; i.
A ring of nettling warts on bell-margin. 8 to 10 tentacles, irregularly arranged in reference
to the 4 to 6 radial-canals. A small abaxial ocellus at the base of each tentacle. The ten-
tacles are 3.5 times as long as the bell-diameter, and they bifurcate at their extremities; one
branch ends in an adhesive disk, and the other in a nettling knob. About (> or ~ medusa-buds
are produced upon the subumbrella side of the ring-canal and project into the bell-cavity.

96

OF THE WORLD.

This process of budding; is independent of the sexual reproduction. The terminal suckers ot the
tentacles are orange. Found on Ulva in the Bay of Naples. Described in detail by Hartlaub.
The structure of the reproductive sac is discussed in describing the character of the genus

Eleutheria.

Eleutheria vallentini Brown.

Eleutheria vallentini, BROWNE, 1901, Annals and Mag. Nat. Hist., scr. 7, vol. 9, p. 279.

Umbrella hemispherical, 3 mm. wide, 2 mm. high. 24 tentacles divided into two branches,
the upper branch with clusters of nematocysts, the lower with a terminal sucker. "An ocellus
on the extreme margin ot the umbrella opposite each tentacle." Stomach-tube conical and
small. Mouth a plain round opening without lips. The gonads occupy "the whole of the
upper part of the umbrella above the stomach." A single specimen was found by Vallentin at

Fii;. 49. Eleutheria claparetlii, after Hartlaub, in Nordisches Plankton.

Stanley Harbor, Falkland Islands, and described by Browne. Color ( ?) Number of radial-
canals ( ?) Brood-pouch ( ?) In the absence of a figure or a more detailed description it will
probably be impossible to redetermine this form unless it be rediscovered in Stanley Harbor.

Genus MNESTRA Krohn, 1853.
Mnestra, KROHN, 1853, Archiv. fiir Naturges., Jahrg. 19, p. 278. GUNTHER, 1903, Mittheil. Zool. Sta. Neapel., Bd. 16, p. 35.

The type species is Mnestra parasites Krohn, from the Mediterranean.

GENERIC CHARACTERS.

Cladonemidae with 4 radial-canals and a ring-canal. No brood-sac above the stomach.
4 to o degenerate, hollow tentacles with a row of netting capsules along their aboral sides.
\\ ith a ring of nettling cells around the margin and 4 linear tracts of nematocysts over the

ANTIKiMI.IM S.K MM.STKA.

exumbrella, one above each of the 4 tentacle-bulbs. The throat of the medusa is blocked by
a spongy mass of entoderm. There is a cup-like depression in the center of the exumbrella.

This medusa attaches itself by its suctorial mouth to the throat of the opisthobranch
mollusk, Phyllirhoe. It may lie derived from some Zanclea-like form \\bich has become
degenerate through its sessile habits. It can not swim, yet it has well-developed circular
muscles in the subumbrella and a distinct velum.

Mnestra parasites Krohn.

Mnf*i>, i /Hi'iifitef, KRIUIN, 1X53, Arclnv. Natures., Jalirg. 19, p. 178. CLAIS, 1875, Vi-rli.indl. /.mil. Buun. Gi-M-ll., \Vu-n.
Bd. 25, p. 9, taf. i. HAF.C'KEL, 1880, Syst. dor MrduM-n, p. 653. GUNTHKK, 1903, Mmhnl. Znol. Sta. Ni-a|vl, lid. id.

p. 35, plates l, 3, 42 figs.
Mncstrii parasitica, FEWKES, 1884, American Naturalist, vol. iS, p. 197, figs. 4, 5.

5'

Fid. 50.- Mnestra parasites, from Florida, after Kc\\kt*s, in Aincr. Natu-
ralist, 1884.

Fn.. 51. Mnestra parasites^ after Giinthrr, in Mitth. Zool. Sta. Nc.ip
Bell turned inside out, showing iiunit]i-|>.irt -, and cauti
exumbrella to be cup-shaped. Medusa hat. 3 \vcll-lc\r1f>pi-<l
tentacles.

FIG. 52. Ctenaria ctenophora, after Hacckc], 1879.

This minute, degenerate, highly variable medusa is parasitic or commensal upon the
mollusk Phvilit'hrjf. Its germ cells appear to migrate into the body of the PAy///rAo where
they develop.

98 MEDrS.E (IF THE WORLD.

It has been rescued from scientific oblivion through an able paper by Giinther, 1903, who
has carried out an elaborate study of its anatomy, life-history, and variations.

The characters of the medusa are those of the genus of which it is the sole representative
(see genus Mnestra).

Fewkes, 1884, describes this medusa upon Ph\lhrhoc on the Florida Reef. It is com-
mon at Naples and Messina, Mediterranean.

Genus CTENARIA Haeckel, 1879.

Clenaria, HAECKEL, 1X79, Syst. der Medusen, p. 107. GUNTHER, 1903, Mittheil. Zool. Sta. Nrapel, Bd. 16. p. 57. HARTLAUB,
1907, Nordisches Plankton, Nr. 12, p. 24.

The type species is Ctenaria ctenophora Haeckel, from the coast of japan. This is the
sole representative of the genus, and Haeckel had but a single preserved specimen.

GENERIC CHARACTERS.

Cladonemidae with 4 bifurcated radial-canals. With 2 feathered marginal tentacles and
simple, unbranched, oral tentacles. There are 8 adradial, meridional lines of nematocysts
over the exumbrella, and a nematocyst tract above the base of each marginal tentacle.
Gonads on the sides of the stomach. There is an apical cavity above the stomach, but it is
not known whether this is an ectodermal brood-pouch, as in Eleutheria, or a mere extension of

the stomach-cavity.

Ctenaria ctenophora Haeckel.

Ctenaria cttnttphora, HAECKEL, 1879, Syst. der Medusen, p. 108, taf. 7, figs. 5-7. HARTLAI-B, 1907, Nordisches Plankton,
Nr. 12, p. 126, fig. 119.

Bell three-fourths-egg-shaped; 6 mm. high, 5 mm. wide. (Fig. 52.) Widest in a zone
slightly below the middle. 8 adradial lines of nettling cells extend up the sides of the
exumbrella from the margin nearly to the apex, and in addition to these there are 1 large
club-shaped clusters of nematocysts extending from the bases of the 2 tentacles half-way up
the sides of exumbrella.

The tentacles are each 2 to 4 times as long as the bell-height and they taper gradually
from base to tip. There are no basal ocelli. A row of numerous, slender filaments arises from
the abaxial side of each tentacle. These filaments superficially resemble those of Ctenophorae,
and do not end in nematocyst capsules as in Zanclea. They are, however, covered with nema-
tocysts throughout their lengths.

The velum is well developed. 4 main radial-canals arise from the stomach, but these
bifurcate so that 8 canals join the marginal circular vessel in the 8 adradii. The edges of these
radial-canals are serrated, being apparently beset with glands.

The stomach is almost spherical and about half as long as the depth of the bell-cavity.
There are 16 short, simple, oral tentacles, each of which ends in a nematocyst-knob.

Haeckel describes 4 swollen, radially placed gonads on the sides of the stomach, but
these are probably adradial or interradial ( '.) There is a well-developed apical cavity above
the stomach, but it is not quite clear whether this is a brood-sac homologous with that of
Eleutheria or whether it is simply an apical extension of the stomach. Haeckel supports the
latter view, but he made no sections, and the point can not be considered as determined.
Color ( ?)

Haeckel describes this medusa from a single alcoholic specimen from Japan.

Genus CLADONEMA Dujardin, 1843.

Cladoncma, Dl'jARDlN, 1843, Anna!, des Sci. Naturelles, tome 20, p. 370. KROHN, 1853, Muller's Archiv. fur Anatomie und
Physiologic, p. 420. GEGENBACR, 1856, Zeit. fur wissen. Zoo]., Bd. 8, p. 230. VAN BENEDEN, 1866, Fauna Littor.
Belgique, p. 139. HINCKS, 1868, British Hydroid Zoophytes, p. 62. ALLMAN, 1871-72, Monograph Tubularian
Hydroids, pp. 216, 357. HAECKEL, 1879, Syst. der Medusen, p. 109. PERKINS, 1902, Johns Hopkins University Circu-
lars, No. 155, p. 21;, 1908, Papers from Tortugas Laboratory Carnegie Institution of Washington, vol. I, p. 136.
HARTLAI-B, 1887, Zool. Anzeiger, Bd. 10, p. 654; 1907, Nordisches Plankton, Nr. 12, p. 131. GUNTHER, 1903,
Mitt. Zool. Sta. Neapel, Bd. 16, p. ^7.

Stauridium cladontma (hydroid), or PLESSIS, 1888, Recueil Zool. Suisse, tome 4, p. 536.

GENERIC CHARACTERS.

Cladonemidae with 4 or 5 or more bifurcated radial-canals or 8 to 10 or more simple
canals, or with some bifurcated and some simple canals. With 8 to 10 or more tentacles which

AXTHOMKIH S.K- ( I.AIHINI.MA. (I'.l

give rise to sucker-bearing or nematocyst-bearing branches or both. With 4 or 5 simple, oral
tentacles which terminate in nematocyst-knobs. There is no brood-sac above the stomach.
The genital products develop within the entoderm of the stomach, and when mature are
found in the ectoderm. The medusa is sometimes observed to be hermaphroditic.

The type species is ClaJonctna radiatum of the coasts of Europe. According to Haeckel
this species is highly variable both in form and color (see Haeckel, 1879, p. 109). The two
American forms described by Perkins are, however, quite constant in their form, although
one of them is somewhat variable in color. The hydroid stock of ('.laJuni-nm belongs to the
tubularian genus StauriJia of Dujardin. It is well described by Allman, 1871 (Tubularian
Hydroids, pp. 216, 357, plate xvn), and by Perkins, 1908.

Cladonema radiatum Dujardin.

Cladonema radiatum, Dt'jARDiN, 1843, Compt. rend. Acad. Sci., p. 1134.

Siauridie (hydroiil), DUJARDIN, Ibid., p. 1133.

C.ladonema radiatum and Stauridie, DIIJARIMN, 1X43, Annal. des Sci. Nat., tome 20, p. 370; 1845, Ibid., ser. 2, tome 4. pp. 271
272, plate 14, fig. C.

C.ladontma radiatum, KROHN, 1853, Muller's Archiv. fur Anat. und Physiol., p. 420, plate 13. KKFKRSTFIN i ND EHI. F, RS, 1861 .
Zoologische Beitrage Neapel, Messina, p. 85, taf. 13, fig. 5. VAN BENEDEN, 1866, Mem. Acad. Roy. Belgnjue, tome 36,
p. i 39, plate 12. HINCKS, 1868, Hist. British Hydroid Zooph., p. 62, plate 1 1 . ALLMAN, 1872, Monog. Tubul. Hydroids,
pp. 216, 357, plate 17, figs. l-io. HAECKEL, 1879, Syst. der Medusen, p. 109. WEI s MANN, 1883, Entstehung Sexual/ ellen,
bei Hydromedusen, Jena, pp. 1 19, 218, taf. 12, figs. 2-5. JICKELI, 1883, Morphol. Jahrbuch, Bd. 8, p. 602, taf. 26 (his-
tology). HARTLAIIB, 1887, Zool. Anzeiger, Bd. 10, p. 655, I fig.; 1907, Nordisches Plankton, Nr. 12, p. 132, fign. 123-125,
(list of authors; excellent description of the medusa). BEDOT, 1905, Revue Suisse de Zool., tome 13, pp. 58, 132 (refer-
ences to 1850). BILLARD, 1905, Bull. Museum d'Hist. Nat. Pans, tome ll, p. 500 (variations). --Mi'LLi R, 1908, Zrit.
fur wissen. Zool., Bd. 89, pp. 30, 73, taf. 3, figs, i, 2; taf. 4, figs. 8-1 1 .

Bell half-egg-shaped or globular and 2 to 3 mm. wide. At Naples, Italy, the medusa
appears to be quite invariable, and according to Haitlaub, 1887, the manubnuin is spindle-
shaped and five-sided with 5 perradial oral tentacles and 5 perradial sac-like outgrowths
upon the sides of the stomach. Thus in the Naples medusae 5 main radial-canals, 72 apart,
arise from the stomach, but 3 of these bifurcate so that 8 canals, 45 apart, join the circular
vessel at the margin. These 5 main radial-canals are arranged as follows: 2 bifurcated
canals 72 apart; 2 simple, unbranched canals 144 apart; i bifurcated radial-canal mid-
way between the two simple canals, i. e., 72 from each. Thus in glancing around the
margin in the direction of the order of succession of the hour-numbers on the dial of a clock,
we may begin with a simple canal, then comes a bifurcated, then a simple, next a bifurcated,
and finally another bifurcated canal. 2 of the bifurcated canals are thus side by side and
the 2 simple canals are separated one from another by an interval occupied by the third
bifurcated canal.

There are 8 tentacles upon the Naples medusae, one at the end of each terminal radial-
canal. The basal bulbs of these tentacles are heavy and swollen, and each bears an abaxial
ocellus, i to 3 sucker-bearing filaments arise from the inner sides of the tentacles and their
outer parts terminate in 4 to 6 branches which are armed with nettling warts and end in
knobs.

The gonad encircles the stomach and is not confined to the sac-like protrusions, as was
believed to be the case by Haeckel. According to Hartlaub, the genital products originate
in the entoderm, but when mature are found in the ectoderm of the manubrium; yet according
to Weismann they originate and remain in the ectoderm.

The medusa exhibits a successive hermaphroditism, according to Hartlaub, although
either sex may precede in the process. (See also Miiller, 1908.)

The stomach, ring-canal, and tentacles are red to brown.

This medusa is abundant oft" the Atlantic coasts of England, Holland, and V ranee, and
in the Mediterranean. Haeckel records a number of departures from the normal form.
These are probably only aberrations, but he gives specific names to each and every one of
them. For example, the oral tentacles and the sac-like outgrowths on the stomach may range
from 4 to 5; and there may be 10 simple, or 4 bifurcated radial-canals.

There appear indeed to be many local races of this medusa, and I am inclined to believe
that Cladonema perkinsii and (<. nmyen of America may prove to be only varieties of C.
radintuin.

100

MK.WS.E OF THK WORLD.

Billard has studied the variations of the medusa at the Bay de la Hougue near St. Vaast
on the northwestern coast of Fiance. Here he found that among 50 medusae 5 had 6 radial-
canals, 4 had 7, 39 had 8, I had 9, o had 10, I had u. In the case of the 39 medusae with
8 radial-canals, 36 had 2 simple and 3 bifurcated canals, as in the Naples medusa.

Miillei, 1908, has studied the origin and structure of the ova in this medusa. The eggs
are small and rounded and widely scattered in the gonad, for the successful eggs devour
the weaker in the ovary. The ooplasma is sparsely vacuolated and there is very little yolk.
53- 55-

54-

";; .- Hydroid of ClaJonema raiiiatum, after Hincks, in British Hydroid Zoophytes.
Showing growth-habit of hydroid and enlarged view of single polypite.
Fir.. 54. ClaJonema raJiatum, after Allman, in Ray Society, 1871-72. Budding

hydroid anil mature me lusa.
Fi<;. 55. Medusa of C.lailontma rnJiatum, after Hincks, in British Hydroid Zoophytes.

The hydroid of Clndoncinti radiattitn isStaurn/ia raJiatum Dujardin = CAndonema radiatum
Hincks.

The hydrorhiza is a slender, creeping filament from which the hydranths arise at inter-
vals. These hydranths are either simple, unbranched, and about 2.5 mm. high, or are borne
upon slightly branching stems 12 to 25 mm. high. The hydranths are club-shaped and
have 2 alternating verticils, each circlet consisting of 4 tentacles. The circlet near the mouth
is composed of long tentacles, each of which ends in a knob, while the circlet at the base of
the hvdranth consists of 4 very short, simple, unbranched, filiform tentacles which alternate
with (i. e. are upon meridians 45 apart from) the oral circlet. The medusae bud out singly
from the sides of the polypite at a short distance above the basal circlet of tentacles. The

\\Tim\ii.Di s i; CI..\I>H\I:M \. 101

body of" the hydranth Is pale reddish and the perisarc bright yellowish-brown. The medusae
are produced in spring and summer.

Dujardin's, 1843, observations upon Cladonema niJintiun are the earlit-st in which the
complete life-history ot' the alternation of generations between hydroid and medusa was
actually observed.

Cladonema perkinsii Mayer.
Plate 9, fig. i .

C.lailonema, sp., PERKINS, 1902, Johns Hopkins Uimersitv Circulars, vol. n. No. 1^5. p. 2^, ne,^. in ti-it.

Cladonema per kin sii, MAVFR, 1904, Memoir- N.it. Sci. Brooklyn Inst. Arts ami SM .. vol. i , XL. i . p. i X, plate 4, fig. 35. H*KT-
LAi'B, 1907, Nonlisches Plankton, Nr. 12, p. i ;^, tit:, it>.

Bell half-egg-shaped; less than 2 mm. in diameter, with thin, uniform walls. 8 lar<;e,
stout, marginal tentacles. The inner and lateral parts of each of these tentacles bear 3 to 10
small, flexible cirri, which are besprinkled with wait-like clusters of nematocysts and termi-
nate in a knob. 1 here is a large, cup-like, ectodermal ocellus upon the outer side of each
main tentacle shaft near the bell-margin. The velum is large and shows circular striations.
Manubrium large and spindle-shaped, with about n rounded, protruding pouches at its widest
part. The mouth is surrounded by a circlet of 5 simple, short oral tentacles each terminating
in a knob-like mass of nematocysts. 8 simple radial-canals, 45 apart, arise hum the stomach,
and extend straight unvard the simple circular vessel. The manuhrium is thus 8-sided at
its proximal end, 6-sided in the middle, and 5-sided at the mouth. The genital products
develop in the ectoderm ot a large part ot the manubriurn and also in the hernia-like pouches.
The ocelli are black and all other parts colorless. This species was discovered by Professor
Perkins in Nassau Harbor, Bahamas, in July, 1902, upon the surface in shallow water at
night.

Cladonema mayeri Perkins.
Plate 9, figs. 2 an, I -, .

Cladonema, .</>., FKWKES, iSS^, Bull. Mus. Comp. Zool. at Harvard College, vol. 11, p. 87.

Cladonema ma\eri, PKRKINS, 1906, Year Book of tile Carnegie Institution of Washington, No. 4, IIJCK. p. llS. KjoS, Papers
from Tort u gas I. ah. Carnegie In ^t. Washington, \ol. i , p. i ;(>, plate- i an, I 2. plate 4, figs. 2 t , 22. H\ ,!roul an, I medusa.

Bell thin-walled, higher than a hemisphere, and with small, solid, apical projection;
top rounded and dome-like, and total height about 2.5 mm. There are 9 marginal tentacles,
one at the toot of each radial-canal. Each ot these tentacles has a large, spindle-shaped basal
bulb, the entoderm of which contains a mass of white concretions. On the outer side of
each basal bulb near the circular canal there is a deep reddish-brown, ectodermal pigment
spot. The distal, inner side of each basal bulb gives rise to about 6 small tapering peduncles
which terminate in small knob-like adhesive disks. These enable the medusa to cling to the
sides or bottom of the aquarium. The main shaft of each tentacle extends outward from the
spindle-shaped end of the basal bulb. It is umtorm, thread-like, and slender, and gives rise
to 4 to 8 thread-like side branches each ot which, together with the central shaft, terminates
in a knob-like cluster ot nematocvsts. There are 4 to 6 nematocyst-warts upon each side
branch and a greater number upon the main shaft. Both the main shaft and the side brandies
are highly contractile and can be expanded so as to become longer than the bell-height, or
contracted into a close bunch. The velum is very wide and its orifice small. 6 radial-canals
arise from the stomach, but every alternate canal bifurcates near its point ot origin, and
thus () equally spaced canals reach the circular vessel. The manubrium normal!) extends
to the level of the velar opening, but may contract somewhat at times, as is shown in our
figure. The mouth is surrounded by <> simple oral tentacles, each of which terminates in
a large nematocyst-knob. Near the middle of the stomach there is a circlet of 6 radially
arranged, short, blunt, hernia-like projections. The genital products are developed in the
ectoderm of the walls of the manubrium.

The color is quite variable. The entoderm ot the basal bulbs ot tin- marginal tentacles
is milky-yellow streaked with dark-brown pigment-granules, while the entoderm of the
manubrium is milky-yellow or dull-milky ocher, streaked longitudinally in each of the ft
radii by dark-brown, almost black, pigment-granules. The disposition and arrangement ot
this pigment is highly variable. Perkins, iijoX, nives the most complete account ot this
medusa and its hydroid.

102 MF.nrs.K UK TIIK WORLD.

This medusa, together with its hydroid stage, was found in great numbers by Prof.
Henry F. Perkins in the salt-water moat of Fort Jefferson, Tortugas, during the summer of
1905. The hydroid is a minute Stauridiu which grows upon algae. The same medusa was
described by Fewkes, 1883, as being in association with Cassiopea at Fleming's Key, near
Key West, Florida.

Genus DENDRONEMA Haeckel, 1879.

Dtmlronema, HAFCKEL, 1879, Syst. der Metlusen, p. MO. GUNTHER, 1903, Mitth. Zool. Sta. Neapel, Bil. 16, p. 57.

The only known form is Dendronema styludendron Haeckel, from the Canary Islands,
Atlantic Ocean.

GENERIC CHARACTERS.

Cladonemidae with branched oral tentacles and branched marginal tentacles, the
branches ending in nematocyst-knobs or adhesive disks, or both. With bifurcated radial-
canals. Gonads in the stomach-wall.

