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MEDUSAE OF THE WORLD

VOLUMis, 11!

LHe SCYPHOMEDUSAE

BY

ALFRED GOLDSBOROUGH MAYER

SS ae =
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WASHINGTON, D. C.
ste

PUBLISHED BY THE CARNEGIE INSTITUTION OF WASHINGTON

IgIo

CARNEGIE INSTITUTION OF WASHINGTON

Pusuication No. tog, Vou. III

Copies of this Book
were first issued

AUG 25 1910

ew yor

CONTENTS.

VOLUME III.

Orderd Cary bdetd cet saree (a sisitites wialorsysi1s ois oes. sicse Geusieieieceis Hae ne oe ere gle cise ong eile ve ae 504-518
Procharybdis tetraptera.... . 506 Carybdea alata var. pyramis.511 Chiropsalmus quadrumanus.515
Carybdea marsupialis....... 507 Van PLAN dS ere este) ns-)= Sir buntendij kieen er ete seer 515

PAStOM apts eee seiste cre 508 Vale MM OSEMtls)o elise) 512 QUuadnigatus: creer ss: 516
Mal VINA GAN diary ta (oleic = 509 TUCLAY ata tee sree ataet tel 512 Zygoneinal seers 517
FAM els Ao onkte poneeee 510 Tamoyahaplonema........ 513 Chirodropus gorilla........518
Al ACA vai ayers tage asian seats eons 510 Tripedalia cystophora..... 514

Orderi Sta wrommed sae spat seeks orc ceyete ee ckcte yoy ote oc bets ahs el Serer © sich ate crs, VaR eueias ein: Siokats lelondokepare atalais 519-540
Eleutherocarpidz.......... 519 Lucernaria walteri......... 529 Haliclystus stejnegeri. ... ...535
Gleistocarpidzes::2- a-mias- 519 kukenthalieeeraeiae erie 529 ANtALCHCUSs «fe eeieteerr- 536
Mllesserareracas yee nacht 522 Inaeckelign smnsnscscrs an. 529 KerguelemsiSueeet eine 530
Messeranthal./0. -) eta. 2255 522 msn ibulunome sires 529 Halimocyathus platypus. . . . 537
Messeramta s:forerys.ssteasys.yenerue 522 campanulata........... 530 lapenabienrm tet scrr recite 537
Depastrum cyathiforme..... 524 AUSEhal iss verte eer 530 Craterlophus tethys........ 538
Stenoscyphus inabai........ 525 Kishinouyea nagatensis..... 531 IMA CLOCYStiS ue spey teas 538

hexaradiatus...........52§ ialiclystus auricula........ 592) Capuasturdziien.. se - 539
Lucernaria quadricornis. . . . 527 Octoradiatus). 3/2. se rao 534 Lipkearuspoliana.........540
Pykamidalliches eter 528 Sal PINK jaro caesarean ysis 535

@rdenCoronateca cere rere <eersts. store eele ena erates Aa ats Sieletess sie te tts AMOMEMIOS Gres clapalersie i or svad 541-569
Periphyllide.......... 541-548 Ephyropside.......... 550-560 Linucheaquila............ 560
Pericolpa quadrigata....... 542 Palephyraantiqua......... Stee Collaspideseemeem-- bi 561-567

Campanave.t. savas ce 542 pelagical eco ters ier p52) eAtollal baind items ley ties: 563
iGurlint@enocasoaos aos 542 INdiGasaewee ey ee eee 553 forma valdivie...... 565
Periphylla hyacinthina...... 544 Nausithoe punctata........ 554 PUP ANICE Agape Se vate = <a) a 565
forma dodecabostry- Chau Sle teysteisrciayers) taicst en: 550 GUID: cokep hon tebe eayays taney 566
diClanacaamect ther 546 challengenteee ere 550 Wey valletrrchs 0, ar5 scape 2 eiees 2 566
forma regina........ 546 albatrossliewyjscle ere 557 forma verrillii....... 567
Periphyllopsis braueri. . . . . . 547 TU Yale atameye nic cetis ers 5S 7ieeatorel idee mee os eee 567-569
Nauphantopsis diomedez. . . 548 PlCtan betyerct acne sie. 557 Atorella subglobosa........ 568
Paraphyllinide........ 548-550 Linuche unguiculata.......558 Vanhofientme caer ee cer 568
Paraphyllina intermedia. . . . 549

Ordergsemacostom eas wv aeyer-poite mishap Aa oe.Nans Rite aO els s Sys o/s! -naisins eaiay vise halo fers 569-630
Pee la pa daeteylatsrssepete ir 569-591 Dactylometralactea........583 Ulmaride............ 604-630
Pelapramoctilitcass..... 454 - 570 quingWecinrhasersi cries 585 Floresca parthenia......... 605

Var.neplecta.......- 574 Ehin(htlod edad ceeucene. 588 Discomedusa lobata........ 607
Gyanell ale te syepessexsryorscare 574 fernUipimastena i eres te 588 phulippimaleese yee: 607
PAMOPylae es ae oso 7 lonsicinral- ee seer 589 Undosa undulata..........609

Van. placental. «4 ..-)-< 575) Wuragea depressa...4-.- + 4- 589 Dipulmaris antarctica...... 610
Haveolag a. raster 576 Sanderia malayensis........ 590 Sthenoniaalbida........... 611
Peviamere mec ysae eccne SO) | Gy anreidaeee cet f-y-4/-) 5 591-604 Phacellophora camtschatica . 613
phosphora sr... 576 Desmonema gaudichaudii. . . 593 SIGUA. weieiers crores sislere 613
(CLASSab rere on ye ate aes 576 chierchianales. eee a 593 AND HAAAAo. Soeapeenboc 615

Chrysaora hysoscella....... 579 Cyaneacapillata........... 596 Omataleya sions s ae ae. chers 616
var. blossevillei...... 581 THIBNUN Clos onc oodods 600 Poraliarufescens..........617

Vea tges tl orl daletes ee) -h-1 = 581 var. versicolor....... GoorAurelliataunttas. ee eects - 623
blossevillei var. plocamia . 581 Val. NOZakier ee = 4-1. 601 forma marginalis. . . .627
helvolamrrcg rte rts ta ee 581 annaskalalyaaeeieiee se 601 ub tatves tavern ost scieies 627

var. calliparea....... 582 Drymonema dalmatina..... 603 SOlid aici 142 i ceisle las 627

var. chinensis........ 582 (Hot)o CON DE ADIEN OY OD anG 604 Jalbiataiis:.jatcvosisielsysieisie.s 628
melanaStenens tyes eiste. sie - G82) sPateran \.sjerc.cs cvs asAsyeescts 6 604 MmaldivensiSeee erie 629

var. gilberti......... 582, Donacostoma. .-. 25. ... 604 Aurosafurcata............ 630

CONTENTS,

IV
Order Rhizostom2. « «sos acic's.cs sas ven vip bv b anid > Weide eles Deep ii apie te ata aie tale ele teistai ee eae 631-714
Rhizostomata pinnata. .635-650 Catostylus cruciatus....... 667 Versuraanadyomene....... 686
Toreuma dieuphila......... 636 paloiipesis.)-11-7cn eee 667 HAAS» dinner eee 687
Cassiopea andromeda... .. . .637 PETG REE prom AS Sees 668 Labonemasmithii.........689
var. zanzibarica..... .639 stuhimanni. >) 45.5 4s 669 Rhizostomata lorifera. . .691-697
var. malayensis. .... .639 OFS: 5 goers 669 ‘Thysanostoma thysanura. . .692
var. maldivensis. . . . .639 stiphropterus........... 670 Loriferalorifera........... 694
var. acycloblia...... .640 viridescens; <% 5/<:,.. ++: 670 Wak. Pacifica... <2 1 695
xamachana..........-. 641 ornatells. a. .cesaeee 670 Hapellata. 0% 5 ccn ener 695
fronddsac. 2 cee ere ae 647 CKIPteMS.© 5h ae ee ae 671 Leptobrachia leptopus..... . 696
QMiiata ee tte eee eae 648 PUNDURUS so eee eee 671 Rhizostomata scapulata.6g7—711
var. digitata......... 648 Lychnorhizalucerna.......673 Rhizostoma pulmo......... 699
Gepressaesa% «iectere elects 649 bartschii-etncseis tortor 674 varvluted.. 5-096. 0- 703
var. picta...........649 Crambione mastigophora. . .676 var. octopus......... 793
mertensity -../.e eee 649 COOK. << san one secs nes 677 War commas cet 703
Ue Here tance: Sear 2a nine 650 Mastigias papua........... 678 var. Capensis........ 703
Rhizostomata dichotoma 650-63 VATS SIGereas sent 679 Rhopilemaesculenta....... 704
Cephea octostyla........... 652 var. siboge.........680 hispidum...) --eee 706
var. coerulescens..... 653 ocellatayce tne sss Voces 680 verrillits: 220. sonqan ee 707
CODNGd ee ane eee ee 654 pantherinals +. ieee 681 Eupilemascapulare........ 709
Var COMMeL ale tes eras 655 Pracilen sc ayaa wee 681 Stomolophus meleagris. . ... 710
var. dumokuroa...... 656 oor eo tindauncacipdloc 681 var. fritillaria........ 711
var. coerulea. .......657 Pseudorhiza aurosa........682 Rhizostomata simplicia. 712-714
var. setouchiana. . .. .657 haeckelit:.:. --70 Reet 683 Archirhiza aurosa.......... 712
typhlodendrium. .......658 Phyllorhiza punctata....... 684 Haplorhiza simplex........ 713
Cotylorhiza tuberculata..... 659 Versura palmata........... 685 punctata.” sj. aecse a 713
Polyrhiza vesiculosa...... . .663 vesiGata:<isa3,1-e ener 686 Cannorhizaconnexa........ 714
Rhizostomata triptera. .663-691 DUT Eicloseder crane noe 686 Stomatonema reticulatum.. .714
Catostylus mosaicus........ 666
Fossil. Medusa... 00°) 2's. 0 e's Gag oven tinier crcl cited on ne Sere atone nee ee Tne eee ee ee 715-718
Medusina radiata.......... 715 Acraspedites antiquus...... 710) aotira cambriaac eee 717
PHNCE DSc ee ne tetera: 715 Semeostomites zitteli....... 716 Dactyloidites asteroides..... 717
deperditats.- eee ee 715 Eulithota fasciculata.......716 Rhizostomites admirandus. .718
Paraphyllites distinctus. ....715 Myogramma speciosum.... . 716 Brooksella alternata........ 718
Cannostomites multicirrata..716 Medusinacostata.......... 717 Confiisals sates: oeeeeney ta 718
Atollites minor............. 716 peryonidessasc: -cetanie 717 thendnasd: +24 Senne ee 718
MAECEL I 52 eta she tavicge etn sds 716
7.41) 2) 1) See RE ROMO CUMS: GUmIn or oe GD OUMOCOEDD OOO THADG sabtpes dor ae Sa NC ac ~se Ene ae 719-726
Preoccupied Generic Names.719 Urashimea globosa......... 722 Microhydra ryderi......... 726
Corynitis = Linvillea....... 719 Luruitopsispacifica. ..---6- 722) Joimnocnidan: ete ieee 726
Slabberia=Dipurena....... 719 Rathkea octopunctata...... 722, “MSGinacitrear letaces se eee 726
Turris and Tiara=Clavula. .719 Val. pratass.anedeaas 722 Scyphomeduse ....... 726-728
Wapdiceal.0 cares ke eee 719 Nemopsis dofleini.......... 723 Carybdea rastonii.......... 726
Hydromeduse.........720-726 Willsia pacifica............ 723 alataz a: ie. See eee 726
Pennanaltiacella. octet 720 Polyorchis karafutoensis....723 Haliclystus octoradiatus. . . .726
Corymorpha nutans........ 720 Spirocodon saltatrix........ 724 Thaumatoscyphus dis-
Sarsia rosaria..............720 Obeliacongdont........... 724 tinctUS:Aces. 455.22 727
\apONicdtay sent ites ee 720 Staurophora mertensii. .... . 724 Parumbrosa polylobata.... . 728
Bleuthenacaeccer erence 721 Cubaia gemmifera......... 725 Ephyropsites jurassicus. ... . 728
Cladonema radiatum....... 721 Craspedacusta sowerbii..... 725