There is an apical cavity above the stomach, but we do not know whether this is a mere
extension of the stomach itself, or a reproductive sac similar in anatomy to that of Eleutkena.

Dendronema stylodendron Haeckel.

])?rnlionetiiti st \lrjiletidT on, HAVCKKL, 1879, Syst. iler Medusen, p. no, taf. 7, fi^. 8.

Bell miter-shaped, with pointed conical apex. 9 mm. high, 6 mm. wide. Stomach spindle-
shaped. There are apparently 4 radially placed oral tentacles each of which branches dichot-
omously 6 to 7 times and terminates in (50 to 60) nematocyst-knobs in each quadrant.
Haeckel's description is, however, vague upon this point. Haeckel states that there are 4 egg-
shaped "gonads" on the 4 perradial sides of the stomach. These may, however, be homol-
ogous with the protuberances seen in Cladonema ( ?) HaeckePs figure shows them distended
with ova. 4 principal radial-canals, 90 apart, leave the stomach and bifurcate so that 8
vessels reach the ring-canal 45 apart. There are 8 marginal tentacles, one at the base of each
radial-canal. There is an ectodermal abaxial ocellus at the base of each tentacle. Each ten-
tacle gives off a stout, short branch on its inner (velar) side, .and this branch bifurcates twice
and terminates in 4 nematocyst-bearing, knob-like "suckers," or adhesive organs. The main
shaft of the tentacle also branches dichotomously many times and is besprinkled with nettling
warts, and the ends terminate in large knobs. There is a spindle-shaped cavity in the apex
of the bell, above the stomach, but its anatomical character is unknown. The mouth, stomach,
apical cavity, and gonads are reddish-yellow. Tentacles, radial-canals, and ring-canal brown-
ish-red. Ocelli black.

This medusa was found by Haeckel in the Canary Islands, Atlantic Ocean, in February,
1867.

Family OCEANIDjE, sens. Vanhoffen, 1891.

OeeaniiLr (in part), ESCHSCHOLTZ, 1829, Syst. der Acalephcn, p. 96. FORBES, 1848, British Naked-eyed Medusa-, p. 21.

GEGENBAUR, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 219.

Tiarid.e+ Margt'lid l p + Cannotidtf (in part), HAECKEL, 1879, Syst. der Medusen, pp. 40, 68, 1^6-159.
Ocrnniiir, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 443.

FAMILY CHARACTERS.

Anthomedusa? with 4, or 4 pairs of, isolated gonads upon the interradial or adradial sides
of the stomach. The marginal tentacles may arise singly or in clusters, but are neither
branched nor feathered.

The Oceanidae constitute the third family of the Anthomedusae, and they are more com
plex and more highly differentiated than the simple and more primitive Codonidae.

AXTHOMKlll S.K -OCEANID.S. HIM

We distinguish three subfamilies ot" Oceanidae.

1. Tiarirue. Lips without oral tentacles. Simple unbranched radial-canals. Ten-

tacles arise separately from the hell-margin and are not grouped in clusters.

2. Margelinti'. With oral tentacles. Simple, unhranched radial-canals. In some

genera the marginal tentacles arise singly; in others they are grouped in clusters.

3. DenJrostaurtnte. Lips without oral tentacles. The radial-canals branch.

Direct development ot the medusa trom the egg is unknown in this tamilv. The following
genera are known to develop through Tubularian hydroids: Stomotoca, Turrn. Podocaryne,
Stylactis, Bougainvillia, Neinopsts, Lizzia, and ftillia.

Medusae are produced asexually upon the interradial sides of the manubrium in ('\tttis,
PoJocoryne, Bougainvillia, and Rathkea, In some species of Bougainvillia the eggs undergo
part of their development within the ectoderm of the parent medusa, and are discharged as
well-developed planulae. In Ntobia the tentacle-bulbs develop into new medusae, and are set
free one after another to repeat this process. In H illia and Prohoscidactyla, medusa-huds
are produced upon hollow stolons which may arise from the corners ot the stomach at the
points ot |uncture with the radial-canals, or from the places where the canals fork, as in P.
flavicirrata, var. stolonifera, Maas.

Haeckel, 1879, describes the gonads ot the Oceanidae as being penadial in position anil on
the sides of the stomach in the same sectors with the radial-canals, but Vanhoffen, 1891 (Zool.
Anzeiger, Bd. 14), showed that this is an error, for the gonads are almost universally adradial
or interradial in position and alternate with the sectors of the radial-canals.

The specialized conditions displayed by the medusae ot the Oceanidae, such as the corru-
gated, folded gonads ot the Tiannae, the clustered marginal tentacles and the oral tentacles of
the Margelmae, and the forked radial-canals and other peculiar structures in the Dendro-
staurinae.all indicate that the Oceanidae are derived from simpler forms, such as the Codonidae.
Indeed, the conditions seen in theOceanidae are largelytoreshadowed in the more differentiated
Codonidae and in the Cladonemidae. In common with the Codonidae and Cladonemidae
the Oceanidae are derived trom Tubularian hydroids.

Subfamily TIARINjE Haeckel, 1879.

(liemnd.r (in part), ESCHSC HOLTZ, 1829, Syst. tier Acalephen, p. <)(>. GK^NBACR, lNcf>, 7.ni. fiir uissrn. /mil., Hd. S,p. 219.

\mleiffrtF (in part), LKSSON, 1X45, Hist. Nat. Zooph. Acal., p. 28}.

Tjitridii-, HAF-CKRL, 1879, Syst. der Medusen, p. 40.

Sfns. reslr,, drlomrrinthiu, VANHO? UK, 1891, Xool. Anzeiger, Bd. 14, p. 44V

Tiiiriiiit, HARTLAUB, 1892, Nachrichten kgl. Gesell. Wissenschafl. Univ. Gottingen, pp. 19-22; 1907, Nordisihes 1'l.inkton,

Nr. 12, p. 5.

.SVfn. restr. t Tiurijcp, MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. iS, |>. ri.
Tiilrin.r, VON LENDENPELDj |NS4, X.uol. An/fiytT, Bd. 7, p. 446.

CHARACTERS OK THF. TIARIN/E.

Anthomedusae with 4 lips, without oral tentacles. With 4 or more unbranched radial-
canals and with simple, hollow, unhranched tentacles which are not grouped in clusters, but
arise separately from the bell-margin. With interradial or adradial gonads forming swollen,
corrugated regions in the walls of the stomach.

The following table shows the genera of the Tiarinae.

1 . Tnhc PROTIARIDI :

The outer surfaces of the four interradial gonads are smooth. Not corrugated or folded. There are 4 simple lips, 4 r.uii.d-
canals, and a ring-canal, 4 or more tentacles.

Proriara HAtcKEL, 1879. With 4 radial-canals and 4 radially situated marginal tentacles. 4 simple lips not com-
plexly folded. Hydroid unknown.

Hetrroliara MAAS, 1905. With 4 radial-canals, and 8 marginal tentacles. Ring-canal gives rise to interradial, blindly-
ending, centripetal canals. Only species is H. anon\mn, MAAS, 1905, Craspedoten Medusen Siboga Expedition,
Monog. lo, p. 19, taf. 3, figs. 19-21, Malay Archipelago. Hydroid unknown.

2. Tribe AMPHINEMIDI:

With 2 long and numerous rudimentary tentacles. There are 4 radial-canals.

Slomoloca L. AOASSIZ, 1862 = Amphincma + Stomoloca+ Codonorchis, HAICKII, 1^79. With two well-developed ten-
tacles. Manubrium mav or may not be mounted upon a peduncle. Gonads corrugated, folded, swollen regions upon
interradial or adradial sides of stomach.

Dissontma HAECKEL, 1879. With 2 well-developed and numerous rudimentary tentacles. During their development
gonads migrate outward from sides of stomach down the 4 radial-canals.

104 MKDT'S.K OF THE WOULD.

V Tribe PAND.IIDI:

With 4 or more well-developed tentacles. The gonails are interradial, corrugated, or folded ridges in the wall of the stomach

4 radial-canals.
Pundta LESSON, 1843 (1837?). 4 interradial gonads in stomach-wall, not completely separated in the 4 principal

radii. Hydntid unknown.
Conis BRANDT, 1838. Ocelli borne upon ends of special, short clubs which arise from bases of tentacles. Hydroid

unknown.

Turris LESSON, i^^=Tiara, LESSON -4- Catablema, HAECKEL. With 4 interradial crescent-shaped gonads in the ecto-
derm of the stomach-wall. Each crescent is composed of partially fused, swollen ridges. Hydroid: Clavttla ( ?),
Carnpaniclava ; or Perigonimus ( r)

4. Tribe CAI.YCOPSIDI:

With more than 4 simple, unbranched radial-canals. Adradial, transversely folded gonads.

Cnhcopsis FEWKES, 1882. With 16 simple, separate, unbranched radial-canals, 4 radial, 4 interradial, 8 adradial.

Haeckel, 1879, established the family Tiandae for Anthomedusa? with 4 wide, crenulated
lips; with 4 separated or 8 cleft gonads in the stomach-wall; with 4 wide, hand-like radial-
canals; and with simple, unbranched tentacles.

Vanhoffen, 1891, showed that the gonads were interradial, not radial in position as was
supposed to be the case by Haeckel; and in 1892 Hartlaub gave important details of the
structure of the gonads, showing that Panden had simpler gonads than either Tiara or Turris.
For example, he showed that the gonads of Pandea conica consist of 4 interradial, horseshoe-
shaped, network-like, swollen regions in the ectoderm of the stomach-wall. The gonads of
T urns cceca, however, consist not only in the interradial network, hut chiefly in a double row
of fused longitudinal swellings in each interradial quadrant of the stomach. In Tiara pileata,
on the other hand, we find none of these network-like gonads, but instead a horseshoe-shaped
gonad in each interradial quadrant, the apex being upward and the sides of the horseshoe
being made up of a series of laterally-fused, horizontal swellings. Maas, 1904, loc. cit., gives
clear figures of these conditions and supports Hartlaub's observations.

In the more complex and specialized Tiarinae, represented by the tribes Amphinemidi and
Pandaeidi, the gonads are thrown into complex folds or corrugations, and the lips have become
recurved, with folded edges.

Vanhoffen concluded that Amplnnema and Codonorchls of Haeckel are equivalent to
Stomotoca L. Agassiz; and in this I concur. He also maintained that Pandea Lesson was
identical with Tiara Lesson, but in 1892 Hartlaub showed that the gonads of Pandea were
quite different in structure from those of Tiara; and Maas, 1904, supports this conclusion.

Maas, 1904, calls attention to the fact that there are no definite distinctions between
Turns and Tiara. He shows that the gonads of Turris consist of 4 interradial horseshoes, the
sides of each horseshoe being made up of partially fused, transverse, branched, ectodermal
ridges and the upper apex of the horseshoe being composed of an open network of ridges. In
Tiara the gonads are horseshoe-shaped, but commonly lack the network-like ridges. The
transverse ridges do anastomose to some degree, however, in Tiara, so that a partial network is
often found. See Maas, 1904, lor. at., plate 2, fig. II.

Maas describes the gonads of Catablema Haeckel as horseshoe-shaped and composed of
partially fused, vertical ridges. In the fully-grown medusa, however, I find that the ridges
tend to become transverse, as in Turris or Tiara.

Maas, 1904, retains the genus Catablema of Haeckel to include medusae resembling
Turns or Tiara, but with blindly-ending, lateral diverticula upon their radial-canal and ring-
canal. As a matter of fact these diverticula are highly variable in different individuals of the
same species and are seen in a more or less well-developed condition in the majority of Tiarinae.
I therefore believe that confusion will be avoided it we combine the genera Turris, Tiara, and
Catablema to form a single genus. This should be called "Turris," for Lesson used this name
on page 283 of his Hist. Zooph. Acal., and on page 17 of his "Prodrome," 1837 Tiara he
defines later on page 286 of his "Histoire," 1843, and on page 20 of his " Prodrome, "1837.

I propose, therefore, following the lead of Vanhoffen, Hartlaub, and Maas, to reduce the
13 genera of Tiarinae enumerated by Haeckel, 1879, to 8, as follows: Stomotoca, Modeeria,
Protiara, Heterotiara, Pandea, Turns, Corns, and Calycopsis.

The Tiarinae are widely distributed, but are abundant only along continental coasts, for
in so far as is known, they arise by budding from Tubularian hydroids of the genera Pcngoni-
nius, Clavula ( ?), and Campaniclava ( r). Asexual budding or direct development in the
medusa-stage is unknown.

ANTIKiMI Dl - E -I'lioTIAUA. ] Of)

The medusae <>t the various genera of Tiarinae hear a close resemhlance one to another.
Their hells are usually miter-shaped; ectodermal ocelli are often found upon their hollow ten-
tacle-bulbs, and their radial-canals arc usually hroad and Hat, and often with more or less
jagged outlines.

All of" the tentacles arise from the lower edge of" the bell-margin when young, but as
growth proceeds, the upper parts of" the basal bulbs of the older ones are crowded and forced a
short distance up the sides of the bell, while the smaller tentacles still remain upon the lower
edge of the hell-margin. 1 his gives the appearance of two rows of tentacles.

Hartlaub demonstrated that the so-called mesenteries of I lacckel. 1X70,, arc only the wide.
funnel-like origins of the radial-canals, where they communicate with the stomach-cavity.

As Maas, 1904, has shown, the Tiarinae have given rise to the more specialized Bytho-
tiandi, wherein the radial-canals have become branched, and the latter arc probably related
to the \\illiadi. I he Iiarmas are themselves derived, probably, from Codonidae in which
the originally ring-like goiuul has become radially separated, so that it lies only in intrn.uli.il
positions on the wall of the stomach. They are thus, apparently, more highly specialized
than the Codonidae. Cnl\; f>m with its 16 simple, unbranched radial-canals may be regarded
as a form intermediate between the Tiarinae and Bythotiaridi.

The Tiarinae are distinguished from the Margehnae by the tact that oral tentacles are
never found in the Tiarinae, but are present in the Margelm;e. Moreover, the tentacles of
the Tiarinae arise singly from the bell-margin, and are not grouped in clusters as is frequently
the case in the more specialized Margelinae. It seems not improbable that the Tiarinae and
Margelinae have arisen independently of each other from the Codonidae. A decided difference
between the Tiarinae and Margelmae is that in the former the entodermal cores of the tentacles
are hollow, and in the Margelinae they are nearly, it" not wholly, solid. When present the
ectodermal ocelli in the Tiarinae are on the untt'r sides of tin- tentacle-bulbs, whereas in the
Margelinae they are on the inner (velar) sides

Genus PROTIARA Haeckel, 1879.

Carminrolhe btroe, SLABBER, 177?, Physikal. BHust., p. 64.

Proliara, HAECKEL, 1879, Svst. ilrr MciluM-n, p. 46. HARC.ITT, 1902, Biological Bulletin, Boston, vol. 4, p. 17; )<)O4, Bull.

I'. S. Bureau of Fisheries, vol. 24, p. 34. LINKO, 190:, /.< -1 . Xii/rim-r, J.ilirt,-. : ^, p. i (12.
( ? ) PlolorniJe, WAI, NCR, 18X5, \VirMlosrn des WIT.M-H Mo-res, BJ. i, p. 74.
Halititirti, KFWKFS, 1882, Bull. Museum Conip. /mil. at Harvanl Collet;.-, M.|. >), p. J-M. V.\^i/ .in.) M^YJR, lS<|i), Hull.

Museum Conip. Zool. at Harvard College, vol. 32, p. 160.

GENERIC CHARACTERS.

Tiarinae with 4 well-developed, radially situated tentacles with hollow, basal bulbs. \\ ith
4 longitudinal, swollen gonads on the 4 interradial sides fit the stomach. The outer surfaces
of these gonads are smooth, not folded, nor corrugated, 1 he 4 lips are simple, not folded noi
crenulated.

Haeckel, 1^79, founded this genus for P. t<ir<i/i<-ni<i, which hail been previously described
by Slabber, 1775, under the name Carminrothe /vro, from the North Sea and Knglish Channel.
According to Vanhorfen, itfiji (X,ool. Anzeiger, Bd. 14, p. 441), this medusa is only a young
('rjr\'iiitis. However this may be, Hargitt. K;oa, discovered a medusa in Yimvaid Sound,
Massachusetts, which accords well with Haeckel's definition of Protiara. The gonads are
described by Hargitt as being found in tour separate, longitudinal, swollen regions in the
interradial (radial ?) sides of the stomach. Hargitt cut no sections of the medusa, and con-
sequently we must merely place this species provisionally in the genus Protiara, for it the
gonads be developed so as to completely surround the stomach and are not separated radially,
the medusa is one of the Codonidae. It it be one of the Tiarinae, i{ appeals to constitute
an interesting intermediate form between some Corynitis like member of the family Codonidae
and the Tiarinae. It has the simple mouth, narrow canals, and smoothly rounded external
surfaces of the manubrium characteristic of the Codonidae, but its 4 separated, interradial
gonads, and its hollow, tapering tentacles allv it to the Tiarinae.

Lmko, IQO2, sectioned a somewhat similar medusa from the Murman coast, between
Russia and Norway, and demonstrated that the 4 gonads are interradial.

The medusa described by Fewkes as f fulitnun fnrninsa is evidently a I'lotiiim.

100 MKDl'S.E OF THIO WORLD.

Protiara^beroe.

Carminrothe beroe, SLABBER, 1775, Physikalische Belustingungen, Niirnberg, p. 64, taf. 14, fig. I.

Oceania tetranema, PERON F.T LESUEUR, 1809, Ann. du Mus. Hist. Nat., tome 14, p. 347. Paris.

Protiara tetranema, HAECKEL, 1879, Syst. der Medusen, p. 47.

(?) Protiara, LINKO, 1902, Zool. Anzeiger, Bd. 25, p. 162, 2 figs.

( >)Proliara borea/is = Phtocnide borealis, Colorless variety, WAGNER, 1885, Wirbellosen des Weissen Meeres, Bd. I, p. 74, taf. 4,

fign. I, 2.
( ?) (Svndictian ?) incerturn, LINKO, 1900, Travaui Soc.Imp.Nat.de St. Petersbourg, tome 29, p. 151, fig. l (this is possibly

Sarsia flannnea ?).
( fyPlotocnidc incena, HARTLAUB, Nordisches Plankton, Nr. 12, p. 70, fig. 66.

Bell 6 to 15 mm. high, 4 to 12 mm. wide. Egg-shaped, with very thick walls, which in
the upper part of the bell are one-third to one-fourth as thick as the hell-height. There are a
few scattered nettle-cells over the exumbrella, these being more numerous in young than in
old specimens. 4 tentacles, each 4 to 5 times as long as the bell-height, with thick basal
bulbs about one-sixth as wide as the greatest width of the bell. No ocelli. The shafts of the
tentacles bear garland-like pads of nettle-cells. There are 4 straight, narrow radial-canals
with smooth edges. The manubrium is short and even in mature medusae does not project
beyond the velar opening. In young medusas it is conical, in mature individuals very wide,
but it narrows greatly at the neck immediately above the mouth. The neck is tubular, and
the mouth is encircled with nematocysts. There are 4 longitudinal, interradial, ectodermal
gonads, which project widely outward in the upper portion of the manubrium, but do not
extend to the mouth. The outer surfaces of the gonads are smooth.

Linko, 1902, sectioned the manubrium and found that the entoderm forms 4 interradial
partial septa which project inward in 4 longitudinal folds toward the axial center of the
stomach, but their inner edges do not fuse. The cells of these septa are digestive and they
serve apparently to increase the area of the stomach-wall. The bell is colorless. Manu-
brium and tentacle-bulbs orange to yellow, radial-canals white.

Common in Barents Sea, North of Russia, between 68 54' and 70 5' N. lat.; and
33 3' to 57 38' l n g- E- from Greenwich. Rare in harbors and fjords.

(Syndiction?) incerturn, Linko, 1900, may possibly be identical with P. beroe. It has
a bell 3 mm. high and somewhat more than 3 mm. wide. The walls are thick and there is a
rounded, dome-like apex sharply set off from the bell itself. The exumbrella is besprinkled
with quite regularly and widely spaced nematocysts. There are 4 thick, tapering tentacles
about one-third to one-half as long as the bell-height. These have large, spherical, non-
ocellated, basal bulbs. Velum well-developed. 4 straight, slender radial-canals. Stomach
mounted upon a short, conical peduncle. The mouth does not reach the level of the velar
opening. Mouth-opening round, without prominent lips. The gonad is figured as being much
swollen above, tapering below, and encircling the stomach. It is not wholly clear from Linko's
description whether there are 4 interradial gonads or only one encircling gonad, for he states
that the medusa resembles P. borealis = (Syndictton boreale Birula) in some of its characters.
Only about 10 very large eggs are produced by the female.

The manubrium is yellow to orange, tentacle-bulbs red, and tentacles yellow. Other
parts are colorless. Found in the White Sea, Northern Russia. It is probable that this form
is a Sarsia, and is possibly S. flammea. See Hartlaub, 1903, 1907. There is, however, no
peduncle in .V. flammea.

Protiara borealis.

Phtocnide borealis, WAGNER, 1885, Wirbellosen des Weissen Meeres, Bd. I, p. 74, taf. 4, fign. I, 2.

Synilictyon boreale, BIRULA, 1896, Annuaire Musee Zool. Acad. Imp. Soc. St. Petersbourg, tome I, p. 336 (Russian).

( ?) Protiara h<fckelii, HARGITT, 1902, Biological Bulletin, vol. 4, p. 16, fig. 4; 1904, Bull. I'. S. Bureau of Fisheries, vol. 24,

p. 34, I fig.
Phtocnide borealis, HARTLAUB, 1907, Nordisches Plankton, Nr. 12, p. 69, fig. 65.

Bell is about 3 mm. high and 1.5 mm. wide, with simple rounded apex and thick walls.
4 radially situated tentacles, each about as long as the bell-height. These tentacles are
besprinkled with nematocysts and have well-developed, conical, basal bulbs, without ocelli.
Wagner's specimens were apparently much younger than Hargitt's and each tentacle termi-
nated in a large knob-like cluster of nematocysts. In Hargitt's specimen the tentacles taper
gradually to their tips. The velum is well-developed. There are 4 straight, narrow radial-

AXTHOMEDL'S-E PROTIARA, HKTKROTIAHA.

107

canals, and a simple ring-canal. The manubrium is mounted upon a short, conical peduncle
and is quite wide; rectangular above, hut nearly circular in cross-section near the mouth.
There are 4 very small, simple lips. In Wagner's specimens the manubrium was only about
halt as long as the depth of the bell-cavity, whereas in Hargitt's the
mouth extended nearly to the level of the velar opening. This differ-
ence may be due to age(?) The gonads are 4 large, interradial,
longitudinal swellings on the sides of the stomach. The outer sur-
faces of the gonads are smooth.

Wagner found a few specimens of this medusa in April, in the
\\ hue Sea, north coast of Russia. Hargitt found an apparently iden-
tical medusa in summer near No Man's Land, an island off the
southern coast of Massachusetts. The gonads and tentacle-bulbs in
Hargitt's medusa are milky in color, other parts being transparent.
This species is distinguished from Protiara bcroe ( = P. " trtrn-
iii-iiin" Haeckel) by its absence of color and lack of ocelli.

Unfortunately no sections have been made of the manubrium.
Wagner's figure and description are unsatisfactory in that he leaves
kaiekeli," tne p os ition of the gonads in uncertainty and we can not tell whether
there are 4 (interradial ?) gonads or only I encircling gonad.

FK;. 56. Protiara

after Hargitt, in Biologi-
cal Bulletin.

Protiara formosa.

Plate 6, figs. 4, 5, and 6; plate 13, figs, l and 2.

Halitiara fortnosa, FEWKKS, 1882, Bull. Mus. Com p. Zool. at Harvard Coll., vol. 9, p. 276, platr 4, fig. 2. -MAYK.R, 1904, Memoirs
Nat. Sci. Mus. Brooklyn Inst. Arts and Sci., vol. I, No. I, p. 8, plate I, fig. 8. A(IASSIZ, A., AND MAYER, 1899, Bull. Mus.
Comp. Zool. at Harvard Coll., vol. 32, p. 160. MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 37, p. 31.

Bell about 3 mm. high and pear-shaped with solid apical projection. There are 4 long,
tapering, radially situated tentacles, about two-thirds as long as the bell-height. These ten-
tacles are hollow, with long, tapering, basal bulbs, and their ends are usually coiled in a close
helix. In addition to these 4 long tentacles there are 24 to 35 short, solid tentacles, not one-
fourth as long as the large ones. These short tentacles are tightly coiled, their axial cells are
chordate and they are more like cirri than tentacles. There are no ocelli or other marginal
bodies. The velum is narrow. There are 4 straight, narrow radial-canals and a slender
circular vessel. The manubrium is pynform to conical and about halt as long as the depth
of the bell-cavity. The mouth is a simple, cruciform opening. The gonads are developed in
the ectoderm on the interradial sides of the manubrium and the ova are large and con-
spicuous. The entoderm of the manubrium and tentacle-bulbs in the female (plate 6, fig. 5)
is green, but in the males light-brown (plate 6, fig. 4).

This medusa is very abundant at Tortugas, Florida, but is not so common in the Bahamas.
An apparently similar form was found by Agassiz and Mayer in the FIJI Islands, South
Pacific, although the Pacific form was duller in color than is usual in Atlantic specimens.

Genus HETEROTIARA Maas, 1905.
Hftnotiara, MAAS, 1905, Craspedoten Medusen der Siboga Exped., Monog. to, p. 19.

The type species is Heterotiara anonyma Maas, from the Malay Archipelago. Only two
imperfect specimens were found by the Siboga Expedition.

GENERIC CHARACTERS.

Tiarinae with 8 marginal tentacles (4 radial, 4 interradial). The ring-canal gives rise to
one (or more ?) blindly-ending, centripetal diverticula. Gonads ( ?)

Heterotiara anonyma Maas.
Hfterotiar/2 <mon\rna, MAAS, 1905, Craspedoten Medusen der Siboga Exped., Monog. 10, p. 19, taf. 3, figs. I9~2r.

Bell 12 to 1 6 mm. high and slightly more than half as wide. Oval, dome-like, with very
thick, gelatinous walls. 8 hollow marginal tentacles arise from the ring-canal and extend
through the gelatinous sides of the bell so as to reach the exterior at a slight distance above the
margin. The basal bulbs of these tentacles are small, and each one has a short, blunt, hollow.

108

MEDUS.E OF THK WORLD.

spur-like projection extending outward. There are no ocelli. All ot the tentacles were broken
off short and their normal length is thus unknown.

The ring-canal gives off a short, straight, blindly-ending diverticulum in one ot the 4
interradii above the base ot one of the interradial tentacles. 4 wide radial-canals as in other
Tiarinae. They are straight-edged and lack the "glandular-pouches" seen in many Tiarmae.

The circular muscles ot the subumbrella are very easily seen near the edges ot the radial-
canals and there are 4 interradial folds in the muscles which present the superficial appearance
of radial-canals, but they are merely radial muscle furrows, in no way to be confused with the
radial-canals.

The manubrium lacks a peduncle and is about halt as long as the depth ot the bell-cavity.
It is 4-sided at its base. The stomach part is urn-shaped, and thereare4toldedhps. Gonads( ?)
The tentacle-bulbs bear dense entodermal pigment granules. Color ( ?)

Two specimens were found by the Siboga expedition in the Malay Archipelago in vertical
nets drawn from 500 fathoms depth to the surface, in lat. o 17.6' S., long. 129 14.5' E.

It is remarkable that in each of these specimens there was but one interradial diverticulum
from the ring-canal. 4 interradial swellings are figured by Maas upon the sides of the
stomach, but he does not mention gonads. Apparently both of his specimens were immature.

FIG. 57. " Perigonimus vestutus" Hydroid and recently

liberated medusa.
FIG. 58. "Perigonimus m'mutus" Hydroid and young

medusa.
Above figures after Allman, in Ray Society, 1871-71.

Genus STOMOTOCA L. Agassiz, 1862.

( ?) Glatide bcroe, SLABBER, 1775, Physikal. Belustig., p. 46.

Saphenia, FORBES (non Eschscholtz), 1848, British Naked-eyed Medusa-, p. 25, plate I, fig. 4. McCRAnv, 1 857, Gymn. Charles-
ton Harbor, p. 27.

Stomotoca, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347. A. AGASSIZ in L. AGASSIZ'S, 1862, Cont. Nat. Hist. U. S.,
vol. 4, p. 347 (foot-note). AGASSIZ, A., 1865, North Amer. Acal., p. 168.

Dinema, VAN BENEDEN, 1867, Mem. Acad. Roy. des Sci. Belgique, torn. 36, par. i, p. 127.

Amphinema+ Stomotoca, HAECKEL, 1879, Syst. der Medusen, pp. 49, 51.

Codonorchis, HAECKEL, 1879, Ibid., p. <;i.

Dinemaiella, FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p. 151.

Stomotoca, VANHOFFEN, 1891, Zool. Anzeiger, p. 443. MAAS, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23,
No. I, p. II.

This genus was founded by L. Agassiz, 1862; the oldest species is possibly Stomotoca
slabbcri of the northern coasts of Europe. This species was first described by Slabber, i/75>
under the name of Gladde beroc, and the hydroid and young medusa were described by Van
Beneden, 1867, as Dinema slabberi. Slabber's description is, however, so unsatisfactory that
we can not be certain that this medusa is actually a Stomotoca, and can not accept it as the
type of the genus. Stomotoca Jincma (Oceania Jincma} Peron and Lesueur may be taken as
the type of the genus. The genus Saphenia Eschscholtz belongs to the Eucopidae.

PLATE 10.

Fig. i. Dissonrtna turnJa. Tortugas, Florida, May 13, 1905. An abnormal

specimen lacking ocelli and marginal cirri.
Fig. 2. Stoniotoi'a dinema, female. Agassiz Laboratory, Newport, Rhode

Island, July, 1892.
Fig. 3. Stomotoca dinema, male. Agassiz Laboratory, Newport, Rhode

Island, September 13, 1896.
Fig. 4. Stomotoca dinema, male. Agassiz Laboratory, Newport, Rhode

Island, July, 1896.
Figs. 5 and 6. Stomotoca rugosa. Agassiz Laboratory, Newport, Rhode

Island, August 2 to 16, 1892.

Drawn from life, by the author.

PLATE 10

.\\THOMKin s K MMMOTOCA.

10!)

FIG. 59. Perigonimus "serpens," after Alltnan, in Ray Socjety, 1871-72.

GENERIC CHARACTERS.

Tiarina; with 2 long, diamcrrically opposed tentacles, and with more or less numerous,
rudimentary tentacles. The basal bulbs of the long tentacles are hollow. The 4 interradial
gonads are complexly folded and are found in the ectoderm of the sides of the stomach. The 4
radial-canals are broad and flat and there is a well-developed circular canal. The hydroid
is Pengonimus.

There are a numberof species
of Pengonimus which probably
produce tree f.S'/omo/o, <i i medns.i-.

'.i/L \.1ia ( '"'/' but the medusa is as yet unknown.

Such are: Perigontmui .\,lni,-t,/iii
Motz-Kossowska, 1905, Archiv.
Zool. Exper.. ser. 4, tome 3, p. 72,
fig. vi; a red-colored Mediterra-
nean hydroid which grows upon
Membranipora and is distin-
guished by a cup-like expansion
of perisarc at the base of each
hydranth. The clavate hydranths
have about 10 tentacles. Another
Mediterranean species is P. nnpol-
itiinns of Margin, 1904, Mirth.
Zool. Sta. Neapel, Bd. 16, p. 571,
taf. 22, fig. 25; also P. steinaehi
Jickeli, 1883, Morphol. Jahrb..
Leipzig, Bd. 8, p. 617, taf. 27, figs. 1-9, from Trieste, Adriatic. We present figures of various
species of Periganimus which produce free medusae, but in which the sexually mature
medusa is undetermined. Perigonimus mitnrcticus (Hickson and Gravely, 1907, National
Antarctic Expedition of 1901 04, vol. 3, Nat. Hist., Hydroid Zoophytes, p. 4, plates I and 4)
is found attached to the stems of Halecium in depths of o to 130 fathoms in McMurdo Bay.
South Victoria Land, Antarctic. It produces fixed gonophores, and is the only form of
Perigonimus known from Polar seas.

Stomotoca dinema L. Agassiz.

Plate 9, figs. 8 to 10; plate 10, figs. I tci 4.

Oceania dinema, PF.ROX KT Ltsi'ti R, 1809, Ann. Ju Museum d'Hist. Nat., tome 14, p. 346. EsCHSCHOLTZ, iSi'j. s i. 1, t

Acalephen, p. 98.

Suf'henia dinema, FORBES, 1848, British Nakcd-eved MeJus.e, p. 25, plate 2, fig. 4 (Kiel. svn.).
tiaphenia titania, Gosst, 1853, Naturalists' Rambles, Devonshire Coast, p. 5X7, plate 26, figs. 7-9.
Stomotoca dinema, AC.ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347.
Amphintma titania, HAF.CKEL, 1879, Syst. der Medusen, p. 50. BEDOT, 1901, Revue Suisse de Zool., tome 9, p. 482; Ibid.,

1905, tome 13, p. 131 (all papers to 1850).

Amphinema dinema, BROWNE, 1896, Proc. Zool. Soc. London, p. 475.
American variety = Stomotoca apicata, L. AGASSIZ.
Male:

Saphenia apifata, McCRADV, 1857, Gymn. Charleston Harbor, p. 129, plate 8, figs. 2, 3.

Stomotoca aficata, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347. AGASSIZ, A., 1865, North Ainer. Acal., p. 168.
Amphinema apicatum, HAECKEL, 1879, Syst. der Medusen, p. 50.
Female:

Dinemalel/a cavosa, FF.WKHS, 1 88 1, Bull. Mus. Conip. Zool. at Harvard College, vnl. 8, p. i?i, plate n, figs. 2. 3; plate 4, fig.

3; 1884, Amer. Naturalist, vol. 19, p. UK, I fig.

Slomotoca apicata, MAYER, 1900, Bull. Mus. Comp. Zool. Harvard College, vol. 37, p. 3, plate . tigs. 3, 4. H
Bull. U.S. Bureau of Fisheries, vol. 24, p. 3?, 2 figs. (Figure .if male is X. apicata, but that of "female" is
NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 371, fig. 77.

r, 1904,
is S.rugoia.)

EUROPEAN MEDUSA.

Bell ^ mm. long and 2 mm. wide, with a narrow, elongate, sharp-pointed apex. 2 long

tentacles, 2 to 10 times as long as the bell-height. About 24 small tentacle-bulbs. Time aie

no ectodermal ocelli. Stomach oval, about half as long as the depth of the bell-ca\it\ and

square in cross-section. 4 well-developed, lanceolate lips. 4 simple, slender radial-canals and

9

110

MEDUS.E OF THE WOULD.

ring-canal. The gonads are 8 adradial, transversely folded, ectodermal regions on the sides of
the stomach. Stomach yellowish-brown or green, tentacle-bulbs crimson or purplish.
Found off the coasts of Great Britain. Hydroid unknown.

I have seen medusae oft" the coast of Cornwall, Eng-
land, which were identical with Stoniotoca apicata of
America.

AMERICAN MEDUSA.

In the adult American medusa the bell is about 4
mm. high and 2 mm. wide. There is a large apical
projection which is hollow in the female, but usually solid
in the male medusae. The bell-walls are thin and the
sides vertical below, but slopinginward above to a pointed
apex. There are 2 long, diametrically opposed ten-
tacles, with large, tapering, hollow basal bulbs. There
are no ocelli. These long tentacles are highly contractile,
but are usually about as long as the bell-height. Their
shafts are covered with small wart-like clusters of nema-
tocysts. In addition to the 2 long tentacles, there are 6
or more small rudimentary tentacle-bulbs upon the bell-

FIG. 60. Stoniotoca dinema, male. From life, by
the author. Mousehole, Cornwall,
England, October 28, 1907.

Synopsis of the Species of Stoniotoca.

S. dinema
Haeckel.

S. "apicata"
L. Agassiz.

S. atra
L. Agassiz.

S. octaedra.

S. rugosa

Mayer.

S.pterophylla
Haeckel.

S. divisa
Maas.

Size of bell
in mm.

3 long, 2 wide.
Apex solid.

4 high, 2 wide
with solid
apex in male
and hollow
apex in
female.

25 high, 22

wide. Apex
solid.

2.5 high, 2
wide. Apex
solid.

5 high, 3 wide.
Apex large,
solid. Sides
thin.

30 wide, 12 high.
Small apical
projection.
Flaring sides.
Thick gelati-
nous substance

30 wide, 20
high. As in
S. ptero-
phylla.

Length of
tentacles

6 to 30.

4

io

5

30

20

AsinS.

pterophylla.

in terms

of bell-ra-

dius (r).

Number of
rudimen-

24

6

80

6

'4

60 to 80.

As in S.
pterophylla.

tary ten-
tacle-

bulbs.

Ectodermal,
abaxial,

None.

None.

None.

8, orange-
colored.

None.

None.

None.

tentacular

ocelli.

Shape of
manu-
brium.

Oval, half as
long as
depth of
bell-cavity.

Flask-shaped.
4 folded,
cruciform
lips.

Half as long as
depth of
bell-cavity.
Gonads
folded as in

As in S. api-
cata.

As in S. atra.

As in S.atra.

As in S.
pterophylla.

Turris.

Color.

Stomach

In male, stom-

Mouth dull-

Stomach and

Stomach, gon-

Stomach and

Gonads

yellowish-
brown to
green. Ten-
tacle-bulbs
purple or
crimson.

ach green
and tenta-
cle-bulbs
purple. In
female, dull
milky-yellow.

yellow,
gonads very
dark-brown,
nearly black,
tentacles
light-brown.

tentacles
green, with
orange cen-
ter in stom-
ach.

ads, and ten-
tacle-bulbs
brick-red
to nearly
black.

tentacle-bulbs
brown. Radial-
canals and
gonads milky-
white.

orange to
cinnabar-
red. Stom-
ach and ten-
tacle-bulbs
yellow.

Where

found.

Coast of Great
Britain.

Atlantic coast
of United
States.
Probably

Port Town-
send, Wash-
ington.
Pacific coast

Tortugas and
the Baha-
hamas, in
summer.

Woods Hole to
Tortugas,
Atlantic
coast of

Bahamas, Tortu-
gas, West
Indies.

Pacific coast
of Central
America.

identical

of America.

United

with S. di-

States.

nema.

PLATE 1 1 .

Figs, i and 2. Stotnotoca rugosa, young and mature. Newport, Rhode
Island, August 4, 1892.

Fig. 3. Young medusa of Perigonimus jonesii. Agassiz Laboratory, New-
port, Rhode Island, August 22, 1896.

Fig. 4. Young medusa of Perigonimus jonesii. Agassiz Laboratory, New-
port. Rhode Island, July, 1896.

Fig. 5. Stotnotoca octaedra, young medusa. Tortugas, Florida, July 17,
1898.

Fig. 6. Stomotoca octaedra, Tortugas, Florida, June, 1897.

Fig. 7. Pandca violacca. Aboral view of gonads of the medusa shown in
figure i, plate 12.

Drawn from life, by the author.

r

\ vrin IM Knrs.K- -ST< >M< >rt ><-.\. Ill

margin. These lack ocelli. The 4 radial-canals are wide and flat. The boundaries of the
radial-canals and of the ling-canal are often bluntly serrated. The manuhrium is flask-
shaped; there is no peduncle and the 4 lips are cruciform, crenulated, and curve slightly
upward. The ectoderm of the upper part of the manuhrium on both sides of the 4 ladial-
canals is thrown into interradial folds or convolutions and the gonads are developed in
this region.

In the male the color of the manuhrium varies from intense green to translucent ocher-
yellow or cream-color. The basal bulbs of the tentacles in the male vary from faint to deep
purple. In the female the manubrium is usually translucent ocher-\ t -\\n\\ or cream-colored,
but occasionally straw-colored or faintly green. The tentacle-bulbs of the female are usualU
translucent milky or ocher, but occasionally one is found having faintly purple tentacle-bulbs.
These sexually dimorphic color peculiarities are seen in the smallest and youngest medusae
as well as in adults. In very young medusae the apical projection is small or absent. There are
2 tentacles and only 2 rudimentary tentacle-bulbs.

This medusa is common at Newport, Rhode Island, from |ulv 15 to September. It is
not common at Tortugas, Florida, and is not seen north of Cape Cod, Massachusetts. On
October 28, 1907, I found two male medusae of this species oft" Mousehole, Mounts Bay,
Cornwall, England.

This species furnishes that which is probably the most remarkable instance of sexual
dichromatism to be met with among the hydromedusae, the color differences affecting parts
other than the gonads.

The two males found off the coast of Cornwall, Kngland, were each 2.2 mm. high and
similar in all respects to the typical American form; with intense green manuhrium and
purple tentacle-bulbs, without ocelli. The bell-apex was solid and gelatinous. 1 believe that
S. apicata of America. IS identical with S. Jlni'inn of Europe. The European medusa may
have more rudimentary tentacle-bulbs, hut these are very variable in number in the Ameri-
can form.

Stomotoca atra Agassiz.

Stottiotora atrn, AGASSI/, I... i862 f Cont. Nat. Hist. U. S. f vol. 4, p. 347.- A<.-\SM/, A., iM>^. N^rth AIIMT. \<,il., p. it>^, figs.
271-27;. H^H-KH., 1X71), Sysl. ilcr Meilusen, p. 53.

Bell 20 to 25 mm. high, 20 to 22 mm. wide. Bell-shaped, higher than a hemisphere.
2 long, highly contractile tentacles, and about 80 small rudimentary tentacle-bulbs. Stomach
about half as long as the depth of the bell-cavity. Swollen, 4-sided, and mounted upon a
peduncle. 4 lips. The gonads a re 8 ad radial, linear cross-foldings upon the sides of the Stomach,
each gonad consisting of I 2 to 15 folds as in Turns. The mouth is dull yellow. Gonads dark-
brown to almost black. Tentacles light-brown.

Found by A. Agassiz in great numbers in Port Townsend, Washington, Pacific coast of
the United States; from |une until September.

Stomotoca octaedra.
Plate 1 1, ti);s. <; ami f>.

Cotlunorchis octnetlrus, HAFCKK.L, 1879, Syst. Jrr MrtliiM-n. p. 51.

Sloniolocn austrtilis, MAYKR, 1900, Bull. Mus. (.'nrn|'. /.mil. .it Harvard College, vol. 37, p. 32, plate I, lii;. i; lintl., 1904,
Mrin. Nat. Sci. Brooklvn Inst. Museum, vol. i, p. 9, platr i, fig. 9.

The following description is derived from studies of specimens found in the Baham.is.
and at Tortugas, Florida.