MEDUSAE OF THE WORLD.

THE SCYPHOMEDUSAE.

INTRODUCTION.

The present volume is a continuation of the work the two former parts of which dealt
with the Hydromedusz. The acknowledgments which I was privileged to render in the
introduction to the first volume need not here be repeated, pleasurable to me as sucha repeti-
tion would be. Suffice it to say that the work was commenced in 1892 at the suggestion of
Dr. Alexander Agassiz whose generous aid enabled me to pursue these studies for many years
under the most advantageous conditions; and apart from the sense of personal gratitude
I cherish toward Dr. Agassiz I hope that enough of scientific worth may be found within
these volumes to cause it to appear that I have made appreciative use of the opportunities he
so magnanimously accorded to me. Throughout the years I have hoped that Dr. Agassiz
might live to see this work and that he might be pleased by this fruit of his inspiration,
but on March 27, 1910, the great student of the oceans died upon the sea, only a few weeks
before the publication of these volumes.

Since the first two volumes passed under the press I have been most kindly aided by the
United States Bureau of Fisheries and by the National Museum of the United States, at
Washington. The authorities of the former permitted me to study the important collection
of Scyphomedusz recently made by the Albatross in the Philippine Islands, and those of the
latter institution were so courteous and helpful as to set aside for my use a table in the
Smithsonian building in order that I might study this collection to the best advantage. It
is due especially to De Hugh M. Smith, Deputy Commissioner of the Bureau of Bishewes
and to Dr. Richard Rathbun, Assistant Secretary of the Smithsonian Institution that I owe
these highly appreciated favors.

Plates 61 and 64A and several text-figures are taken from drawings made from nature
by my friend, the late Prof. William K. Breaks: These drawings were most kindly presented
to me for publication in this work by Prof. E. A. Andrews, of the Johns Hopkins Univ ersity 5
soon after the death of Professor Brooks. They will serve as memorials of the rare skill in
observation and in draftsmanship of the great naturalist who made them.

SCYPHOMEDUS2.

Phanerocarpe, Escuscuoutz, 1829, Syst. der Acal.

Steganopthalma, Forxes, 1848, British Naked-eyed Meduse.

Acraspede, GEGENBAUR, 1856, Zeit. fiir wissen. Zool., Bd. 8.

Discophore, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4.

Acraspede, HarcKEt, 1880, Systems der Medusen, Halfte 2.