Immature nirJ/isn (plate 11, fig. 6). Bell about 2.5 mm. high and with well-developed,
sharp-pointed, apical projection upon aboral side. Bell-walls thin.' There are 2 radially
situated, diametrically opposed tentacles, each about as long as the bell-height and with
long hollow, tapering basal bulbs. There is a single, ectodermal, orange-colored ocellus upon
the outer side of each tentacle-bulb near its point of origin from the umbrella. In addition to
the 2 long tentacles there are 2 short, rudimentary tentacle-bulbs go from the long tentacles.
Each of these rudimentary tentacle-bulbs has an ectodermal, orange-colored ocellus upon its
outer side, and there are 4 other ocelli upon the bell-margin in interradial positions. There
are thus 8 ocelli in all, 4 on the tentacle-bulbs and 4 interradial ones on the bell-margin. 'I he

112 MKDUS.E OF THK WOULD.

velum is well developed. There are 4 straight-edged, wide radial-canals and a broad circular
vessel. The manubrium is urn-shaped, quite wide, and with 4 cruciform, recurved lips. The
mouth is at a point about half-way between the inner apex of the bell-cavity and the velar
opening. The gonads are found in complexly folded regions upon the 4 interradial sides of
the stomach. The manubrium and tentacle-bulbs are yellow or greenish-yellow. The ento-
dermal core of the stomach is often orange.

When about I mm. high (plate 1 1, fig. 5) the bell has a small apical projection. There
are 2 large tentacles and 2 rudimentary tentacle-bulbs, each with an orange ocellus. There
are no other ocelli upon the bell-margin. The manubrium is slender and urn-shaped, without
gonads and with 4 simple, cruciform lips.

The young of this medusa are common at Tortugas, Florida, and in the Bahamas through-
out the summer. Although the medusa has usually but 2 long tentacles, occasionally one is
seen with 4 long, equally developed, radially situated tentacles. The mature medusa has not
been seen and we must remain in doubt concerning its generic position, for it may be a young
Turn's. The presence of ocelli upon the tentacle-bulbs distinguishes it from all other Ameri-
can species of Stornotoca and is a character commonly seen in Turns.

I believe this medusa to be identical with Haeckel's " Codonorchis octaedrus," which he
obtained offthe Atlantic coast of France. Haeckel states that this medusa had 2 well-developed
tentacles and 10 tentacle-bulbs (2 perradial and 8 adradial). He described the ocelli as
brownish-red. Apparently he found but a single specimen and the interradial tentacle-bulbs
may have failed to develop. His medusa is described as having folded gonads, and is said to
be 4 mm. high and 2.5 wide.

Stomotoca rugosa Mayer.
Plate 10, figs. 5 and 6; plate 11, figs, i and 2.

Slomoloca afirala, FEWKF.S, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p. 152, plate 2, figs, i, 4, 9. HARC.ITT,
1901, American Naturalist, vol. 35, p. 581, fig. 40; 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 35, i fig. (errone-
ously labeled Stomotoca apicata, female). RITTENHOUSE, 1907, Proc. Boston Soc. Nat. Hist., vol. 33, pp. 440, 445,
452, 456 (embryology).

Stomotoca rugosa, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 4, plate 2, fig. 5; Ihitl., p. 32.

Amphinema u[titatuin, BROOKS, 1883, Studies Johns Hopkins Biol. Lab., vol. 2, p. 473.

Bell 5 mm. high and 3 mm. broad. Solid, apical projection, in some individuals short and
blunt, in others long and slender. Bell-walls of moderate thickness, and bell itself, exclusive
of the apical projection, somewhat higher than a hemisphere, with relatively vertical, straight
sides. There are 2 long, diametrically opposed tentacles, which are of equal length, and when
stretched are fully ten times as long as the bell-height. They are, however, highly contractile
and may become not more than one-tenth as long as when fully expanded. The basal bulbs
of these long tentacles are large, hollow, and tapering and there are no ocelli.

In addition to the 2 long tentacles there are 14 small, permanently rudimentary tentacles,
2 at the bases of 2 of the radial-canals and 3 in each interradial quadrant. These lack ocelli.
The velum is well developed. There are 4 wide radial-canals and a ring-canal, all with
jagged edges. The manubrium is quadratic and flask-shaped and the mouth in old medusae
is about at the level of the velar opening. The mouth is cruciform and there are 4 prominent,
recurved, crenulated lips. The mature sexual products are found in the ectoderm of the
adradial walls of the stomach on both sides of the places of entrance of each of the 4 radial-
canals where the surface is thrown into 8 series of complex folds and ridges, a double ridge of
folds in each interradius.

The gelatinous substance of the bell is transparent, but the entoderm of the tentacle-bulbs
and stomach is brick-red, often streaked with sooty brown. The radial and circular canals are
faint red in color. Specimens from Tortugas, Florida, often show black streaks through the
brick-red color of the stomach and tentacle-bulbs, and medusae which have been confined in
aquaria for some days often become wholly black in these parts.

Hydrotd and \onng medusa. Professor W. K. Brooks, 1885, describes the hydroid.
It is a Pi-rigfjniiniis, very much like P. niiniitiis Allman, 1871 (Monog. Tubularian Hydroids,
p. 324, plate xi, figs. 4 -6). It was found at Beaufort, North Carolina, growing upon the lower

ANTHOMKIH'S.K STOMOTOfA. 113

surface of the shells of Limulus, fastened to the sand-tubes of Sabfllarin. The stems are simple
and unbranched and are only about 0.2 mm. in height. They are covered for about two-
thirds of their length by a delicate, closely-adherent film of perisarc to which foreign particles
become attached. The stomach occupies about one-fourth or one-fifth of the length of the
stem from which it is separated by a slight constriction. Each polypite has 10 tentacles which
point alternately backwards and forwards, those pointing forwards being a little longer than
the others. The medusae are attached by very short peduncles to the sides of the stems. When
the medusa is set free it is about 0.5 mm. in height and there is no trace of the apical projection,
which develops in about eight days.

In an abnormal individual medusa of this species found at Newport, Rhode Island, in
July, 1892, there were 4 long tentacles, i at the base of each of the 4 radial-canals. This
medusa was maintained alive in an aquarium for more than a month. When first found it had
only 2 long tentacles which were diametrically opposite one another at the bases of i of the
radial-canals. The other 2 tentacles developed later, after the first pair had attained their full
length. The medusa had then 4 radially placed tentacles and 12 rudimentary tentacle-buds.
This variation is interesting, as it illustrates the close relationship between Stiiniutrnu and
Turns.

Stomotoca ruorjsa is common on the southern coast of New England in summer. It is
found all along the coast to southern Florida, but is not very common at Tortugas or among tin-
Bahama Islands. It has never been seen north of Cape Cod, Massachusetts.

Rittenhouse, 1907, finds that the eggs of this medusa are laid between 5'' to 5'' 30"' in the
morning. The egg is chalky-white. The entoderm is formed by cellular ingression. The
planula settles down upon its side and becomes a branched hydrorhi/a from which the p<>!\ -
piles bud out.

Stomotoca pterophylla Haeckel.

Plate 2<), figs. 5 to ^; pl.Ue 30, fig. 7.

Xtomotoca ptfroplnlla, HAKCKF.I., 1879, Syst. der Medusen, p. 52, taf. 4, fig. 10.
Xiumolorti persphyHa, F I:\VKKS, |SX<), Report Comrmss. Fish and Fisheries for 1X86, p. 524.

Adult medusa. Bell conical, with widely flaring sides and small, sharply-pointed apical
projection. It is about 20 to jo mm. in diameter and 10 to 12 mm. in height. The gelatinous
substance is very thick at the aboral pole, but becomes thin at the bell-margin. There are 2
long, tapering, marginal tentacles which are situated at the bases of 2 of the radial-canals, 1 80
apart. When expanded these tentacles are full}- 10 times longer than the bell-diameter. In
addition to the long tentacles, there are about 60 to 80 small rudimentary tentacle-bulbs, 15 to
20 in each quadrant. The 4 radial-canals are wide, ribbon-like, and flat, with smooth, simple,
outer edges. The ring-canal is narrow and its upper edge is smooth. There is a very wide,
conical peduncle which extends about to the level ot the velar opening. The gastric part of the
manubi mm is large and swollen and lies mainly outside ot the bell-cavity. There are 4 promi-
nent, complexly crenulated, lanceolate lips. The gonads occupy 8 adradially situated, longitu-
dinal swellings upon the sides ot the stomach (plate 29, fig. 4). Each gonad consists ot a io\\
of swollen, leaf-like ridges which trend in a transverse direction. The stomach and tentacle-
bulbs are brown. Radial-canals milky. There are no ectodermal ocelli.

This species is found in the West Indies and the warmer parts of the Gulf Stream. I
found many specimens among the Bahamas and at the Tortugas during the spring and early
summer ot 1907.

It is closely related to Stomotoca Jivisa, described by Maas, from the west coast of Mexico
(Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. n, tat. i. tigs, i i<j, 180,7). ' Hlt
its rich-brown entoderm distinguishes it from the Pacific species.

Haeckel describes this medusa from preserved specimens, and this may account for
certain errors in his description and figure. A small apical projection appears to be con-
stantly present and there are no "ocelli." The radial and circular muscles of the velum are
very powerfully developed. In extreme states ot contraction of the bell the peduncle mav
extend beyond the velar opening or tall short of reaching it. The long tentacles are higlih
contractile and when expanded are reduced to mere thread-like filaments.

Ill MKDrS.E OF THE WORLD.

Stomotoca divisa Maas.

Siomoioca Jk'isa, MAAS, 1897, Mem. Museum Comp. Zool. at Harvard College, vol. 23, p. u, taf. i, fign. 1-9.

Bell 20 to 30 mm. wide, 15 to 20 mm. high. Upper part of bell thick, solid, and dome-
like, and separated by an annular furrow from the thin-walled, marginal part of bell. This

furrow may be due to contraction ( ?) A
very small, pointed apical projection arises
sharply from the aboral surface of the
evenly rounded dome of the bell. There
are 2 tapering tentacles about as long as the
bell-height. These appear to have some-
what more swollen basal bulbs than are seen
in S. pterophylla. There are no ocelli. There
are a large number of rudimentary tentacle-
bulbs as in S. pterophylla. The bell is trans-
parent with yellow entoderm, and with
orange to cinnabar-red gonads.

IMI.. 61. Stomotoca Jivisa, after Maas, in Mem. Museum Comp. It IS found in the Bay oi Panama,

Zool. at Harvard College, showing different statesof P ac jfi c coast o f Central America, ill March,
contraction of the bell. -.......,- ... .

It is distinguished from the closely allied

.S\ pterophylla of the West Indies by its brilliant coloration, S. pterophylla being constantly
dul I -brown.

Perigonimus jonesii Osborn and Hargitt.
Plate n, figs. 3 and 4.

Perigonimus jonesii, OSBORN and HARGITT, 1894, Amer. Naturalist, vol. 28, p. 27, figs. 1-12. HARGITT, 1895, Mittheil. Zool.
Station Neapel, Bd. u, p. 479; 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 33, I fig.; 1901, American Naturalist,
vol. 35, p. 308, fig. 4; p. 579, fig. 37. NUTTING, 1901, Bull. U. S. Fish Commission, vol. 19, pp. 331, 372, fig. 80.

} oung medusa. None of the specimens yet seen were mature. Bell of largest 2 mm.
in height and side walls extremely thin and flexible. There is a very small, dome-shaped
apical projection. There are 2 well-developed, diametrically opposed tentacles, which are
situated at the bases of 2 of the radial-canals. These tentacles are of unequal length and
are at times carried curled in a close helix and at other times are extended to their full length,
in which case one of them becomes about as long as the bell-height, while the other attains to
about twice this length. The basal bulbs of these tentacles are long, conical, and hollow
and have no ocelli, and the shafts of the tentacles are thickly covered with nematocyst-cells.
In addition to these well-developed tentacles, there are 2 small tentacle-bulbs, situated at
the bases of the 2 radial-canals 90 away from the long tentacles. There are no ocelli. The
velum is wide and flexible. There are 4 straight, slender radial-canals and a narrow, circular

' O

tube. The manubnum is short and simple, with a wide base and 4 simple, cruciform lips.

The ectoderm of the manubrium and tentacle-bulbs is of a dull ocher-yellow.

This medusa is very rare; only 3 specimens have been seen by me during 3 summers' study.
All of my specimens were found in Newport Harbor, Rhode Island, during July and August.

Hydroid. The hydroid has been described by Osborn and Hargitt, 1894, from Cold
Spring Harbor, Long Island, New York, where it is found very commonly upon the abdomen
and upon the walking legs of the spider-crab (Libinia emarginata). It is a Pengoniiniis.
The stems arise from a creeping hydrorhiza and branch luxuriantly. The oldest polypite is
found at the distal end of the stem. The stems are covered with a thick gelatinous pensarc
which extends up the stems to the level of the tentacles, to which it is fastened. There are
normally 16 tentacles in a single circlet around the base of the hypostome. These assume an
alternately reflected position. The medusa-buds arise in clusters from near the center of the
stems. They are covered with the thick perisarc and each one is attached to the stem by
means of a well-developed peduncle. When set free the medusa has 2 diametrically opposed
tentacles which are usually carried turned inward into the bell-cavity. The hydroid is flesh-
colored.

The medusa can at once be distinguished from S. apictita by the extreme tenuity and
flexibility of the bell-walls. The bell is also higher and narrower than in S. apicata The

ANTIKiMKDIS.K ST< iM( iT( i( 'A, DISSi i\KM \ . II.")

medusa swims by the aid ot a rhythmical series ot wave-like contractions which travel one
after another up the very flexible sides ot the hell, from margin to apex. The efficiency ot
these movements is greatly enhanced In the cooperation of the powerful velum.

The 4ully-developed medusa is unknown and it-is possible that this medusa may not be
Stomotoca. It can not be referred to any Turns known upon our coast, and the tact that the
2 long tentacles are of unequal length, one being considerably longer than the other, separates
it at once from any other known species of Sttjtnotrjici. At present we place it provisionally
in the genus Strjinotoca to which its hydroid appears to refer it. Were it not for the absence
ot ocelli upon the tentacle-bulbs I would be inclined to suspect that this medusa might prme
to he the young ot Turns pileato.

(Stomotoca?) Perigonimus cidaritis Weismann.

Pfrigottiniiis cidaritisi WUSMANN, 1885, Entstehung Sexualzellen bei Hyiimmciluscn, pp. 117, 218, taf. 12. figs. 10, 11.

The stems of the hydroid are about 90 mm. high and arise thickly one by the side of the
other from a root-like hydrorhiza. The stems are often simple and unbranched, but tlu\
commonly branch dichotombusly one or two times, the branches being set off at acute angles
one with another. The hydranths have a circlet ot about 20 tentacles. Perisarc thin and
yellowish. The medusa-buds are borne upon pedicels in a zone at the bases ot the hydranths.
When set free the medusa is spherical, with 4 marginal tentacles and 4 radial-canals. The
manubrium has 4 short, knobbed oral tentacles. The whole hydroid stock is invested with
a thin layer of slimy mud, leaving only the mouth-ends of the hydroids projecting, and some-
times even those are covered, leaving only the tentacle tips tree.

Found at Naples, Italy. Medusa set tree in March.

The sex-cells originate in the ectoderm ot the proximal part of the manubrium of the
medusa-bud and develop in their place of origin, becoming mature in the tree medusa. The
mature medusa has not been determined.

Genus DISSONEMA Haeckel, 1870.

l)ionrma, HAF.I KM, 1X7.), Svst. dcr Mrdusen, p. 126. MAYF.R, 1900, Bull. Mus. C'oinp. /.mil. .11 H.ir\.ir,l (.'iill'-i'r , \nl. ;-,
p. 44. BK;F.UHV, H. B., 1909, Bull. MiiM'um of Comp. Znol. ;it Harvanl Colli-iv.

This genus was founded by Haeckel, \$~(), for Dissotit-itiii .\ii[>lii'n,-llii, of the- coast <>l
Australia.

GENERIC CHARACTERS.

Tiarinas with two hollow, diametrically opposite tentacles. The gonads extend from
the manubrium outward along the canals. Without marginal sense-clubs. Cirri aie some-
times present. With abaxial, ectodennal ocelli upon the bases ot the tentacles.

Rigelow, 1408, finds that the adradial gonads begin to develop upon the sides of the
manubrium and that afterwards they extend outward over the radial-canals. The hollow
tentacles, abaxial ectodermal ocelli, rudimentary tentacle-bulbs, and hollow bell-apeK are
all characteristic of the Tiarinae., and it appears that this genus must be removed trom the
Thaumantiad;e, in which Haeckel placed it.

J)is.friiii-niii bears the same relation to Stin<tt't,<i that .\<//;<//MM does to Bougainvillta.
The hydroid is unknown.

Dissonema saphenella Haeckel.

/>;\w,'M w<; niplifntlla, HAFI KKI, 1879, Svst. tier Mcduscn, p. 126, t:tf. 8, rl^. }.

Bell pyriform, with thick, solid apex and thin, bulging sides. 6mm. high, 4 mm. uulc.
2 well-developed tentacles, several times as long as the bell-height. These tentacles have
large, swollen, conical basal bulbs. There are 2 rudimentary tenta'cle-bulbs yo apart trom
the long tentacles. There is an abaxial (ectodermal ?) "ocellus" ot large si/e upon each ot
the 4 tentacle-bulbs. There are no marginal clubs, cirri, or other appendages.

The velum is well-developed and there are 4 straight, narrow radial-canals and a ring-
canal. The manubrium is cylindrical, half as long as the depth of the bell-cavity and pro-
vided with 4 short, crinkled lips. The 4 folded, spindle-shaped onads are developed upon
the middle three-fourths of the radial-canals. Color ( ?) Coast ot Australia. (See tig. '

Ill)

MEDUS.E OF THE WORLD.

Dissonema turrida Mayer.

Plate 10, fig. l; plate 22, fig. I.

Dissonema turriJa, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 44, plate 2, figs. 3, 4; 1904, Mem.
Nat. Sci. Brooklyn Institute Museum, vol. i, p. 8, plate 2, fig. 10.

Adult medusa. Bell bluntly cone-shaped with a hollow apical projection. It is 4 to 7
mm. high and the side walls are thin and flexible. There are 2 long, hollow, gradually tapering

tentacles which are quite contractile, but
when expanded are 3 to 4 times as long as
the bell-height. Each of these tentacles has
a minute, red ocellus in the ectoderm of its
outer side near the bell-margin. In addition
to the two large tentacles there are 12 to
14 small, solid cirri which arise separately,
at equal intervals, from the bell-margin.
Each cirrus has a red ocellus in the ecto-
derm of its outer (abaxial) side at the bell-
margin. The velum is well developed.
The circular vessel is narrow, but the 4
radial-canals are broad, and the 4 convo-
luted, bag-like gonads occupy three-quar-
ters of their length and also lie upon the
adradial sides of the manubrium. In the
female each gonad contains 4 to 7 large eggs
which project prominently over the surface
of the organ. The manubrium is pyritorm
and the mouth projects beyond the velar
opening. The stomach-walls are thin and
flexible, and the mouth is surrounded by
large recurved lips with crinkled edges. 1 he
entoderm of the manubrium and tentacles
is a delicate green and the genital organs
and circular canal are tinged with green or
pink. The ocelli are red.

In young medusae there are but 2 large
tentacles and 2 cirri. There are 8 ocelli.
The 2 long tentacles have tapering, hollow
basal bulbs, but their main shafts are at
first solid, though later they become hollow.
This medusa is common on the surface
in the Bahamas and at Tortugas, Florida,
in summer.

Dr. H. B. Bigelow shows as a result of
sectioning the medusa of Dissonema that the
gonads begin to develop upon the adradial
sides of the manubrium and later extend
outward down the radial-canals. This fact,
taken in connection with the abaxial ectodermal ocelli, the hollow tentacles, bell-apex, rudi-
mentary tentacles and general form of the medusa, make it evident that Dissonema is one
of the Tiarinne.

Genus PANDEA Lesson, 1843.

Pan/tea, LESSON, 1837, Prodrom. Monog. Meduses, No. 22 (not published); 1843, Hist. Zooph. Acal., p. 288. ACASSIZ, L.,

1862, Cont. Nat. Hist. U. S., vol. 4, p. 347.
Pantlepa, HAECKEL, 1879, Syst. der Medusen, p. 53.
Pandra, AGASSIZ and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 160. HARTI.AI-B, 1892, Nachricht.

kgl. Gesell. Wissenschaft. Univ. Gottingen, p. 21. MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, p. 15.
Diantfa, Quov ET GAIMARD, 1827, Annal. des Sci. Naturelles, tome 10, plate 6.
.\'on Dianefa, LAMARCK, 1816, Hist. Anim. sans Vert., tome 2, p. 505.
Pandja, ZOJA, 1892, Boll. Sc. Paira, Anno 1891, Nos. 3, 4, 3 pp.; 1892, Annals and Mag. Nat. Hist., ser. 6, vol. 9, p. 409.

(Deniiroclam dohrnii gives rise to a medusa resembling Pandea ?)

Fin. 62. Dissonema saphenella, after Haeckel, 1879.

.\\THilMKDCS.E r.\\T)K\.

11;

The type species of PanJea is P. cnnlca Lesson, 1843. This medusa was previously
described by Ouoy and Gaimard (1827, Annal. Sci. Nat., torn. 10, p. 182, plate 6) under the
name Diantea conica. It is found in the Mediterranean. Haeckel introduced the spelling
" PanJ(ta," although Lesson's original spelling is PanJea. The genus "Diantea" was founded
by Lamarck (1816, Hist. Anim. sans Vert., torn. 2, p. 505), the type species being Diancea
tneJra, a medusa which had been previously described by Pt-ron under the generic name of
Lymnorea. Lamarck's genus Duimta also included various species of Geryonta, Oceania,
Pelagia, and Mfdusa of previous authors, and is so hopelessly involved that in my opinion
it must be dropped.