Scyphomedusa@, LANKEsTER, 1881, Encyclopedia Britannica, Ed. 9, Hydrozoa, p. 547-

Medusiform Cnidaria with tentacle-like, entodermal filaments upon the subumbrella floor

of the stomach-cavity. With entodermal gonads, and without a velum such as is characteristic
of the Hydromeduse. Development by strobilization from scyphostoma larve.

There is but little evidence to support the supposition that the Scy phomeduse have been
derived from the Hydromeduse. The medusa shape of the sexual generation in both has in
all probability been acquired independently. Indeed, various animals have assumed the
external appearance and peculiar mode of locomotion by pulsation which is characteristic of
the medusa. Such for example are the protozoan Cras pedotella and the holothurian Pela-
gothuria. Moreover there is reason to believe that the medusa-bell of the Narcomedusz has

499

500 MEDUS OF THE WORLD.

been acquired independently of and is not homologous with the bell of the Leptolina medusz.
(See Goette, 1907, Zeit. fiir wissen. Zool., Bd. 87, p. 289.)

The peculiar velum of the Carybdeidze among the Scyphomedusz is a structure of the
subumbrella, mot of both subumbrella and exumbrella asin Hydromeduse. It may be regarded
as a parallelism, and not genetically related to the velum of Hydromeduse. The exumbrella
nerve-ring, found commonly in Hydromedusz, does not exist in the Scyphomedusz, but is
replaced by a subumbrella plexus of fibers extending between the marginal sense-organs and
also radially i inward from these ganglionic centers. There is also a diffuse, nervous, epithelial,
ectodermal network over the subumbrella. The subumbrella alone is sensitive to stimuli, the
exumbrella being non-sensory, but covered with a nematocyst- bearing epithelium.

In the Scy phomedusee the mature, sexual products are found in the entoderm, whereas
they are usually in the ectoderm in Hydromeduse. The velum which is universally present
in Hydromedusz is absent in Scyphomedusw. The 4 interradial, gastric septa which are
always seen in the scyphostoma larva of Scyphomedusz are not found in the polyp stage of
Hydromeduse.

When we come to consider the relationships of the several orders constituting the Scypho-
medusa themselves we meet with difficulties which render our classification only tentative.
The history of the attempts to classify the Scyphomedusz have recently been reviewed by Maas,
1907 (Ergeb. und Fortschritte der Zool., Bd. 1, p. 189), and by Bigelow, 1909 (Mem. Museum
Comp. Zool. at Harvard College, vol. 37, p. 13). Marked advances over Haeckel’s artificial
classification were made by Claus, 1878, 1883, who showed that the Carybdeide are a very
aberrant group not closely related to other Scyphomedusz, and not descended from the Stauro-
medusz as Haeckel believed. As Claus showed, the interradial, gastric septa in Carybdeidz
may be newly arisen fusions of the entodermal walls of the exumbrella and subumbrella and
not derived from the interradial taeniolz of the scyphostoma. Unfortunately, however, no one
has seen the late scyphostoma stage in Carybdeida, and hence our knowledge of their rela-
tionships must remain doubtful. Claus, 1883, and Vanhéffen, 1892, called attention to the
supposed importance of the presence or absence of the interradial, gastric septa in the classi-
fication of sexually mature Scyphomedusz but we must remember that these are absent in
Semzostomz and found in Coronate, yet these two orders are closely related. Moreover,
interradial gastric septa are found both in Stauromeduse and Carybdeidz yet all modern
students agree that these groups are widely divergent. The sessile Stauromedusz certainly
approach very closely in structure to the type of the scyphostoma, but this may be the result
of degeneracy due to their sedentary habit.

Vanhoffen, 1892, in a masterly paper called attention to the importance of the annular
furrow and marginal, exumbrella sculpturing of the bell in certain of Haeckel’s Discome-
dusz, and he grouped these under the designation Coronate. Claus, 1883, had previously
shown that in Nausithoé, which is a typical member of the Coronatz, the interradial septa of
the central stomach are derived from the 4 teniole of the scyphostoma. These septa are, how-
ever, absent in the medusa-stage in Semzostomez and Rhizostome, although they are found
in the scyphostoma.

All evidence both anatomical and embryological shows that the Rhizostomz are descended
directly from the more simply organized Semzostomez. It is probable that the Coronate and
Semzeostomez are not descended one from the other, but are derived from some common
ancestral stock which has disappeared. The Stauromeduse may represent a highly specialized
derivative from this common ancestral form, their specialization being due to their sessile,
attached habit of life. The Carybdeidz are so aberrant and our knowledge of their develop-
ment is so imperfect that we have not yet been able to determine their relationships to other
Scyphomedusz.

The habits and distribution of the several orders of Scyphomeduse are characteristic.
The Stauromeduse are found only in the Arctic and Antarctic regions and in cold seas,
and are unknown from the tropics; thus furnishing us with an interesting problem in dis-
tribution. The planula is not pelagic but creeping and is devoid of cilia, and the medusz
are sessile or creeping, and do not pulsate rhythmically. They affix themselves to seaweeds
and rocks and rarely move from their places of attachment. They are probably degenerate
forms although their relationship to other Scyphomedusz is uncertain. They are confined to
the shores, but may be carried far from their normal habitats upon drifting seaweed.

SCYPHOMEDUS 4.

The Orders of the Scyphomeduse.

501

Carybdeide.

Stauromedusz.

Coronate.

Semaostomea:.

Rhizostome.

Form of bell.

Margin of bell.

Tentacles.

Sense-organs.

Interradial septa
in stomach.

Mouth.

Cubical with inter-
radial angles, and
perradial sides.

No lappets but with
an annular mem-
brane formed from
the subumbrella
and constituting a
velum.

4 interradial, single
or in clusters. Ten-
tacles hollow, lash-
like, and mounted
upon spatula-shaped
projections of the
subumbrella.

4 perradial clubs set
in niches on sides
of bell. Ocelli and
lithocyst present.

Present.

Cruciform, simple.

Pyramidal, usually
attached to objects
by aboral apex.

Usually with 8 ad-
radial lobes which
bear the tentacles.

Usually 8 adradial
clusters of hollow,
knobbed tentacles.

When present 8 (4
perradial, 4 inter-
radial) anchors
without ocelli or
lithocysts.

Present.

Cruciform, simple.

Discoidal with an-

nular furrow, and
marginal zone of
pedalia in exum-
brella.

Cleft into lappets.

Arising from clefts
between lappets.

Arising from clefts
between lappets.
An entodermal
mass of concretions
always present.

Present.

Cruciform, simple.

Discoidal without
annular furrow or
pedalia.

Cleft into lappets.

Arising from sub-
umbrella or from
clefts between
lappets.

As in Coronate.

Absent.

Unitary. Cruci-
form, usually with
veil-like lips.

As in Semzostomea.

Cleft into lappets.

Absent. In one
genus (Lobonema)
the marginal lap-
pets have been
metamorphosed
into tentacle-like
organs.

As in Coronate.

Absent.

Numerous mouths
surrounded by mo-
tile tentacles and
borne on 8 ad-
radial, fleshy, arm-
like processes.

The Carybdeidz are highly specialized forms which are confined to tropical and warm
seas. They live upon the bottom in shallow water along coasts, and are not commonly found
upon the surface until nearly mature. Their developmental stages are practically unknown
and their exact relationship to the other orders of Scyphomedusz is problematical. In com-
mon with the Stauromedusz and Coronatz they have 4 perradial stomach-pouches. They
bear a remarkable, but wholly superficial, resemblance to the Hydromedusz in the shape of
the bell and in the presence of an annular diaphragm which constricts the aperture of the
bell-cavity. This velar diaphragm is, however, only an extended part of the subumbrella
in Carybdeidze whereas both exumbrella and subumbrella take part in its formation in
Hydromeduse.

The Coronatz are deep-sea and pelagic forms and are therefore of world-wide distri-
bution in common with other creatures of similar habits.

The Semzostomee are mainly coastal forms which develop through strobilization from
scyphostoma larve. The genus Pelagia is peculiar, however, in that the larva is not attached,
but remains free-swimming and develops directly into the medusa; and thus it is that medusz
of this genus are widely distributed, independently of the proximity of land and are found in
all warm seas. Certain families, such as the Cyaneidz, are practically confined to the
Arctic and Antarctic, and to cold seas; whereas the Pelagidz are found only in warm oceans.
On the other hand the genus Aurellia is of world-wide distribution.

The Rhizostome form a well-defined order which are clearly descended from the Semz-
ostomez. They are confined to tropical and warm seas and are most abundant in the Malay
Archipelago. They develop through strobilization from scyphostoma larve, and are therefore
confined to coastal waters of a depth suitable for the maturation of their attached larve.

In the Semzostomez and Rhizostome the egg develops into a pyriform, ciliated planula
which is set free from the mouth-folds of its parent and swims freely through the water. Soon
it attaches itself by means of its forward end. An ectodermal invagination is said by Goette to

502 MEDUS OF THE WORLD.

occur at the non-attached, upper pole of the larva. 4 perradial stomach-pouches develop, 2
being partially ectodermal and derived in some measure at least from the invaginated ectoderm
of the cesophagus, the other 2 pouches being entodermal and derived from the entoderm of the
stomach of the larva. These perradial pouches are separated by 4 interradial septa which
finally become perforated near the margin forming a peripheral ring-sinus. 4 funnel-like

cavities lined by longitudinal muscle- fibers sink downward into these 4 initerradial septa from
the hypostome of the larva, and tentacles dev elop around the margin of the hypostome. Finally
the larva splits by transverse clefts into a series of discoidal, free-swimming, ephyra larva,
and the original corona of tentacles is cast off and dies while a new set of tentacles and a 4-
cornered mouth may develop at the upper end of the remaining part of the larva. In some
species, and under certain conditions, this strobilization may be monodiscus and give rise to
but a single ephyra, while in others from 10 to 30 ephyrz may arise from a single scyphostoma.
Details of this peculiar process of delamination and regeneration, and of the larval stages in
general, will be found in Korschelt and Heider’s text-book of the embryology of invertebrates,
part 1, 1895, pages 102 to 122, and throughout the text of the present work. See especially
Cyanea arctica, Aurellia aurita, Pelagia, Chrysaora, Cotylorhiza tuberculata, and Cassiopea
xamachana.