GENERIC CHARACTERS.

Tiarinae with 4 or more well-developed, marginal tentacles. There are 4 interradial,
genital ridges in the ectodermal wall of the stomach, but these gonads are only imperfectly
separated in the 4 principal radii; so that the stomach is completely encircled by the genital
organs, which are, however, better developed in the interradii than in the principal radii.

DenJroclava is possibly the hydroid of some species of PnnJea, Coins, or Turn's-, but the
mature medusa is unknown.

When set free the medusa of DenJroclava dohrnii (Weismann, 1883, pp. 26, 216, taf. 12,
fign. 6-9) has 8 marginal tentacles, 4 radial-canals, and a manubrium with 4 band-like gonads

i<;. 6j. Pandeei conica.

A. Side view of half-grown medusa. Bell 13 mm. high. Gonads interradial.

H. Manubrium of mature medusa, with bell 21 mm. high. Gonads continent, surrounding stomach.
C. Manubrium of young medusa, showing interradial gonads. Bell (> nun. lir\;h. l-'rom lit- . I'.
the author. Zoological Station, Naples. Winter of 11)07-08.

on its interradial sides. The sexual cells originate in the ectoderm of the manubrium of the
medusa-bud and develop in the ectoderm of the free-swimming medusa, without wandering
from their place of origin. The hydroid of DenJroclava dohrnii was found by Weismann at
Naples in the summer of 1881, attached to a stock of Isis which was dredged from a depth ot
35 to 40 fathoms. The stems are about 35 mm. high and arise from a short, creeping hydro-
rhiza, which together with the stems is covered by a smooth, horny perisarc. The main stems

118

MKDUS.E OF THE WOULD.

give rise to alternate side branches. The hydranths are cluh-shaped and bear 12 to 20 short,
filiform tentacles, scattered over their whole sides. The medusa-buds arise singly from the
side branches close under the neck of the lateral hydranths. Pictet, 1893, found a very similar
or identical hydroid at Amboina, Malay Archipelago. The principal references to this clava-
like hydroid are as follows:

DenJroflava Johrnii, WEISMANN, 1893, Entstehung Sexualzellen bei Hydromedusen, Jena, pp.

26, 215, taf. 12, fign. 6-9. DU PLESSIS, 1888, Reeueil Zool. Suisse, tome 4, p. 531 (from

Villefranclie).
DenJroclava dohrni, PICTET, 1893, Revue Suisse Zool., tome I, p. 6, plate I, figs. I, 2 (from

Amboina).

It does not seem probable that Dcndroclava dohrni produces a Pandea-like medusa, for it
is well established by Brooks that Tumtopsis nutricula comes from a DenJroclava hydroid.

Pandea conica Lesson.

Dian<ta conica, p_i OY IT GAIMARD, 1827, Annal. des Sci. Nat., tome 10, p. 182, plate 6 A, figs. 3, 4.

Pandea conica, LESSON, 1843, Hist. Zooph. Acal., p. 288. HAECKEL, 1879, Syst. der Medusen, p. 54. MAAS, 1904, Result.
Camp. Sci. Prince de Monaco, fasc. 28, p. 16, plate i, figs. 6, 7 (figures of gonads). BEDOT, 1905, Revue Suisse de
Zool., tome 13, p. 141 (literature, 1827-50). LOBIANCO, 1903, Mitth. Zool. Sta. Neapel, Bd. 16, p. 217.

Oceania setiecimcostata, KOLLIKER, 1853, Zeit. fiir wissen. Zool., Bd. 4, p. 324.

Oceania conica, GEGENBATR, 1856, Zeit. fiir wissen. Zool., Bd. 8, p. 221, taf. 7, fign. 1-3.

Bell with barrel-shaped sides, and bluntly pointed or concave apex; 21 mm. high, 10 mm.
wide. There are 8 to 24 well-developed, longitudinal rib-like ridges in the tentacular radii on
the exumbrella, and an equal number of marginal tentacles with abaxial ectodermal ocelli.
Stomach wide and short with 4 folded lips having sinuous margins. The gonads consist ot 4
crescentic, reticulated, swollen regions in the ectoderm of the 4 interradial sides of stomach,
the convexity ot the horseshoe pointing aborally. These gonads are not completely separated
in the adult, in the 4 principal radii, so that they surround the stomach on all sides.
The best figures illustrating their structure are given by Maas, 1904.

Stomach brownish. Gonads and
tentacle-bulbs brownish-red. Tentacles
yellowish-milky. Ocelli dark-purple.
Common in the Mediterranean.
This medusa was quite common
during the winter of 1907-1908 at
Naples, Italy. When young the 4 inter-
radial gonads are completely separated,
but later they fuse more or less over the
perradii, forming a complete, swollen
network which girdles the stomach.
The prominent longitudinal ridges over the exumbrella arise after the tentacles begin to
develop and extend upward from the bases of the tentacles toward the bell-apex. The table
gives a growth-record obtained by the author from specimens of this medusa observed at
Naples. (See fig. 63, page 117.)

Pandea saltatoria Lesson.

Oceania saltatoria, SARS, 1835, Beskriv og Jagtt., p. 25, plate 4, fig. loa-r.

Pandea sallatoria, LESSON, 1843, Hist. Zooph. Acal., p. 290. HAECKEL, 1879, Syst. der Medusen, p. 54. BEDOT, 1901, Revue
Suisse de Zool., tome 9, p. 485; Ibid., 1905, tome 13, p. 141 (all literature to 1850).

Bell 8 mm. high, 6 mm. wide, with a pointed apex and rounded sides. Exumbrella with
only 12 to 16 longitudinal lines of nettling-cells instead of about twice as many as in P. conica.
24 to 32 tentacles; twice as many as the lines of nematocysts, instead of being equal to these in
number as in P. conica. Ocelli on outer sides of tentacle-bulbs. Stomach slender, lips small
and simple. Immature (?) Color (?) Coast of Norway, Bergen.

Pandea minima von Lendenfeld.

Pand<ra minima, vox LEXDFNKELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 916, plate 42, figs. 10-12.

Bell semi-ovate, 3 mm. high, 2 mm. wide. 8 tentacles about as long as bell is high. 8
adradial lines of nettle-cells upon the exumbrella. A bundle of nerve fibers is said to extend
under each linear series of nettle-cells connecting the clusters one with another! The man-

Height of
bell in mm.

No. of

tentacles.

Condition of the gonads.

6

16 4 small interradial.

X

16 4 larger interradial.

'3

21 4 interradial nearly touching in the

perradii.

21

24

A unitary network of ridges sur-

rounding the stomach.

PLATE 12.

Fig. i. Pandea violacca, mature female. Tortugas, Florida, June 5, 1906.

Fig. 2. Turris vesicaria, young medusa. Eastport, Maine, September 19,
1898.

Fig. 3. Turris vesicaria, half-grown medusa. Agassiz Laboratory, New-
port, Rhode Island, June 24, 1893.

Fig. 4. Turris pileata, half-grown medusa. Agassiz Laboratory, Newport,
Rhode Island, July 13, 1895.

Fig. 5. Podocoryne fulgurans. Basal part of one of the marginal tentacles.

Figs. 6 to 9. Podocoryne fulgurans. Views of manubrium showing medusa-
buds. Agassiz Laboratory, Newport, Rhode Island, September,
1892.

Drawn from life, by the author.

fin

x

ANTHOMKDU.S.E PANDEA.

II '.I

ubrium is slender and about half as long as the depth of the bell-cavity. 4 small lips. 4
longitudinal gonads on the stomach. These are narrow folds with smooth outer surfaces.
Bell light-pink, with the 8 adradial lines of nematocysts more intense in color. Stomach and
tentacle-bulbs light-brown.

Found in Sydney Harbor, New South Wales, Australia, in August and September.

Pandea violacea Agassiz and Mayer.

Plate ll, fig. 7; plate 12, fig. i.

Pandra violacea, AGASSIZ, A., and MAV>R, iX<)<l, Bull. Mus. Comp. <>!. at Harvard C'"llrt;i\ vol. ;2, p. i(>o. M\vtR, 1900,

Bull. Mus. Comp. Zocil. at Harvard College, vol. 37, p. 34, plate i, fig. i.
Pandea, s/>., LoBiANCo, 1903, Mitth. Zoolog. Sta. Neapel, Bil. 16, p. 217, lav. 7, fig. i.

Bell pear-shaped and 7.5 mm. high, with moderately thick
walls. There are about 32 tentacles. 8 to 12 of these are
about 3 times as long as the bell-height and 24 are small and
rudimentary. The long tentacles are all of equal length. Their
basal bulbs are long, tapering, and hollow. There are i to 3 rudi-
mentary tentacle-bulbs between each successive pair of long ten-
tacles. There are about 32 ectodermal, purple ocelli, one on tin-
outer side of each tentacle-bulb. The velum is well developed.
The manubrium is flask-shaped and quadratic in cross-section.
The outer surfaces of the adradial gonads are smooth and with-
out corrugations (plate n, fig. 7). The mouth is at the extrem-
ity of a well-developed neck and is at about the level of the velar
opening. There are 4 large, slightly recurved lips with smooth,
simple edges. There are 4 straight, narrow radial-canals and a
broad circular vessel. The entoderm of the manubrium and ten-
tacle-bulbs is delicate pink and in some specimens a green streak
extends along the outer surface of the entodermal lining of the
radial-canals. This medusa is common at the Tortugas, Florida,
and among the Bahamas throughout the summer. An appar-
ently identical species is found at Suva in the Fiji Islands, South
Pacific, although the Pacific medusa is not so highly colored.

A closely related, if not identical, medusa is described by
Lobianco, 1903, from two specimens drawn from depths of 500 to
600 fathoms near Capri, Bay of Naples, Italy. Dr. Lobianco
kindly permitted me to studythese specimens. The bell is 1 1 mm.
high, 7 mm. wide; thick-walled, with dome-like apex. There are
12 to 13 long tentacles, 4 to 5 times as long as the bell-height.
These have long, hollow, tapering basal bulbs, each with an
abaxial, dark-red ectodermal ocellus. In addition to the long
tentacles there are about 24 to 36 very short rudimentary marginal
tentacles, each with an abaxial ocellus. The 4 radial-canals are
quite wide and with slightly jagged outlines. The manubrium
is half as long as the depth of the bell-cavity. The 4 complexly
folded lips are at the end of a well-developed neck. The outer
surfaces of the adradial gonads are smooth. The entoderm is
strawberry-pink. I can not separate this medusa from Pandea
violacea of Tortugas, Florida, some specimens of which are fully
as pink in color as is the one shown in Lobianco's figure.

FK;. 64. Pandra vtolacea, after
Lobianco, Mitth. Zool. Sta.
Neapel, 1903.

Pandea maasi.
Tiara, j/)., MAAS, 1904, Result. Camp. Sci. Prince dc Monaco, fasc. 28, p. 13, plate 2, fig. M.

Bell 10 to 13 mm. high and only about half as wide as high, with a smooth exumbrella
surface. Bell-walls quite thick, with an evenly rounded apex, without an apical pro]i-ctmn.
4 thick, hollow tentacles at the bases of the 4 radial-canals. These have large, swollen basal
bulbs, but no ocelli were observed. The tentacles are about as long as the bell-height and

120 MEDUS.E OF THE WOULD.

their shafts are of about uniform width throughout, the outer ends being very blunt, not taper-
ing. There are no other tentacles or marginal appendages. The 4 radial-canals and the ring-
canal are simple, narrow, and straight-edged; without glandular diverticula.

The stomach is wide and barrel-shaped to cylindrical and is about three-fourths as long
as the depth of the bell-cavity. There is no peduncle and the 4 radial-canals run directly into
the stomach without enlarging as they approach its base. The mouth is well developed, but
the lips are simple, without the foldings commonly seen in other Tiarinae.

The gonad is developed as an open network of more or less transverse folds over the ecto-
dermal wall of the stomach, excepting only the neighborhood of the mouth. The gonad is not
distinctly separated into 4 interradial parts, but is more or less fused over the perradial lines,
thus encircling the stomach very much as in medusae of the Codonidae or in Pandca con i en.

The entoderm is yellow, the gonads being deep in color and the tentacles lighter. Other
parts are transparent.

Found by the Prince of Monaco at Bear Island, between Norway and Spitsbergen, in
July, 1898. It is well figured and described by Maas, 1904. Hartlaub, 1907, considers this
medusa to be identical with Sarsia flammea Hartlaub. The general proportions of the two
medusae appear to be much alike, but the details of structure of the gonads and the color of
P. maasi appear to distinguish it from other medusae.

Genus TURRIS Lesson, 1843, sens, amend.

Turr'n+Tiara (in part), LESSON, 1837, Prodrom. Monog. Meduses, Nos. 17, 20 (not published); 1843, Hist. Zooph. Acal.,

pp. 283, 286.

Turris + Oceania (in part), FORBES, 1848, British Naked-eyed Medusae, pp. 21, 26.
Turrh+ Tiara, AGASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 346, 347. HARTLAUB, 1892, Nachricht. kgl. Gescll.

Wissenschaft. Univ. Gottingen, pp. 20, 22.
Turris, AGASSIZ, A., 1865, North Amer. Acal., p. 164.

Tiara+ Turris+ Catablema, HAECKEL, 1879, Syst. der Medusen, pp. 56, 60, 62.
Turris, FEWKES, 1881, Bull. Mus. Comp. Zoo], at Harvard College, vol. 8, p. 147.
Tiara, WAGNER, 1885, Wirbellosen des Weissen Meeres, p. 78. MAAS, 1905, Craspedoten Medusen der Siboqa Expedition,

Monoe. 10, p. 14. BOVERI, 1890, Jena. Zeit. Naturw., Bd. 24, p. 339. (The egg gives rise to 2 polar globules.)
Cu1ablema-\- Turris + Tiara, VANHOFFEN, 1891, Zool. Anzeiger, Bd. 14, p. 444.
Catablema+ Tiara+ Turris, MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, pp. 12, 15, 16.

GENERIC CHARACTERS.

Tiannae with 4 or more marginal tentacles. With 4 interradial horseshoe-shaped gonads
in the stomach-wall. Each horseshoe composed of partially fused swollen ridges.

Lesson, 1843, describes three medusae under the name Turns, only one of which belongs
to the genus as we define it. This one is Turns neglecta of the north Atlantic coasts of Europe.
The name Tiara was also proposed by Lesson, 1843, and is used to describe a medusa which
was first observed by Forskal, 1775, under the name Medusa pileata. It is found off the
Atlantic coast of Europe and in the Mediterranean.

I use Turris as equivalent to Turris Lesson + Tiara Lesson + Catablema Haeckel.
Haeckel, 1879, p. 62, establishes Catablema for a Greenland medusa which he calls Catablema
campanula.

Maas, 1904, who embodies the results of the studies of Vanhbffen, Hartlaub, and himself,
defines Turns as having horseshoe-shaped interradial gonads, the sides of the horseshoe
being made up of partially fused, transverse ridges while the arch connecting the two sides is
composed of an irregular network of ridges. In Tiara the network is lacking, the entire
horseshoe being composed of more or less transverse ridges. The ridges do, however, anas-
tomose to some extent, so that an imperfect, partial network may be formed. Catablema he
distinguishes by the jagged edges of its radial-canals and circular vessel. However, medusa;
of Turns and Tiara often display lagged edges upon their canals, and this character is very
variable and subject to much individual irregularity in development. In the young medusa
of Catablema the ridges forming its interradial horseshoe-shaped gonads tend more or less
longitudinally, but in later development they come to he almost transversely as in Turns
or Tiara. In order to terminate the confusion that has been introduced by these intergrading
and too precise criteria for distinguishing the genera Turris, Tiara, and Catahlrma, I propose
to unite them all under one genus and call it Turns, the name first used by Lesson to dis-
tinguish any of these medusae.

ANTHOMKUUS.E '1THUIS.
Tubular Synopsis of the Species of Turris.

121

T. papua Lesson.

T.rotunda = Tiara
rotunda Haeckel.*

T.prismatica =
Tiara prismatica
Maas.f

T.reticulata
Haeckel.

T. vesicaria
A. Agassiz.

Size of bell in mm.

28 high, 14 wide.

Pointed apex.

20 high, 20 wide.

Cubical.

20 high, 12 wide.

12 high, 10 wide.

25 high, 12 wide.

Number of tenta-
cles.

8 to 32

8

12 to 1 6

16

16 long, 48 short.

Ectodcrmal, ab-

Present.

Prcbcnt.

?

Present.

Present.

axial ocelli on

tentacle-bulbs.

Character of edges

Serrated.

Smooth.

Serrated.

Smooth.

Jagged.

Character of
gonads.

4 interradial horsc-
shoes of complex,
mainly transverse
swellings which
anastomose to
some extent.

4 pairs of adradial
groups of cross-
folds.

As in T. pilcMta.

8 separate triangular
areas in adradii of
stomach, with
complex anasto-
mosing ridges.

Swollen, transverse,

papilliform,
anastomosing
folds forming net-
work in 8 adradii
of stomach.

Color.

Tentacle-bulbs an.l

Tentacle-bulbE]

?

?

Stomach and ten-

stomach pink to
dark-red. Gonads
pink to dark-
brownish-yellow.

stomach, and
gonads carmine.
Radial-canals and
ring-canal yellow.

tacle-bulbs cin-
namon-brown.
Gonads darker.
<krlli dark-red.

Where found.

Pacific and Indian

Straits, of Gibraltar.

Gulf Stream, At-

South Atlantic nc.u

An tic Ocean to

Oceans in tropical
regions.

lantic.

Tristan
d' Acunha.

southern coast of
New England,
United States.

T.superba Mayer .J

T.coeca Hartlaub.

T. pelagica Agassiz
and Mayer.

T. conifera= Tiara
conifera Haeckel .J

T.campanula =
Catablema cam-
panula Haeckel.

Size of bell in mm.

^ high, 3.5 wide.

^o high, 20 wide.

16 high, 9 wide.

25 high, 2O \v ulr.

20 high, 20 wide.

Number of tenta-
cles.

4 long, 12 short.

24 to 30 of various
lengths.

30 short. All of
same length.

24 to 48. All long.

24 to 48* All long.

Ectodermal, ab-
aiial ocelli on

Present.

Not present.

Not present.

_'

Not present.

tentacle-bulbs.

Character of
edges of radial-
canals.

Smooth.

Jagged.

Jagged.

?

Jagged.

Character of

gonads.

4 interradial horse-
shoes of partially
fused, mainly
transverse ridges.

AsinT. superba.

As in T. vesicaria.

Simple transverse
ridges across the
8 adradii of
stomach.

4 crescent-shaped
gonads in the 4

intcrradii of
stomach. Each
composed of
longitudinal
ridges side bv
side. No trans-

verse folds.

Color.

Stomach emerald-
green and pink to
rose. Tentacle-

Stomach pink,
rose-colored,
wine-colored.

All entodermal parts
light port-wine-
colored.

p

Entodermal parts
yellow.

bulbs rose-color. Tentacle-bulbs

Entire gelatinous , yellow.

substance delicate

*

pink.

\VlnTr found.

Tortugas and
Bahamas in sum-

Mediterranean.

Pacific coast of
Lower California.

Arctic Ocean,
Russia, Green-

Arctic Ocean.

mer.

land.

* Probably an abnnrmal or stunted T. fiilrata.
J Probably ulrntical with T. pilcala.

s this an abnormal individual of T. fileala (?)

122

MEDUS.E OF THE WORLD.

According to Wright, 1859 (Edinburgh New Philosoph. Journ., plate 8, fig. i) and
Hincks, 1868 (British Hyd. Zooph., p. 14, plate 3, fig. l), the egg of Turris neglecta develops
into a Tubulanan hydroid. Clavula gossei (see also Allman, 1871, Monog. Tubularian
Hydroids, p. 259; also Metschnilcoff, 1886, Arbeit. Zool. Inst., Wien, Bd. 6, p. 239) found
that the eggs of Tiara pileata develop into a clava-like hydroid polyp. The hydroid polypites
of Turris (Clavula) arise singly from a creeping stolon. Each polypite is slender, elongated,
and claviform with scattered, elongate tentacles. The coenosarc is invested by a polypary.
It is possible that the hydroid described by Gegenbaur, 1854 (Verhandl. phys.-med. Gesell.
Wiirzburg, Bd. 4, p. 165, taf. I, figs. 3, 4) may prove to be the hydroid of Turris. The short,
clavate polypites arise separately from a somewhat branching stolon, and each has 5 to 8
scattered, slender tentacles. The medusa-buds arise from the stolon and when set free each
medusa has 2 long and 2 immature tentacles, and a line of nematocysts extends up from each
tentacle-bulb to the top of the bell. Gegenbaur calls this form Clcodoru tricuspiJata or
Syncoryna cleodorir. Allman, 1871 (Monog. Tubularian Hydroids. p. 260) calls it Cam-
paniclava.

It is probable that certain species of Perigonimus give rise to medusae of the genus Tun-is.

Corydendrium minor Nutting, 1905 (U. S. Fish Commission Bulletin for 1903, p. 941,
plate 2, fig. I ; plate 7, figs. 8, 9) from the Hawaiian Islands may be the hydroid of some Turris.

Turris neglecta Lesson.

Kfnnin beroe, SLABBER, 1778, Physikal. Belustig., Niirnberg, p. lio, plate 13, fig. 3.

Turris neg/ecla, LESSON, 1843, Hist. Zooph. Acal., p. 284. FORBES, 1848, British Naked-eyed Medusa 1 , p. 23, plate 3, figs.

za-2.1.