With reference to the relationships between Hydromeduse, Scyphomedusz, and actinians,
Goette’s announcement that the cesophagus of the scyphostoma of Scyphomedusz is formed
of invaginated ectoderm seemed to suggest a close relationship between the scyphopolyp and
the actinozoa. Hadzi, 1907, however, re-affirms the work of Claus and supports the view that
the cesophagus of the scyphostoma is lined on the inside with entoderm and is not inyaginated
but evaginated from the primary stomach. All of the stomach-pouches are therefore ento-
dermal according to Claus and Hadzi, whereas, 2 of the primary stomach-pouches are at least
partially ectodermal and 2 wholly entodermal according to Goette.

We know, however, from Conklin’s study of the development of Linuche, and Hyde’s
research upon Auurallias that gastrulation in one and the same species may occur either through
invagination or by ingression, and judging from the mass of evidence which has been accumu-
lated we can not doubt but that the mouth of the scyphostoma of one and the same species
may be formed either by invagination, or by a simple breaking through of both entoderm and
ectoderm in which the entoderm takes the more active part, or even by an evagination of
entoderm. The three processes may thus be closely related and the manner of operation
dependent upon which layer takes the initiative and assumes the more active part. It is there-
fore misleading to attach any deep morphological significance or to draw any sweeping con-
clusions in respect to the phylogeny of the Scyphomedusz from this process. Indeed, long
ago our faith in the rigid application of the germ-layer theory has become so shaken that the
discussion between the Goette and the Claus schools has lost most of its significance.

According to the Claus-Hadzi view the Scyphomeduse are more closely related to the
Hydromeduse than they are to the actinians. Certainly the histological characters of the
germ layers of the scyphostoma resemble the hydropolyps rather than the actinozoa, although
both are so closely similar that no great weight can be attached to this fact. Hérouard, 1909
(Comptes Rendus, Paris, tome 148, page 1225), supports the opposite view from histological
evidence, for he finds that the cellular investment of the pharynx of Scyphomedusz and scypho-
stoma laryz is similar in character to that of the gullet of Anthozoa, thus indicating a genetic
relationship between the Anthozoa and Scyphomedusz.

According to Goette the 4 primary stomach-pouches are actively evaginated from the cen-
tral stomach-cavity and are of mixed ectodermal and entodermal origin. According to Claus-
Hadzi, however, they are wholly entodermal, and are only passively separated one from another
by the infolding of the 4 entodermal foldings i in the stomach-wall which constitute the teniola.

It is extraordinary that this contention should have persisted so long without a final settle-
ment of views.

The table on the opposite page will serve to present the question more clearly.

The scyphostoma larva of Nausithoé bears a remarkable superficial resemblance to
hydroids, being elongate, branched, and incased by a horny perisarc. It infests sponges.

In the majority of the Scy phomedusze the sexes are separate, but in Chrysaora and in
certain Rhizostoma the meduse are hermaphroditic.

SCYPHOMEDUSA,

508

| Hydropolyp
(hydroids).

Inner lining of cesopha-
gus.
Teniolx.

Muscle-strands.

Stomach-cavity.

Evaginated and ento-
dermal.

None.
|

| derm and entoderm
cells.

| Unitary, but often with

| irregular, longitudinal
| tidges of entodermal

cells. (See Hamann,
1882, Jena. Zeitsch.

Formed from basal elon- |
gations of both ecto- |

| Naturw.,Bd.15, p.509-))

Scyphopolyp (Scypho-
stoma), according to
Claus-Hadzi.

Evaginated) « and ento-
dermal.

4 wholly of entodermal
origin, but with ecto-

dermal muscle-strands.

OF ectodermal origin.

With 4 primary pouches

of purely entodermal
origin simply separated
one from another by 4

entodermal infoldings. |

| Sey Fiepein p (Sey pho-
stoma), according to
Goette.

Invaginated and ecto-
dermal.

4 of mixed ectodermal |

and entodermal origin, |

but with ectodermal
muscle-strands.

Of ectodermal origin.

2 of the primary pouches

| Anthopdly Pp
(Actinozoa).

Invaginated and ecto-

dermal.

Septa composed mainly
of entoderm, with ento-|
dermal anteils S.

Mainly entodermal but
some ectodermal in
origin.

With lateral pouches.

are partially ectoder- |

mal and 2 of purely
entodermal origin.
They grow actively

outward from sides of |

central stomach.