Oceania globulosa (young?), FORBES, 1848, British Naked-eyed Medusa 1 , p. 29, plate 3, fig. 3.
Turris neglecta (hydroid ?), GOSSE, 1853, Nat. Rambles Devonshire Coast, p. 348, plate 13, figs. 6-10.
Clavula gossei, WRIGHT, 1859, Edinburgh New Phil. Journ, plate 8, fig. l.
Turris neglecta, HINCKS, 1868, Hist. British Hydroid Zooph., p. 13, plate 3, fig. I. ALLMAN, 1872, Monog. Tubularian Hydroids,

p. 259. HAECKEL, 1879, Syst. der Medusen, p. 62. BEDOT, 1901, Revue Suisse de Zool., tome 9, p. 488, Ibid., 1905,

tome 13, p. 152 (all literature to 1850).

Tabular Synopsis of the Species of Turns Continued.

T.eurystoma = Cat-
ablema eurystoma
Hacckel.

T. pilrata = Tiara
pileata L. Agas-
siz.*

T. neglecta Lesson.-]

T. digitalis Forbes.

T.brevicornisMur-
bach and Shearer.

Size of bell in mm.

40 high, 20 wj e.

40 high, 25 wide.

6 high, 4 wide.

40 high, 20 wide.

45 ni 'g h <35 wide.

Number of tenta-
cles.

24 to 48. All long
and 24 to 32 ten-
tacle-bulbs.

12, to 48. Usually
241032. All well
developed.

60 to 70 short.

50 to 100 short.

140 +

Ectodermal, ab-
axial ocelli on

Not present.

Present.

Present.

?

Not present.

tentacle-bulbs.

Character of edges
of radial-canals.

Jagged-

Slightly notched
and jagged.

Jagged.

Jagged.

Character of
gonads.

Developed in longi-
tudinal curtain-
like folds of in-

4 interradial horse-
shoes of mainly
transverse, some-

As in T. pileata.

8 adradia rows of
transverse ridges.

As in T. pileata.

terradial sides of

what anastomos-

short stomach.

ing ridges.

Color.

Stomach and
gonads cherry-
red. Canals and
tentacles bright-
red.

Variable. Stomach
almost colorless,
yellow, red, purple,
brown. Radial-
canals colorless,

Stomach, gonads,
tentacle-bulbs
scarlet, purple.

Stomach purple,
reddish-brown.
Tentacle-bulbs
yellow, orange.

Tentacles, gonads,
and stomach dark-
red, purple.

yellow, green.
Tentacles color-

less, pink.

Where found.

Coast of Greenland.

North Atlantic to

British coasts.

Greenland to North

St. Paul Island,

Mediterranean.

Sea, British

Pribylofflslands.

coasts.

North Pacific.

*T. conifera rotunda, superba and prhmatica are probably identical with T. pileata. Hydroid: Perigonimus ref>ens?
(Hydroid: Clavula gossei Wright.

PLATE 13.

Fig. i. Protiara formosn, mature medusa. Tortugas, Florida, May 21, 1907.

Oral view of female.
Fig. 2. Protiara forniosa. One of the marginal cirri showing the chordate

entodermal cells.
Figs. 3 to 5. Podocoryiif fulgnraiis. Agassiz Laboratory, Newport, Rhode

Island, September, 1892.
Fig. 6. Turns pileata, young medusa. Agassiz Laboratory, Newport,

Rhode Island, September 25, 1896.
Fig. 7. Turris vesicaria, mature medusa. Eastport, Maine, August, 1897.

Drawn from life, by the author.

PLATE 13

.

ANTHOMEDU8/E TDRRI8.

123

Bell 4 to 6 mm. high, 5 to 4 mm. wide. Globular or with a conical apex and bulging
sides; margin somewhat constricted. 60 to 70 short, crowded tentacles, shorter than the

bell-height, with abaxial ocelli. Stomach wide, cruciform in
cross-section, and one-half to two-thirds as long as the depth
of the bell-cavity. 4 simple lanceolate lips, not complexly
tolded. Gonads as in T. pilrtitu. Entoderm of stomach
and tentacle-bulbs reddish-brown to rich purple. Found
or!" the British coasts.

The hydroid was obtained by Grosse and Wright, who
reared it from the egg. The planula is bright crimson.
The hydranths are club-shaped and only about 2.5 mm. high.
They arise singly at intervals from a creeping filiform h\dm-
rhiza and have 12 or more scattered, filiform tentacles. The

Fn.. 65. Hvdroid of
' Hincks,
Zoophytes.

rj negfecta, after hydrocaulus and hvdrorhiza are invested by a perisaic.

Hincks, in British Hvdroid .; .

Hydranths bright crimson.

Turris pileata.
Plate n, fig. 4; plate 13, fig. d.
Medusa pileata, FORSKAL, 1775, Descriptions Animal., p. no; 1776, Icones rerum naturalium, plan- 33, fig. I).

ania pileata+ P. Lesueur, PERON ET LESUEUR, 1809, Ann. du Museum d'Hist. Nat., tome 14, p.

nia (Charvbdea) pisifera, OKEN, 1815, Lehrbuch iler Naturges., Tril 3, p. 125.
ania ampitHacea, SARS, 1835, Beskriv og Jagttagelser, p. 22, plate 4, fig. fta-f.
ra f>,ifia/ii+T. sarsii, LESSON, 1843, Hist. Zooph. AcaL, p. 287.

ania turrita+0. octona+O. episcopates, FORBES, 1848, British Naked-eyed Mediis.r, pp. 27, 28, plate 2, figs. 1-3.
ania coccinea, LEI'CKART, 1856, Archiv. fur Naturges., Jahrg. 22, p. 24.
alidium arnpullaceum + Tiara pileata, A<;ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 352, 347.

0:

T

Or

Or

Ph

Tiara smara^dina, HAECKEL, 1864, Jena. Zeit. fiir Naturvv., Bd. i, p. 336.

Oceania pileata, SPAT.NOLINI, 1876, Catalogo Acalefi Mediterraneo, p. 21, tav. 3, fig. i, 2.

Tiata pileata, HAECKEL, 1879, Syst. der Medusen, p. 58, taf. 3, fign. 6-8. HAMANN, 1883, Zeit. fiir wissen. Zoo!., Bd. 38, p. 426,
taf. 23, fign. 16-20 (development of the planula larva). METSCHNIKOFF, E., 1886, Embryol. Sludien an Medusen, pp.
29, 48, etc., taf. l, fign. 1-17. HARTLAUB, 1894, Wissen. Meeresuntersuch. Komni. Mrrrr Ku-l, Helgoland, Neue Folge,
Bd. l, p. 189. FORSKAL= Oceania epiicopalis FORBES; BROWNE, 1896, Irish Naturalist, p. 180; 1895, Proc. and
Trans. Liverpool Biol. Soc., vol. 9, p. 258. BEDOT, 1901, Revue Suisse de Zoo]., tome 9, p. 487; Ibid., 1905, tome 13,
p. 150 (all literature to 1850). BROWNE, 1903, Bergens Museums Aarbog, No. 4, p. 11. MAAS, 1904, Result. Camp
Sci. Prince de Monaco, fasc. 28, p. 15, plate l, fig. 9. BROWNE, 1905, Proc. Royal Soc. Edinburgh] vol. 25, p. 760.

(?) Tiara intermedia, BROWNE, 1902, Annals and Mag. Nat. Hist., ser. 7, vol. 9, p. 277 (Falkland Islands, South Atlantic).

Gervonia octona, FLEMINC, 1823, Edinburgh Philosoph. Journ., vol. 8, p. 299.

Oceania octona, FORBES, 1848, British Naked-eved Medusa-, p. 27, plate 2, figs. 3^-3^. The plate is wronglv laln-lt-d *'O. epis-
copal is ."

Oceania turrita (young medusa), FORBES, 1848, British Naked-eyed Medus.e, p. 28, plate 2, figs, la-zc.

Tiara octona, A<;ASSIZ, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 347.

Oceania coronala, ALLMAN, 1871, Monog. Tubularian Hvdroids, p. 33, fig. 8.

Tiara octona, H^FCKEI., 1879, Svst. der Medusen, p. 57.

Yon Tiara octona, MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, p. 13, plate 2, fig. ll.

Bell about 15 to 40 mm. high and 10 to 20 mm. wide. Side walls quite thin, but there is
usually a well-developed, solid, apical projection, although this may lie absent. This pro-
jection may be conical or cylindrical, with a basal constriction, or pineapple-shaped, etc.
There are 12 to 48, usually 24 to 32, tentacles with hollow, laterally compressed, tapering
basal bulbs. These tentacles are longer than the bell-height and there is an abaxial ocellus
on the outer side of the base of each bulb. The velum is narrow and simple.

The 4 radial-canals and ring-canal are wide, flat, and ribbon-like and their edges are often
more or less notched and jagged. As they approach the sides of the stomach the radial-canals
widen out into funnel-like expansions, so that they embrace the upper halves of the sides of
the stomach in the 4 principal radii.

The stomach is wide, balloon-shaped, and 4-sided, and fills the upper one-half to two-
thirds of the bell-cavity. The 4 lips are at the end of a relatively- narrow, short neck and
are complexly folded, crenated and recurved upward.

The gonads are 4 horseshoe-shaped regions on the sides of the stomach, the sides of each
horseshoe being adradial and the apex aboral and interradial. I he horseshoe consists of an
area of more or less transverse folds which, however, anastomose more or less so as to form
a network of ridges. In young medusz the ridges of the gonads are more or less parallel
and transverse, but in later life they anastomose to form an irregular network.

124

OF THE WORLD.

This medusa is very variable in color. The stomach ranges from yellow to red, or from

brownish-red to purple. The radial and circular canals may be colorless, to yellow or green.

The ocelli are red or dark brownish-purple. The tentacles range from colorless to light-purple.

It is widely distributed over the North Atlantic and in the Mediterranean, but it

appears to be much commoner on the European than along the American coast.

The development has been studied by Hamann, 1883, and Metschnikoff, 1886. Seg-
mentation is total and equal and a single-layered, elongate, ciliated larva is found. The
entoderm develops from cells which wander into the central segmentation cavity from the
hinder end of the larva, so that finally a two-layered planula is formed. The planula attaches
itself and the hinder end develops into a hydrorhiza, while the forward end becomes the
polypite. The hydrorhiza becomes invested with a thin pensarc and the polypite develops

a mouth and 2 or 3 long thin tentacles, which arise at the base
of the conical throat-tube. This stage was reached at the
end of 16 days in Metschnikoft's specimens, and apparently
the hydroid is one ot the Clavidae.

This hydroid may possibly be the very abundant and
widely distributed Pengonimits rrpens (fig. 66) found oft the
Atlantic coasts of Europe from Russia southward, in the
Mediterranean, off the Pacific coast of the United States,
and at the Falkland Islands and Terra del Fuego, South

o

America.

The stems of the hydroid branch and are covered with
a gelatinous investment to which foreign bodies become
attached. The polypites are spindle-shaped, with a single
oral circlet of tentacles. The medusa-buds arise singly from
the sides of the stems.

According- to Hartlaub and Browne the medusa is set

P>

free with but 2 tentacles and 2 small tentacle-bulbs. Forbes
described the 4-tentacle stage as Oceania titrnta. the 8-ten-

o

tacle condition as O. octona, and the medusa with 12 tentacles

as 0. episcopalu.

Some of the recent literature relating to the hydroid P.
repens is here presented, though it must be borne in mind that this has not been proven to
be the hydroid of the medusa in question.

Pengonimits re pens + P. minutus, ALLMAN, 1872, Monog. Tubul. Hydroids, p. 521, plate II.

Perigommus repens, GRAEFFE, 1884, Arbeit. Zool. Inst. Wien., Bd. 5, p. 350. (At Trieste, Adri-
atic. Rare. Medusae produced in May). DU PLESSIS, 1888, Recueil Zool. Suisse, tome 4,
p. 539. BROWNE, 1896, Proc. Zool. Soc. London, p. 462 (P. minutus ALLMAN = P. repens).
BONNEVIE, 1899, The Norwegian North Atlantic Expedition, 1876-1878, vol. 26, Hydroids,
p. 41 (from Norway). CALKINS, 1899, Proc. Boston Soc. Nat. Hist., vol. 28, p. 539, plate
I, figs. 3-3 D (from Puget Sound). TORREY, 1902, California Univ. Pub. Zool., vol. I,
p. 29 (Pacific coast, California to Washington). HARTLAUB, 1905, Zoolog. Jahrbiichern,
Suppl. 6, p. 530, fign. K, L, M (from the Falkland Islands and Terra del Fuego).

Turris rotunda.

l>i,in,ra rotunda, QUOY ct GAIMARD, 1827, Annal. des Sci. Nat., tome 10, p. 181, plate 6 A, figs, i, z.

Tiara rotunda, HAECKEL, 1879, Syst. der Medusen, p. 57, taf. 3, fign. 9, lo. BEDOT, 1905, Revue Suisse de Zool., tome I},
p. 151 (all literature to 1850).

Bell cubical without a pointed apex; 20 mm. high, 20 mm. wide. Bell-cavity only two-
thirds as deep as the bell-height. 8 tentacles much longer than the bell-height, with conical
basal bulbs. A large, dark-red ocellus on the outer side of each tentacle-bulb. Stomach
small, swollen, globular, and confined to the upper third of the bell-cavity. The lips are short,
much folded, and with a row of nettling warts around their edges. The gonads consist of
4 pairs ot longitudinal, swollen areas, each thrown into 8 to 10 cross-foldings, on the adradial
sides ot the stomach. The stomach, gonads, and tentacle-bulbs are carmine, lips rose-red.

1 i'.. i<>>. Perigonimus repent, after Him k
in British Hydroid Zoophytes.

.\\TIHtMKIirs.K TI'KlilS. I!'.".

Canals yellow. Found by Ouoy and Gaimard, and later by Haeckel, in the Straits of Gib-
raltar. Development unknown.

Turris papua Lesson.

Tuna papua, LFSSON, 1843, Hist. Zooph. Acalephes, p. 283; Prodromus, 1837, No. 36. EYDOI x I:T SOILEVKT, 1X41.
Voyage de la Bonne, tome 2, p. (t\q, plate i, figs. 1-3. HAECKKL, 1879, Sysl. der Meduscn, p. 58. MAAS, 1905, ( i
doten M.-dusen der Xibaga Expedition, Monog. 10, p. 14, taf. l, fig. 13; 1906, Kevue Smsse ilr Zool., tome 14, p. 8X.
BF-UOT, 1905, Revue Suisse de Zool., tome 13, p. 150 (all literature to 1850).

.fyuorea tniiTti, LESSON, 1829, Voyage de la Coqiiille, Zool., p. 127, plate 14, fig. 4.

(?) Tiara ocranica, AGASSIZ, A., and MAYER, 1902, Mem. Mus. Comp. Zool. at Harvard College, vol. 16, p. 141, plate I, fig. I.

(?)Tf<vd intermedia, BROWNE, 1902, Annals and Mag. Nat. Hist., MT. 7, vol. 9, p. 277.

Bell 28 mm. high, 14 mm. wide, with a solid apical projection and thin vertical walls. The
longest tentacles are about as long as the hell-height and have elongate, conical, basal bulbs,
with dark-red ectodermal ocelli on their outer sides. According to Lesson and to Eydoux et
Souleyet there are 8 tentacles: 4 radial, 4 interradial. Maas found 16 tentacles: 4 radial, 4
interradial, and 8 adradial. Agassiz and Mayer tound 32 tentacles: 8 long radial and
interradial, 8 short adradial, and lu very short intermediate tentacles. These differences may
he local variations, but can hardly be due to the medusae being in various stages of develop-
ment, for Agassiz's specimen was only 5 mm. high and had 52 tentacles, while Browne's
and Maas's specimens were 7 mm. high with only 16 tentacles. Moreover, Lesson's medusae
were larger and had only 8 tentacles. The 4 radial-canals are quite wide and have serrau-il
edges. The stomach is wide, swollen, and the lips complexly folded, and about at the level of"
velar opening. The gonads are in 4 interradial, horseshoe-shaped series of transverse swell-
ings on the sides ot the stomach, the open ends ot the horseshoe being directed downward
toward the mouth and the apex being near the aboral end ot the stomach.

The entoderm of tentacle-bulbs and stomach is pink to dark-red. The gonads are pink t<>
dark brownish-yellow.

Widely distributed over the Indian Ocean and tropical Pacific.

It is probable that this form displays considerable variability in the number of its tentacles
and in the color of its gonads. It is closely allied to 'Turns superb a ot Tortugas, Florida, and
mav be identical with T. intermedia Browne, from the Falkland Islands.

Turris prismatica.
Tiara pTiimalifa, MAAS, 1893, Ergeh. der Plankton Exped., Bd. 2, K. c.. p. f8, taf. 6, fign. 10, 1 1.

Bell about 20 mm. in height, 12 mm. in diameter; 4-sided and prismatic in shape. There
is no apical projection and the bell-walls are remarkably thick. In the single specimen
described by Maas there are 12 tentacles, one at the foot of each of the 4 radial-canals. 3
others in one quadrant, 3 in another, and one in each of the other quadrants. It seems probable
that the specimen is abnormal and that there are normally in tentacles. The tentacles arise at
a slight distance above the bell-margin. Ocelli ( ?) The gonads are similar to those ot Tuna
pilnitii. In the preserved specimen the bell has a 4-sided prismatic shape. Color ( ?) A
single individual of this species was found by the Plankton Expedition on August 4, 1 880., in the
Gulf Stream. May it not be an abnormal specimen of Turris pileata ( ?)

Turris reticulata Haeckel.

Tiara retifitlala, HATI-WEI., 1879, Syst. der Medusen, p. 60, taf. 3, fig. 1 1 .

Medusa bell-shaped, 12 mm. high and 10 mm. broad. There is a small, conical, apical
projection, about 1.25 times as high as it is broad. The stomach is 4-sided and pyramidal,
widest below, and its radial edges are bound throughout their length by the 4 radial-canals.
The 4 lips are large and much folded. There are 16 tentacles which are longer than the width
of the bell, and have large, laterally compressed bases, with ocelli on their outer sides. Their
are 8 separate, triangular gonads in the outer wall of the stomach. The surfaces ot the gonads
display complex anastomosing folds and swellings.

This species was described by Haeckel from preserved specimens obtained in the South
Atlantic near the Island ot Tristan d'Acunha.

12f) MEDUSA OF THE WOULD.

Turris vesicaria A. Agassiz.

Plate ll, figs. 2 and 3; plate 13, fig. 7.

Turris vesicaria, AC.ASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 97; 1865, North Amer. Acal., p. 164, figs. 261-268.

NUTTING, 1901, Bull. U. S. Fish Commission for 1899, vol. 19, p. 375, fig. 86 (abnormal medusa).

C.nlablema vesicarium, HAECKF.L, 1879, Syst.der Medusen, p. 64. GRONBERG, 1898,200!. Jahrb., Abth. Syst., Bd. n, p. 461.
Turris efitcopalit, FEWKES, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p. 147, plate 3, figs. 1-6.
Turris vesicaria, and T. efiscofalis, HARC.ITT, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 36, 2 figs.
Catahlema I'tsicarium (young medusa), MAAS, 1904, Result. Camp. Sci. Prince de Monaco, fasc. 28, p. 12, planches I, 2.

Adult medusa (plate 13, fig. 7). Bell about 25 mm. in height and 12 mm. in diameter.
There is a large, solid, apical projection which varies greatly in shape, being globular in some
individuals, while in others it is cylindrical or conical. Its upper end is often seen to terminate
in a small button-shaped projection. The side walls of the bell are quite thin and flexible.
There are 16 well-developed tentacles, the basal bulbs of which are quite large and are
flattened in a radial direction, so that the tentacle appears to arise from the side of the bell
slightly above the bell-margin. These well-developed tentacles are very flexible, and when
ejctended are about 2 to 4 times as long as the bell-height. These basal bulbs are hollow and
bear each an abaxial ocellus. In addition to these long tentacles there are 48 very short, rudi-
mentary tentacles, which arise from the bell-margin at a slightly lower level than the long ten-
tacles. There are 3 of these rudimentary tentacles between each successive pair of long
tentacles. There is a single dark-red ocellus upon the outer side of each and every tentacle-
bulb, 64 in all. The velum is narrow. There are 4 broad, flat radial-tubes and a broad,
circular canal, all of which display jagged edges. The 4 radial-canals enter the stomach by 4
wide, funnel-like openings. The manubrium is large and urn-shaped and there is no peduncle.
The mouth is surrounded by 4 prominent, crenulated lips. The gonads occupy 4 interradially
situated, longitudinal regions upon the walls of the stomach. Each gonad consists ot a pair
of swollen, papilliform, adradial, ectodermal ridges, the outer surfaces of which are thrown
into complex, but mainly transverse, folds and corrugations. The color of the manubrium and
tentacle-bulbs is cinnamon-brown, the gonads being of a somewhat darker shade. The
ocelli upon the tentacle-bulbs are dark-red.

y otnig medusa (plate 12, fig. 2). In the youngest medusa observed the bell is about 2 mm.
in height. The walls are thin, and there is a short, conical, apical projection. There are only
4 well-developed tentacles, one at the base of each of the radial-canals. These tentacles have
large conical basal bulbs which are hollow. The axes of the tentacle-bulbs are at a slight dis-
tance above the bell-margin. In addition to the 4 well-developed, radial tentacles, there are 4
intermediate interradial tentacles which arise from the bell-margin at a slightly lower level than
the radial tentacles. The 4 radial-canals are broad and their edges are jagged. The manubrium
is large and quadratic in cross-section. The lips are simple.

In a medusa 3 mm. in height (plate 12, fig. 3), there were 8 well-developed and 8 immature
tentacles. The apical projection was large and miter-shaped, and the lips had folded, notched
margins.