Ripe sexual products. | In ectoderm. In entoderm. | In entoderm. In entoderm.

| ee
| Structureless, free from | Containing cells.
cells. |

Intermediate lamella. | Containing cells. Containing cells.

Mode of reproduction. | Meduse develop by
budding.

Medusa develop by

strobilization.

No meduse developed.
strobilization.

|

Romanes and Eimer found that if the marginal sense-clubs of Scyphomedusz be removed
a more or less complete paralysis results, although the disk remains capable of responding
temporarily to any stimulus. Bethe, 1903, found that in Rhizostoma pulmo and Cotylorhiza
tuberculata the pulsation-stimulus is nervous in nature, and there are many analogies between
the thythmic movement of these medusz and that of the vertebrate heart. For example the

“all or none” principle applies to medusz, as does also the phenomenon of the insensibility
of the medusa to stimulation while in systole, and the extra systole and compensating period
of rest. Mayer, working upon Casstopia xamachana, finds that the sea-water is a balanced
fluid, the stimulating tendency of its sodium being exactly counterbalanced by the inhibiting
effects of its magnesium, calcium, and potassium. The pulsation stimulus arises in the mar-
ginal sense-clubs and is due to the constant formation of sodium oxalate in the entodermal
cells at the distal end of each club. This precipitates the calcium chlorides and sulphates
of the sea-water forming the insoluble calcic oxalate crystals of the sense-club, and setting
free sodium chloride and sulphate the cations of which are powerful nervous stimulants, and
produce the periodic response of the nervous elements which causes the rhythmic pulsation.

In the hydromedusz on the other hand the function of the control of pulsation is not
localized in the sense-organs for any part of the bell-rim will serve to maintain the bell in
pulsation.

It is interesting to note that whatever the effect of the several cations, sodium, magnesium,
calcium, and potassium, may be upon the neuromuscular system, their effects upon the moye-
ment of cilia is the exact opposite. Whatever stimulates muscles inhibits cilia, and vice versa.

The gastric tentacles, which arise from the subumbrella wall of the stomach on the sides
of the interradial septa are very characteristic and constant structures in Scyphomedusz.
They are entodermal with solid cores of mesogloea, and are in no sense comparable with the
marginal tentacles, but their function is unknown.

The marginal tentacles themselves are to be regarded as structures of the subumbrella.
They are usually but not invariably hollow and consist of an axial core of entodermal cells
encased outwardly by ectoderm which is richly besprinkled with nematocysts. The marginal
sense-clubs are highly specialized tentacles which have been transformed into sensory centers.
Ocelli may or may not be present and when found they may be of ectodermal or of entodermal
structure; but concretions of crystalline nature are iny ariably found in the entodermal cells
at the distal end of the club in all forms exhibiting rhythmical pulsation. The gonads are
follicular foldings of the entodermal subumbrella floor of the stomach.

Meduse develop by |

504 MEDUSZ OF THE WORLD.

Order CARYBDEIDZ Gegenbaur, 1856.

Carybdeide, Gecennaur, 1856, Zeit. fiir wissen. Zool., Bd. 8, p. 214.

Charybdea, Craus, 1886, Arbeit. Zool. Inst. Univ. Wien, Bd. 7, p. 110.—VANuOrFEN, 1892, Ergeb. der Plankton Expedition,
Bd. 2, K. d., p. 23.

Marsu pialida, renee L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 174-

Cubomeduse, Hacker, 1880, Syst. der Medusen, p. 423.—Goerre, 1887, Abhandl. zur Entwickelungsgesch. der Thiere, p. 66,
Leipzig. —Scuweixorr, 1889, Morphol. Jahrb., Bd. 15, p. 25.—Conant, 1898, Mem. Johns Hopkins Univ. Biol. Lab.,
vol. 4, No. 1, p. 3-—Haacke, 1887, Jenaische Zeitschrift, Bd. 20, p. 5g0.—Maas, 1907, Ergeb. und Fortschritte der Zool.,
Bd. 1, p. 197-

CHARACTERS OF THE ORDER.

Scyphomeduse with 4 perradial, knob-shaped, marginal sense-organs which are situated
within niches upon the sides of the bell. Each sense-organ contains an entodermal concretion,
and one or more ectodermal ocelli. There are 4 interradial tentacles or groups of tentacles.
4 wide perradial sacs extend outward from the central stomach into the interstitial space of the
bell. These sacs are incompletely separated one from another by 4 narrow interradial septa.
There are 8 gonads each one of which is leaf-shaped and attached along one side to an inter-
radial septum, from which it extends outward into the gastrovascular space of the bell. The
bell-margin is not cleft into lappets. The subumbrella forms an annular diaphragm called
the velarium which partially closes the opening of the bell-cavity.