This medusa is probably an Arctic species. It has been found but rarely south of Cape
Cod, and then only in May and June. It is abundant, however, in the harbor of Eastport,
Maine, in August, and ample opportunities for observing its growth were afforded.

The medusa is remarkable for its voracity and will devour Nanomia cara with great
avidity. It is a common thing to observe the medusa in the act of devouring one of these large
Siphonophores which may be many times its own size. Maas, 1904, records this medusa off"
Bear Island, between Norway and Spitzbergen, so that its distribution is probably circum-
polar. Grcinberg records it as being rare in August off" the coasts of Greenland and Spitz-
bergen.

Turris superba (a variety of T. pileata).

Plate 27, fig. 8; plate 28, figs. 3 and 4.

Tiara superba, MAYER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 34, plate 16, fig. 39; Memoirs Nat.
Sci. Museum Brooklyn Inst. Arts and Sci., vol. I, No. I, p. 8, plate 2, fig. 1 1.

Bell 5 to 7 mm. high and with a well-developed apical projection. There are 4 long,
hollow and 12 small, solid, rudimentary tentacles. The long tentacles are slender, with taper-
ing basal bulbs. There is an ectodermal ocellus upon the outer side of the tentacle-bulb of

ANTHOMEDU8.* TURRI8.

127

each of the 1 6 tentacles. The velum is well developed. There are 4 broad, flat, smooth-edged
radial-canals and a broad, simple, circular vessel. The 4 radial-canals enter the stomach by
4 wide tunnel-like openings. The manubrium is wide and tills the greater part of the bell-
cavity. The 4 lips are recurved and their edges much tolded. The 4 gonads are developed
on the interradial sides of the stomach. Each gonad is horseshoe-shaped, the apex of the
horseshoe being uppermost, and the sides composed of partially fused, transverse ridges upon
the ectoderm of the stomach-wall. The entire gelatinous substance of the medusa is of a
delicate rose-pink.

The entoderm of the manubrium and tentacles is of a rich rose-color and the entodermal
nut' of the stomach is emerald-green. This medusa is quite common at Tortugas, Honda,
and among the Bahamas in the summer months. It is one of the most beautiful of American
medusae.

Turris breviconis Murbach and Shearer.

Turrit hrei-iconis, Mi'RBAfl! AND SHKAKKK, 1902, Annals ami Mag. Nat. Hist., ser. 7, vol. 9, p. 73; 1903, Proc. /..nl. s.u .
London, vol. 2, p. 170, plate 18, figs. I, 2.

Bell cubical, 45 mm. high, 35 mm. wide. More than 140 tentacles in double rows, with
contractile muscles on their inner sides and enlarged ectoderm on their outer sides. No
ocelli in preserved specimens. Gonads and stomach occupy less than halt of the upper part
of the hell-cavity. 4 broad radial-canals with unbranched lateral diverticula. Bell bluish,
tentacles, gonads, and stomach dark-red or purple.

St. Paul Island, Piihylott' Islands, North Pacific.

Turris coeca Hartlaub.

Turns rrecu, HARTIM-R, 1892, Nachrichten kgl. Gesell. Wissen. Gimingen, p. 19, fig. i. MAAS, 1904, Result. Camp. Sci.

Prince de Monaco, fasc. 28, p. 17, planchr I, fig. 5.
Tinrclhi fHirlhfrifjf'tifti, TRINCJ, 1907, Archiv. Ital. Anat. Embr. Firenze, vol. 5, p. 533 (development and structure of ooevtes).

Bell about 30 mm. high and with 24 to 30 tentacles of various lengths. The tentacle-
bulbs l<ifk ocelli. The radial-canals are wide and flat and there
are a few globular lobes along their edges. The stomach is
wide and the lips complexly folded. The 4 interradial gonads
are composed of horseshoe-shaped swellings. The ridges are
lateral cross-folds in the 8 adradii, but in the mterradius at the
apex of the horseshoe, near the base of the stomach, they form
a reticulated network.

The stomach is rose-colored to wine-colored, and the ten-
tacle-bulbs are yellow.

Found at Naples and Monaco, Mediterranean, in Febru-
ary and March.

Turris pelagica Agassiz and Mayer.

Turrit prla^irn, AGASSI?, A., and MAYF.R, 1902, Mem. Museum Comp. 7.ool. at
Harvard College, vol. 2(1, p. 142, plate \, fig. 2.

Bell id mm. in height and o mm. wide, and the sides are

O *

I'K,. (17. --Gonail of Tmrii creca, after barrel-shaped, being wider at the middle than at either end.
Maas, in Result. Camp. There is a small, solid, apical projection. The bell-walls are
very thin and <|uite flexible. There are about 30 short tentacles,

less than half as long as the bell-height, all being of the same size. These tentacles all arise
from the bell-margin and their bases are large and conical. There are no ocelli. The velum
is well developed. There are 4 radial tubes which are flat and qujte broad, being narrower
near the circular canal than at any other place. Their outer edges are jagged, excepting in
the narrow parts near the circular canal. The manubrium is large and fills the greater
part of the bell-cavity. It is joined to the 4 radial-canals by means of 4 wide funnel-shaped
ducts. The gonads occupy 8 adradially situated rows which extend about two-thirds of the
distance from the inner apex of the bell-cavity to the velar opening. The outer surfaces
of the gonads are transversely folded and give rise to numerous papillae. There are 4 well-

12S MF.nrs.r, OF THF, WORLD.

developed, complexly folded lips. The lips and tentacle-bulbs are of a light port-wine color
.nul the gonads and radial-canals are of a still lighter shade. A single specimen of this
medusa was found in a surface haul on August 26, 1899, off the Pacific coast of Lower Cali-
fornia; N. lat. 51 10', \V. long. 125. It is separated from T. papua and T. [>ilf<ita by the
absence of ocelli.

Turris conifera Haeckel.

Tiara coni frni, HAF.CKFI., 1879, Syst. der Meilusen, p. 59. FEWKES, 1888, Report on Medusa? of the Lady Franklin Bay Ei-
pi-dition, p. 43. LFVINSF.N, 1893, Vid. Meddel. Nat. For. Kjobenliavn (5), Bd. 4, p. i44.--GKijNiir.RU, 1898, Zoolog.
[ahrh., Abth. Svst., Bil. i I, pp. 454, 460. LINK", 1904, Zool. Anzriger, Bd. 28, p. 215.

Bell cone-shaped, with a large, conical, apical projection. It is about 25 mm. in height
and 20 mm. in diameter. There are 24 to 48 tentacles which are longer than the bell-diameter
and which have wide, conical bases. Ocelli ( ?) The manubrium is quadratic in cross-
section and is somewhat longer than wide. The base is wide and cruciform, and the 4 radial
coiners are wide where the radial-canals enter the stomach. The 4 lips are very prominent
and are thrown into complex crenulations and folds. The gonads are composed ot about
10 pairs of regularly arranged, simple transverse or oblique ridges across each ot the 8 adradial
regions of the stomach and are separated in the 4 interradn.

This form is very closely related to Turris pileata, with which it may, indeed, prove
identical. Haeckel says that it is distinguished from both T. pileata and T. reticulata by the
regular feathered gonads and by the smallness of the 4 funnel-like origins of the radial-
canals where they join the stomach. The oral lappets are smaller and less crenulated than
in 7 . pili-titii or T . rrticulatn. The conical, apical projection is half as high as the bell. Color ( ?)

This species is found off the coast of Greenland. Linko, 1904, also found it in Barents
Sea, north of Lapland, and its distribution is, therefore, probably circumpolar in common
with other Arctic species ot Turns.

Turris campanula.

Calablema campanula^ HAF.CKFJ, 1879, Syst. der Medusen, p. 63, taf. 4, fign. 4, 5. LF.VINSEN, 1893, Vid. Meddel. Nat. For.
Kjohenhavn (5), Bd. 4, p. 144. GRONBERG, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 1 1, p. 460. LINKO, 1904, Zool. An-
zeiger, Bd. 28, p. 21?. BF.DOT, 1901, Revue Suisse de Zool., tome 9, p. 482; Ibid., 1905, tome 13, p. 132 (list of refer-
ences to 1850).

Bell about 20 mm. wide and 20 mm. in height. There is a well-developed, conical,
apical projection, which is hollow and contains a prolongation ot the gastric cavity. There
ate 24 to 48 tentacles which are longer than the bell-diameter. Their bases are long and
thick. No ocelli. The 4 radial-canals and the circular tube are wide and flat and have
numerous complex, or simple, blindly-ending side branches resembling those of T. cury-
stomn. The manubrium is wide and shallow and the 4 lips are thrown into complex
crenulations and folds. The 4 gonads are found within 4 crescent-shaped swellings upon
the 4 interradial sides of the stomach. A number of simple, straight, longitudinal folds or ridges
extend across the crescent-shaped gonad; the horns of each crescent are directed downward,
toward the velar opening. The manubrium, gonads, canals, and tentacles are yellow.

This species is found off the coast of Greenland and at Spitzbergen, and Linko found
it in Barents Sea, north of Lapland.

Turris eurystoma.

Calablema eurystoma, HAECKEL, 1879, Syst. der Medusen, p. 64, taf. 4, fign. 6, 7. LF.VINSEN, 1893, Vid. Meddel. Nat. For.
Kjobenhavn (5), Bd. 4, p. 144. GRONBERO, 1898, Zoolog. Jahrb., Abth. Syst., Bd. 11, p. 462.

Bell about 20 to 25 mm. in diameter, and spheroidal, being flatter than it is high. There
is a spheroidal, apical projection which varies in size from about one-fourth to almost as large
as the bell itself. This apical projection is always solid. The sides of the bell are quite thin.
There are about 24 to 48 well-developed tentacles which are much longer than the bell-
diameter. These tentacles have long, conical basal bulbs. In addition to the long tentacles
there are usually 24 to 32 small, immature, or rudimentary tentacle-bulbs which alternate
with the long tentacles. Ocelli ( ?) There are 4 radial-canals, each of which is about 3 mm.
wide in the middle and only 2 mm. wide at either end. Both the radial-canals and the circular

AVTHHMKWS.TC TTTJIMS.

129

tube give rise to numerous simple or branched, blindly-ending side branches ot o.} to i mm.
in length. Some ot these side branches are simple sacs, but others give rise to secondary
branches. The stomach is wide and cruciform, but the side walls are remarkably short, so
that the mouth is a gaping, cruciform slit, reminding one of" the mouth-opening of Stauro-
pliwa. The 4 interradial sides of the stomach form 4 curtains within which the sperm or
ova are developed. The tree, lower edges of these curtains are crenulated and thrown into
complex folds. The stomach and gonads are cherry-red and the canals and tentacles are
light-red in color. This species is found on the coast of Greenland.

Turris digitalis Forbes.

Ttirris <//t;fM/is, FORHF.S, 1848, British N.ikei!-eyed Medusa-, p. 21, plate 3, figs. lil-lt. A<;ASSIZ, L., 1X62, Cunt. Nat. Hi t
U. S., vol. 4, p. }4<>. H\KI k M , 1*74, Sv\t . .Icr Mr ilusen, p. fl , taf. 4, figs. 2, 3. \>* VINSI N, lSn, Vui . Mrddrl . N .M
For. Kjdhenh.ivn (5), Bd. 4, p. 144. -Km KIMUKC., iSSo, Vergleu hen, I phvsiolog. Suidien /u Turnv, etc., Aht. 5, p. 124
(reactions to curare and strychnine).

Bell cylindrical, with a well-developed, conical, apical projection. It is 30 to 40 mm. in
height and 15 to 20 mm. in diameter. There are 50 to 100 short tentacles, which appear as if
arranged in two rows owing to the tact that the bases ot the larger ones extend a slight dis-
tance up the sides ot the bell above the margin.

KM.. 68. Turrii digitalis, after Haeckel, 1879.

Fn;. 6<). Corri* i\ilf,/>liilnilr>ia t after Haeckel, in S\st. lrr MeJuseti.

There are 8 strands ot longitudinal muscle fibers within the wall of the subumbrella.
These strands lie close to, and upon both sides ot, the 4 radial-canals. The radial-canals are
broad and flat. In its upper halt each radial-canal is a simple, flat tube with a straight edge.
In their lower halves, however, the canals become broader and give rise to many small side
branches which end blindly. The manubnum is very large and fills the greater part ot the
hell-cavity. The mouth extends to the level of" the velar opening. There are 4 prominent
crenulated lips. The gonads occupy 4 interradially situated, longitudinal regions upon the
wall of" the stomach. Each gonad consists ot" a double row of papilliform ridges, the outer
surfaces of which are thrown into complex folds and corrugations.

The stomach is purple or reddish-brown and the lips are rose-colored. The gonads
are darker in hue than the stomach. The tentacle-bulbs are orange or yellow and their
outer ends are white. The bell is transparent. The 8 muscle-strands in the subumbrella
are yellow.

130 MEDUS.TC OF THE WORLD.

The species is found in the North Atlantic. It has been taken off the Shetland Islands,
between Greenland and Iceland, between Iceland and the Hebrides, in the North Sea, and off
the west coast of Greenland. It is probably an Arctic form, of circumpolar distribution.

Genus CONIS Brandt, 1834.

(.'own, BRANDT, 1834, Recueil des Actes-seances publ., Acad. Imp. Sci. St. Petersbourg, p. 19 (of the "separate"); 1838, Mem.
Acad. Imp. Sci. St. Petershourg, ser. 6, vul. 4, par. 2, p. 355. HAF.CKKL, 1879, Syst. der Medusen, p. 55.

The type species is Conis niitrnta Brandt of the Bonin Islands, North Pacific.

GENERIC CHARACTERS.

Similar to Pandea, but the tentacle-bulbs give rise to special ocellar clubs, which project
outward around the margin. The gonads consist of simple, longitudinal swellings on the
sides of the stomach and have smooth outer surfaces. The hydroid is unknown.

Conis mitrata Brandt.

Conit mitrata, BRANDT, 1834, Recueil Actes publique, Acad. Sci. St. Petersbourg, p. 19 (separate); 1838, Mem. Acad. Imp.
Sci. St. Petersbourg, ser. 6, vol. 4, par. 2, p. 355, pi. 2. HAECKEL, 1879, Syst. der Meduseo, p. 55. BEDOT, 1905,
Revue Suisse de Zool., tome 13, p. 133 (literature 1834-50).

Bell 50 mm. high, 35 mm. wide, with a short, conical apex. 64 short tentacles, each with
a club-like ocellar bulb projecting from the outer side of the basal bulb at the margin. The
tentacles are shorter than the bell-height. The stomach is barrel-shaped, with 4 not very
complexly folded lips, having their margins studded with nematocyst-warts. Lips are at a
level about half-way between the velar opening, and the inner apex of the bell-cavity. Stomach
light-red. Tentacle-bulbs blue. Ocelli black. Bonin Islands, North Pacific.

Conis cyclophthalma Haeckel.
Cmit r\rlo[>hihalma, HAECKEL, 1879, Syst. der Medusen, p. 55, taf. 4, fig. i.

Bell pyriform, with thin sides and without an apical projection; 15 mm. high, 12 mm.
wide. There are 52 tentacles with very elongate, conical basal bulbs, one-third as long as the
tentacles themselves. The outer ends of the tentacles are thin and flexible while the basal
bulbs are stiff. The tentacles are longer than the bell-height. At the base of each tentacle
on the abaxial side of the margin there is a short club-like projection which contains a black,
ectodermal ocellus. There are 4 straight-edged radial-canals and a simple ring-canal. The
stomach is very large and almost fills the bell-cavity. There are 4 very large, complexly
folded lips with their margins studded with nematocyst-warts. The lips are at about the
level of the velar opening. (See fig. 69.)

Haeckel figures and describes 4 longitudinal, swollen gonads on the "perradial" sides
of the stomach. Their outer surfaces are smooth and they are filled with ova. He represents
them as being completely separated in the interradii. The gonads are probably adradial or
interradial ( ?). Color ( ?)

Haeckel studied a preserved specimen in the Museum of Copenhagen. It was obtained
near Gibraltar in lat. 30 iff N., long. 2 23' W.

Genus CALYCOPSIS Fewkes, 1882.
Calycopsis, FEvvk'ts, 1882, Bull. Mus. Comp. Zoo], at Harvard College, vol. 9, p. 304.

The type species is Calycopsis typti Fewkes, from the Gulf Stream off the coast of the
United States.

GENERIC CHARACTERS.

Tiarinae with 16 separate, simple, unbranched radial-canals; 4 radial, 4 interradial, and
8 adradial. Gonads are composed of a row of leaf-like, transverse foldings along each of the
8 adradial sides of the stomach. Lips ( ?) Tentacles simple. Ring-canal simple, without
blindly-ending diverticula. Development unknown.

ANTHOMEDUSjE CALYCOP818.

131

Calycopsis typa Fewkes.

(,Wv<'//m< t\f><i, l''>.wKts, lSS2, Bull. Mus. Comp. Zool. at Harvard College, vol. 9, p. 304, platr i, fig. 54.

Bell higher than a hemisphere, with evenly rounded top and vertical sides. External
surface smooth, bell-walls thin, thicker above than near the margin. Size ( ?) There are 16
radial-canals and 16 marginal tentacles, one at the base of each canal. These tentacles are

about as long as the bell-height and are flexible with some-
what club-shaped outer ends and very small basal bulbs.
No lithocysts or marginal bodies other than the id ten-
tacles. The stomach is very wide and fills almost the whole
ot the inner half of the bell-cavity. The gonads are devel-
oped upon the sides of the stomach and appear as a series
of transverse, leaf-like ndges on both sides of each of the 4
principal radial lines of the stomach. They are thus
adradial in position and their manner of folding resembles
the condition seen in the gonads of Turns. The sides of
the stomach are bound to the subumbrella by 4 radially
situated partitions. There are 16 straight, narrow radial-
canals: 4 of these are perradial, 4 are mterradial, and 8
adradial. In alcoholic specimens the ovaries were dark-
brown. Tentacles and bell-walls white with a tinge of
bluish color.

4 specimens were found by the Allxitross, in 1880 81,
off the Atlantic coast of the United States, in deep water along the Gulf Stream.
Form of lips ( ?) Size ( ?) Ocelli ( ?)

Fi(i. 70. Cai\rof>si* t\f>ti, after Fewkes, in
Bull. Museum Comp. Zoo), at
Harvard College.

Subfamily MARGELIN^ Haeckel, 1879.

Oceanidx with oral tentacles and with 4 simple, unhranched radial-canals. Gonads
on the mterradial, or adradial sides of the manubnum; the mature genital products are
tound in the ectoderm.

i . Tribe Cvttftdi:

Marginal tentacles arise singly from bell-margin and are not grouped into clusters. Oral tentacles are unbranched, simple, or

degenerate. They are found in all forms excepting in the degenerate medusa; of Stvlactis.
Cyttris, ESCHSCHOLTZ, 1829. Cyt<ris+ Cubogaster HAECKEL. 4 marginal tentacles. Oral tentacles present. Hydroid

unknown.
Podocorvne, SARS, 1846 = Dysmorf>hosa Cytcrandra HAECKEL, 1879. 8 or more marginal tentacles. Oral tentacles

present. When present the peduncle is solid. Hydroid: Podocorynt.

Turritopsis, McCRADY, 1856. 8 or more marginal tentacles. Entodermal walls of the 4 radial-canals, above stomach,
are composed of highly vacuolated cells forming a peduncle. Mouth studded with a row of nematocyst-bearing
knobs. Hydroid: Dendroclara.
Oceania, sens., KOLLIKER, 1853; GECJENBAUR, 1856. Medusa similar to Turritopsh, but with a simple, solid, gelatinous

(not vacuolated) peduncle. Hydroid: Clava-like.

Stvlactis, ALLMAN, 1864, degenerate medusae with 4 to 8 rudimentary marginal tentacles and no oral tentacles. Medusa:
mature upon liberation from hydroid. Hydroid: Stylactts; closely related to Podocoryne.

2. Tribe Thamnostomeidi:

With branching oral tentacles. With simple marginal tentacles, not grouped in clusters.

Thamnost\ltis, HAECKEL, 1879. 2 diametrically opposed, marginal tentacles. Development unknown.
Thamnitis, HAECKEL, 1879. 4 radially situated, marginal tentacles. Development unknown.

L\mnorea, PERON and LESUEUR, i8og= Ltmnorea+ Thamnostoma, HAECKEL, 1879. 8 or more marginal tentacles.
Development unknown.

3. Tribe BougatnviUidi:

Marginal tentacles grouped into clusters. Each cluster consisting of 2 or more tentacles. The 4 oral tentacles may be simple

or branched.
Bougainvillia, LESSON, 1836= Margflis + Lizusa Hippocrene, HAECKKL, 1879. With branching, oral tentacles. With 4

radially situated clusters of marginal tentacles, all of one sort. Hydroid: Bougainvillia.
Ntniopsis, L. AGASSIZ, 1849. With branched oral tentacles. With 4 radially placed clusters of marginal tentacles,

consisting of a median pair of club-shaped tentacles and a number of lateral filiform tentacles in each cluster.

Hydroid: Bougainvillia,
Ratfikfti, BRANDT, 1857 = Mnr^fll turner Lizzia+ Lizztlla+ Rathkeu, HAM-KM., 1^79- With X dusters of marginal ten-

tacles. Mature hydroid unknown.
Cfiiarella t MAAS, 1X97. With 16 (8 double) clusters of marginal tentacles. Circular can.il gi\rs off 4 intrrradi.il, blindly

ending, centripetal branches.

132

MKDUS.E OF THE WUHLI).

Genus CYTjEIS Eschscholtz, 1829.

\_uiiiu. /juui . tii iidi v di u \^<_mcgc, vui. ji, u< iwi . uttuvv i> t, i ywjj ixcfi . i \_c^iun i cd

London, Supplementary Report 27, p. 135. CHUN, 1895, Bibliotheca Zoologica, Heft 19, p. 11 (law of budding).
(1 \tiris. + Cubogaster f HAECKEL, 1879, Syst. der Meduscn, pp. 73, 75.

A r on Cytais, MAVER, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 39.
C.\t,ris, VANHOFKEN, 1891, Zool. Anzeiger, Bd. 14, p. 444.
\on C \tj-is, SARS, 183^, Beskriv og Jagttag., p. 28.

Ctibogasttr, HAECKEL, 1864, Jena. Zeit. fur Naturw., Bd. i, p. 341 (vuung of Cy/.r;.0-
\igritina, STEENSTRUF, 1837, Acta et Catal. Musei Hafmensis, Hauniic.

GENERIC CHARACTERS.

MargeliiiEE with 4 simple, radially situated, marginal tentacles and with 4 or more simple,
unbranched oral tentacles. The hydroid is unknown.

The type species is Cyta-is tetrastyla Eschscholtz, 1829, of the Mediterranean and eastern
parts of the tropical Atlantic. Eschscholtz states that it has 8 simple oral tentacles and 4
marginal tentacles, but he figures at least 10 oral tentacles. More recently species of Cyttcis
have been found with 8, 16, or even 32 oral tentacles.

Tubular Synopsis of the Medusae of Cytirts.

C. tetrastyla

C.atlantica =

C. pusilla

C. vulgaris

C. herdmani

Eschscholtz,

C. macrogaster

Gegenbaur, 1856.

Agassiz and May-

Browne, 1905.

1819.*

Haeckel, 1879

er, iSgg.-f-

= C. tetrastyla

Eydoux ct Soule-

yet, 1841

= C. nigritina

Haeckel, 1879^

Shape and size of

Cubical to domelike

Pyrtform. 5 high,

Oval. 4 high, 3

Prismatic, flat above

Oval. 3.5 high, 3.5

bell in mm.

2 to 3 high, 2 to 3

6 wi e.

wide.

5 high, 3.5 wide.

wide.

wide.

Character of the 4

Similar each to each.

As in C. tetrastyla,

2M- long. With

IT long, with large

As in C. atlantica.

marginal tenta-

2r+ long. Basal

but with large

small basal bulbs.

basal bulbs.

cles. Length in

bulbs small.

pynform basal

terms of bell-

swellings on ex-

raJius (r).

umbrella above

each tentacle.

Shape and size of

Flask-shaped,

Very large, flask-

Stomach oval.

Stomach oval to

Wide flask-shaped,

manubrium.

mounted on

shaped. Two-

Mounted upon

spindle-shape ',

on short peduncle

peduncle. Mouth

thirdsto i .5 times

conical peduncle

half to two-thirds

Two-thirds as

at level of velar

as long asdeptli of

not quite as long

aslong asdepth of

long as depth of

opening.

bcll-cavitv. No

as stomach.

bell-cavity.

bell-cavity.

peduncle in

Mouth at middle

Young medusa has

mature medusa.

of bell-cavity.

short peduncle, but

this is not seen m

mature animal.

Number of oral

4 to 6

8 to 32

12 to 24

3 2

50 to 60

tentacles.

Character of

4 interradial swell-

Numerous medusa-

4 interradial swell-

8 adradial gonads

4 perradial? or 8

gonads and

ings.

buds on upper

ings.

near base, on sides

adradial ? g<>n.i;K

medusa-buds.

half of manubrium.

of stomach.

No medusa-buds.

Color.

Stomach and tenta-

Stomach yellowish.

Stomach and gonads

Stomach and tenta-

?

cle-bulbs brown.

Gonads and tenta-

vellowish-brown.

cle-bulbs red and

cle-bulbs violet-

Tentacle-bulbs

green. Abaxial

brown, red, or

black.

entoderm red,

nearly black.

superficial green.

Where found.

Mediterranean, and

Mediterranean to

Mediterranean.

Fiji Islands to Malav

Coast of Ceylon.

tropical Atlantic.

Atlantic to

Archipelago, trop-

Canary Islands.

Brazilian coast.

ical Pacific.

*Probably young and immature. Possibly young of C. atlantica?

f For details see text.

AXTHO.MKDI-S.K cvr.Kis. i:j;;

The gonads ot C\t<cis are developed upon the mterradial sides of the stomach. In
('\tiris vulgaris, according to Maas, 1905, they occur as 4 pairs of slightly swollen, corrugated
regions, a pair being found on each interradial side of the stomach. The mature genital
products are found in the ectoderm ot the stomach-wall. Many of the species develop medusa-
buds upon the interradial sides ot their stomach. Haeckel's statement that the gonads are
radial in position appears to be erroneous.

Cyttris is closely related to PoJocofyne and is distinguished only by its smaller number
(4) of marginal tentacles, whereas PoJoforync has 8 or more marginal tentacles. MOICOMI
there are commonly more than 4 oral tentacles in ('.\ttcis, whereas the majority of the species
ot Podocoryne have only 4 oral tentacles. C\t<ns is possibly the original genus from which
I'tJvi-rji vii,- and more complex forms may have been derived.

Cttbrjgtisti-r Haeckel, 1864, appears to be only the young ot (,'v/<r/>, excepting " (*.uln\:i>t, i
gemmascens" which I take to be an abnormal young Rathkc/i octoputittntu with only 2 well-
developed and 6 retarded marginal tentacles. Broch, 1905 (Bergens Museums Aarbog. No.
11, p. 4), states that the marginal tentacle-bulbs may range from 4 to 8, and that well-
developed tentacles may be absent.

Cytaeis tetrastyla Eschscholtz.

( 'v/,M \ trrr<i*t\l<i, Ls( ns( HOI. rz, 1829, Syst. der Acalrphrn, p. 104, taf. 8, fig. 2. HAM K M , i S^j. S\ i . ,lcr MrduM-n, p. ?} .

BKDOT, 1905, Revue Smsse dr Zool., tome 13, p. 133 (literature, 1829-50).
Bougainrillca mcditerranea, BUSCH, 1851, Bcobacll. wirbell. Seethiere, p. 21, taf. 2, fig. 10.

This medusa is recorded trom the Mediterranean and the eastern parts ot the tropical
Atlantic. For details see tabular synopsis ot the medusae ot Cyttris.

Cytaeis atlantica.

iVigriiinti (itltintift2 > STKKNSTRUP, 1837, Catal. Musei Hafniensis.

C\(.rn ici',i^\l,i, KVDOUX et SOULEYET, 1841, Voyage de la Botnir, tonic 2, p. ^41, pl.ite 2, figs. 4 to i^.

C.M.ris marrogauer, HAKCKEL, 1879, Syst. der Medusen, p. 74, taf. 6, fig. I. CHIN, 1X1)5, Bihliotheca Zo-ilogna, Mi-ft iij, p. 4^

(budding of medusa from the manubrium).
(;\[,ris nigritina, HAECKEL, 1879, Syst. der Medusen, p. 74, taf. 6, fign. 2-5. MAAS, 1904, Result. Cam p. Sa. Pririo- il<- Mi'iiani,

l.i i . 28, p. 8, plate I, figs. 3, 4.

Bell bluntly conical or pear-shaped, about 6 mm. high and 5 mm. wide. There are 4
stout radially placed marginal tentacles longer than the bell-diameter. The bases ot these
tentacles are very thick and extend up the sides ot the bell as large, bulbous, pear-shaped
swellings. These swellings bear a large crescent-shaped mass of entodermal pigment on
their lower, outer sides. These bulbous swellings are each about one-eighth to one-fourth as
high as the bell. The velum is well developed. There are 4 simple, straight radial-canals
and a simple ring-canal. The manubrium is large and lacks a peduncle. It is 0.5 to i.^
times as long as the depth of the bell-cavity. Normally it is probably about as long as the
depth of the bell-cavity. There are numerous medusa-buds upon the upper halt ot the stomach-
wall and these are seen in various stages ot development. 1 he mouth is surrounded In .1
circlet of 8 to 32 short, slender, oral tentacles, each of which terminates in a nematocyst-knob.
The stomach is reddish-brown and the tentacles brown, with tentacle-bulbs nearly black.
This medusa is found in the Mediterranean and Atlantic trom the shores ot Africa to flu-
Brazilian coast.

Haeckel describes 4 gonads at the points ot |uncture ot the 4 radial-canals with the
stomach, but this is probably erroneous, tor it is improbable that the gonads are m the principal
radii.

Haeckel distinguishes C. nigritimi with manubrium half as long as the depth ot the
bell-cavity, and with 16 to }2 oral tentacles; and ('.. niiu-rngantfr with manubrium 1.5 tinns
as long as the depth of the bell-cavity and with 8 to 16 oral tentacles. _ In 1904, Maas dcsciilu s
a very similar medusa, under the name ot (',. nigi'itnm, in which the manubrium is about
two-thirds as long as the depth of the bell-cavity and has i i oral tentacles. These medusa-
are so closely similar in all other respects that I am of the opinion that they are men-l\
growth and contraction phases of one and the same species. Following the usual rule of pre-
cedence, it appears that the specific name should be "atlantica," this being the designation
proposed by Steenstrup, 1837.

134

MEDUS.E OF THE WOULD.

Cytaeis pusilla Gegenbaur.

(.'v/.n'j fuiilla, Gr.iiiNBAi-R, 1856, Zeit. fur wissen. Zool., Bd. 8, p. 228, taf. 8, fig. 8. KEKERSTEIN UNO EIILERS, 1861, Zoolug.

Beitrag., p. 84, taf. 13, fign. 8, 9. HAECKEL, 1879, Syst. der Medusen, p. 75.
(?) Cubogasler diuonema (young medusa), HAECKEL, Ibid., p. 76.

This medusa is recorded from the Mediterranean. For details see tabular synopsis of
the medusae of Cyttcis.

Haeckel describes "Cubogaster dissonema" a small medusa 2 mm. high, with 2 well-
developed and 2 immature or rudimentary marginal tentacles, and 8 oral tentacles. This
he found at Croisic, Bretagne, Atlantic coast of France. It may be the young of C. pusilla ( ?)

7'-

Flc. 71. Cvttfis atlantica, after Maas, in Result. Camp. Sci. Prince de Monaco.

FIG. 72. Cytefis " macrogaster" ((7. atlantica), after Haeckel, 1879.

FIG. 73. C.ytas'n pusilla, after Gegenbaur, in Zeit. fur wissen. Zool., Bd. 8.

Cytseis vulgaris Agassiz and Mayer.

Cyttsis vulgaris, AGASSIZ and MAYER, 1899, Bull. Mus. Comp. Zool. at Harvard Coll., vol. 32, p. 161, plate 2, figs. 3-5. MAAS,
1905, Craspedoten Medusen der Siboga Exped., Monog. 10, p. 8, taf. i, fign. 4-7; 1906, Revue Suisse de Zool., tome 4,
p. 85, fig. 3, plate 2.

Mature specimens of this medusa are described by Maas from the Malay Archipelago.
Agassiz and Mayer described only half-grown medusae from the Fiji Islands.

Mature medusa. Bell 5 mm. high, prismatic, 4-sided, and with a flat top. There are
4 thick, radially placed, marginal tentacles, each somewhat shorter than the bell-height. These
tentacles end in blunt tips and have large hollow basal bulbs. Maas describes a thick lens-
shaped swelling on the abaxial side of each tentacle-bulb, but his specimens are somewhat
contracted through preservation in formalin. The basal bulbs of the living medusas studied
by Agassiz and Mayer were large and swollen, but lacked the lenticular, ectodermal swellings.
On the other hand Agassiz and Mayer found only half-grown medusae, and it is possible that
the peculiar form of the basal bulbs described by Maas may be characteristic of the full-grown,

ANTHOMKDI s.K (VI.KIS, I'l i| M ( >IM NK. 135

but not of the half-grown, medusa. The velum is well developed. The 4 radial-canals and
ring-canal are slender and simple. The manubrium is spindle-shaped and shorter than the
depth of the bell-cavity. In the halt-grown medusa there is a short conical peduncle, but this
was not seen by Maas in his large specimens.

The mouth is at the extremity of a bluntly pointed, conical neck, and is a simple round
opening. 32 or more short, slender, filiform, unbranched oral tentacles arise from a zone
above the mouth. Each oral tentacle is solid and ends in a knob-like cluster of ncmatocysts.

1 he gonads are found on the upper interradial sides of the stomach and appear as a pair
of longitudinal swellings in each of the 4 interradii. They are not developed on the 4 radial
lines of the stomach. The eggs protrude over the surfaces.

In the formalin specimens studied by Maas, the gonads were yellow, the axial entoderm
of the stomach dark-red, as were also the entodermal parts of the tentacle-bulbs. There is
some green pigment in the ectoderm of the tentacle-bulbs. In the immature specimens from
Fiji, studied from life by Dr. Agassiz and his assistant, the entoderm of the outer part of the
manubrium and of the tentacle-tips was green, while there was red pigment in the axial part of
the stomach and in the tentacle-bulbs and adjacent parts of the 4 radial-canals. In the young
specimens from Fiji the oral tentacles ranged from 4 to id, while in those found by Maas in
the Malay Archipelago, they ranged from 8 to 32.

Found in abundance on the surface among the Fiji Islands in November to January.
Maas found numerous specimens in the Siboga collection from the Malay Archipelago,
Sulu, Damar, Daiam. Sabuida, Gisser, Mampa Straits, and Saleyer; and from Amboma, 1906.

Cytaeis herdmani Browne.

(.'w//\ ftciil'tifini, BKDWNK, 1905, IV.ul OvMer I'ishenes, GuH nf Mjnaar, Km . Soi . LMIU|IUI, Suppl. Report 2^, p. i;^, pl.itr I.
fig. i ; plate 4, fig. 12.

Umbrella about 3.5 mm. in width and height. Somewhat bell-shaped with fairly thick
walls. 4 marginal, long tentacles with very large, spear-head-shaped, basal bulbs one-third as
lung as the bell-height. The ectoderm of the tentacles is thickly crowded with nematocysts.
The entoderm of the tentacles and basal bulbs is pigmented. Velum narrow. 4 wide, straight
radial-canals. Stomach about as wide as long placed upon a short peduncle and extending
a little more than half-way down the cavity of the bell. A row of 50 to fto short, equalh
spaced oral tentacles, each ending in a knob of nematocysts. Gonads form 4 (perradial ?)
swellings extending the entire length of the stomach. Color ( ?) Coast of Ceylon; Chilaw
Paar, and Cheval Paar; March to November.

Genus PODOCORYNE Sars, 1846.

( >)l)\smtirfho;a, PIIILIPPI, 1842, Archiv. fiir Naturgesch., Jalirg. 8, Bd. i. p. ',7, t.if. i, tic,. 3.

I'aitarortnr, SARS, 1846, Fauna littor. Nurveg., tome I, p. 4, t.if. I, figs. 7-18. Hl\< ks, iXdS, British Hvilroul /.,, phvtes, p.

29. ALLMAN, 1871, Monograph Tubul. Hydroids, p. 348. Hi MIM., iK'jj. |<>nm. M.nplml., vol. 9, p. 20^. HMtiixir.,

1905, Znolog. Jahrbuchern, Supplement 6, p. 523.

l'u,lm-or\na, KROHN, 1851, Arch, fiir Naturges., Jahrg. 17, Bil. I, p. 263.
Dvimorpliosa, Ai.^si/, A., 1865, North Amrr. A. ,il., p. id}. I'm N. 1X9^, Bihli.itlu-ta X.n.il., Hi-ft n), I.fi;. I, pp. 12, ;;.

MAAS, 1905, Craspcdotcn M.'iluscn ilor fiibo^a KxpcJitmn, Mnnni;. 10, p. 8. VMIM> > N, |S.||. /...ol. An/i-ri;iT, Bil. 14,

p. 444. BEUOT, 1905, Rcvur Suisse dc Zo..]., tmn<- i ;, p. 7-, ; I'-niotorynt, Ibid., p. 102 (literature to 1850). MAAS, 1906,

Revue Smssr .It- /.mil., form- 14, p. 86, plate }, fit^. 8.

Dysmorphoso+ Cyteeandra, HAKCKKI-, 1879, Syst. iler Medusen, p[i. 76, 79.
Tuiritopsis, AI^ASSIZ, A., 1865, Nnrth Amer. Acal., p. 167.

The name l)\<snitirf>litis<i was first used In Philippi, 1^42, in his description ol I). <on-
<-/i/t olt/, a hydroid polyp which may have been either a Podocoryne or a Hydr actinia in the
modern sense. In 1846, Sars described a closely related, medusa-bearing hydroid under the
name Podocorvne cornea. The name Dysmorphosa was first applied to the free medusa by A.
Agassiz, 1865. The great majority of authors have included both the hydroid and the tree
medusa under the generic name I'oJm <>i \n< . In the absence of proof as to whether Philippi's
hydroid develops medusae or not, we are unable to decide with certainty which name should
stand, but there is no probability that Philippi's hydroid will ever be determined and we niav
avoid confusion by considering it obsolete.

l.'JG MIClirS.K I IF THK WORLD.

GENERIC CHARACTERS.

Margelinae with 4 or more simple, unbranched oral tentacles and with 8 or more simple
marginal tentacles. When present the peduncle of the manubrium is simple, solid, and gelat-
inous. The hydroid stock is Podocoryne.

Haeckel, 1879, restricts the genus "Dysmorphosa" to include only those medusae which
have 8 marginal tentacles. Medusze bearing more than 8 tentacles he places in a new genus
"(1 \iirunJi n." Inasmuch as specimens of Podocoryne fulgurans and P. cnrnea are often found
which have more than 8 tentacles, we consider Haeckel's definitions to be too restrictive. Van-
hoffen, 1891, defines Dysmorphosa, as we do Podocoryne, to include medusa, with 8 or more
marginal tentacles.

PoJ'jiftr \iii- may be distinguished from Lymtiorca by its simple unbranched oral tentacles.
In Turritopsis there are no oral tentacles, but only 4 simple lips studded with nematocyst-
knobs, and the presence of simple oral tentacles in Podocoryne at once distinguishes it from
Turritopsis. It is distinguished from Cyta-is by its having eight or more marginal tentacles,
whereas C ytieis is restricted to medusae bearing 4 marginal tentacles. In Cvtccis the oral ten-
tacles are commonly more than 4, whereas there are only 3 known species of Dysniorp/ios/i
bearing more than 4 oral tentacles. Podocoryne is possibly derived from the more simply
organized C \ta-is.

Podocoryne carnea Sars.

Plate 14, figs. 2 to 6; plate 15, fig. 14.
SYNONYMS OK THK EUROPEAN FORM.

( ) Dysmorphosa conchicola, PHILIPPI, 1X42, Archiv. fiir Naturgesch., Jahrg. 8, Bd. i, p. 37, taf. i, fig. 3.

Podocoryne carnra, SAKS, 1846, Fauna Littor. Norveg., p. 4, taf. i, figs. 7-18. KROHN, 1851, Archiv. fiir Naturgesch., Jahrg.
17, Bd. I, p. 266. HINCKS, 1868, British Hydroid Zooph., p. 29, plate 5, 6 figs. ALLMAN, 1871, Monograph Tubul. Hy-
droids. p. 349, plate 16, figs. 1-9. DEVARENNE, 1882, Archiv. de Zool. Ejpcr., tome lo, pp. 645, 674, 683, plate 33, figs.
6-15; plates 34, 36-38; 1882, Compt. rend. Paris, tome 94, p. 892. WF.ISMANN, 1883, Seiualzellen bei den Hydromedu-
sen, pp. 63-72, taf. 19, fign. 1-13. BROWNK, 1896, Proc. Zool. Soc. London, p. 463. DEVARENNE, 1881, Annals and
Mag. Nat. Hist., ser. 5, vol. 9, p. 134. HAMANN, 1882, Jena. Zeitschrift fur Naturwissen., Bd. 15, p. 517, taf. 20, fign. I,
3, 4(histology of hydroid). JICKELI, 1883, Morphol. Jahrb., Bd. 8. p. 621, taf. 27 (histology of the hydroid). GRAEFFE,
1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 347. ISHIKAWA, 1888, Zeit. fiir wissen. Zool., Bd. 47, p. 621, 6 fign. (origin nf
egg cells). BEDOT, 1905, Revue Suisse de Zoo]., tome 13, p. 103 (literature 1842-1850). RITCHIE, 1907, Trans. Roy.
Soc. Edinburgh, vol. 45, p. 1^23 (from Cape Colony, South Africa).

Dysmorphosa carnea, HAECKEL, 1879, Syst. der Medusen, p. 77.

(:\[,ti! exi^nti, HAECKEL, Ibid., p. 634.

Podocoryne h'fckcln, HAMANN, 1882, Jena. Zeit. fur Naturwissen., Bd. 15, p. 519 (young stocks of I', carnea ?).

(?) Podocoryne conchicola (Philippi) in part, HARGITT, 1904, Mitth. Zool. Station Neapel, Bd. 16, p. c8l, fig. 26, taf. 22.

SYNONYMS OF THE AMFRICAN REPRESENTATIVE.

Turritopsis nutricula, AGASSIZ, A., 1862, Proc. Boston Soc. Nat. Hist., vol. 9, p. 97, figs. 22, 23; 1865, North Amer. Acal.,p. 167*
figs. 269, 270. FEWKES, 1881, Bull. Mus. Comp. Zool., vol. 8, p. 153, plate 4, figs. 4, 7-10.

t'.nli \<tion formosuii, FEWKES, 1882, Bull. Mus. Comp. Zool., vo