The perradial sense-organs in the Carybdeide are knob-shaped, and are set within
niches upon the sides of the bell. The stalk of each knob is hollow and connected with the
gastrovascular system of the bell. The knobbed end of each club contains a large entodermal
concretion and one or more ectodermal eyes (plate 56, fig. 6”). These eyes are placed on
the inner or centripetal side of the club and look inward into the bell-cavity. The larger
eyes are provided with a doubly convex lens, back of which is a cup-shaped space occupied
by the lens-capsule and the retina. The retina is made up of a single layer of cells forming a
hollow cup, into which the lens with its capsule fits. The central space of the retinal cup
is clear and glassy. This transparent region is enveloped by a deeply pigmented part and
back of this lies a muscular zone. The retina is made up of three kinds of cells, some long
and prism-shaped, others short and pyramidal, and still others long and pyramidal. The
deep-lying ends of these cells are more or less pigmented, while their outer parts are trans-
parent, and being packed closely together, they form the vitreous center of the retinal cup.
(See Conant, 1898; Berger, 1900; etc.)

The tentacles of the Carybdeidz are interradial and arise at a level slightly above the
apparent bell-margin, although they are actually structures of the subumbrella. Their proxi-
mal parts are developed into tough, gelatinous, wing-shaped or spatula-shaped structures
(pedalia), which probably serve as keels to steer the animal in its course through the water.
The long, distal ends of the tentacles are cylindrical and highly contractile, and their outer
surface is covered with rings of nematocysts. The tentacles are hollow and their lumen is
connected with the gastrovascular space of the bell. This is accomplished by the interradial
septa being incomplete near the bases of the tentacles and thus the lumen of each tentacle
is placed in direct connection with the gastrovascular spaces of the two adjacent perradial
pouches. The flexible parts of the tentacles are armed with nematocysts, the stinging power
of which is so great that the name ‘Sea Wasp” is commonly given to these medusz.

The velarium is an annular membrane which extends inward from the bell-margin,
forming a diaphragm which partially closes the cavity of the subumbrella; superficially it
bears a close resemblance to the velum of the Hydromedusz, but is separated from the exum-
brella by means of a continuous sheet of entodermal tissue which penetrates the gelatinous
substance of the bell all around the bell-margin, and connects the entoderm of the intermediate
lamella with the ectoderm of the outer surface of the bell. It thus forms a ring of entoderm
around the bell-margin which completely separates the ectoderm of the subumbrella side of
the velarium from that of the exumbrella. At points other than those occupied by sense-
organs or tentacles this vascular, entodermal lamella forms a simple, flat sheet, but when it
comes to the tentacle-bases, or sense-organs, it makes a loop upward over them. Thus the
tentacles and sense-clubs are structures of the subumbrella only, for they lie below the ento-
dermal lamella which isolates them completely from the exumbrella. What the philogenetic
significance of this lamella may be we do not know, and until this is discovered we can not be
certain that the velarium of the Carybdeidz is not strictly homologous with the velum of the

a

9
f { ae Cie Of ee Ine c oo ae 7
Hb) ar wa. -- « oi Be ings. 4 tel -
ia OS A es jill ST @ ; a
in: P18 > Sisas 2S . ei it 5 Te 7
Ae : ¢ jf
eof a
q? TT). 7 ‘ :
a a
ae
-_
-
"
e

5

PraTE 56.

. 1. Side view of Haliclystus salpinx, from eelgrass near Ram Island, off

Manchester, Massachusetts, September 7, 1905.
. Oral view of medusa shown in figure 1.
. Section of stalk of medusa shown in figure 1.

2
3

. 4. Side view of marginal organs of medusa shown in figure 1.
5

. Young medusa of Carybdea xaymacana, showing prominent nema-
tocyst-warts over the exumbrella. Tortugas, Florida, May 24,

1899.

. 6. Quarter-grown medusa of Carybdea xaymacana. Nassau Harbor,

Bahamas, March, 1893.

. 6’. Side view of one of the sense-clubs of medusa shown in figure 6.
. 6”. Section of sense-club of medusa shown in figure 6. /, ectodermal

lens of cup-like eye; oc, ocellus; of, otolith. Entoderm brown,
ectoderm gray.

. 7. Carybdea xaymacana showing gonads; twice natural size. Nassau

Harbor, Bahamas, April 24, 1907.

. 7’. Side view of the sense-club of the medusa shown in figure 7.
. 7". View of the sense-club of the medusa shown in figure 7, showing side

that faces inward in such manner as to observe objects within the
bell-cavity.

Drawn from life, by the author.

PLATE 56

MAYER

2%
Abeen tts

f ee \

) pep {/
oe) ie td |]
aaa ond Ir) |

(Gs)

Sy” ||

lyr”
o
SS

Pa 2\3

« \
A a
' } °
: ~ & x -q bs
@ (eA7h \\ Ws ee jae.
et § St =) N Vet m d A
ae - © C F
el, ae \\ | Re, Read
% SS Vv ‘ = \ raat
= iY | ’ es | a setetveggee
as iS / a WS - on
¢ mC2e) ee) a oan = 2 <p Ry, Re.
a —— a = ae Ps > @) ay
E-a e . 4 —————————-_*- eT passe 2 aI '
‘. = ‘ —— : . Sawn +) 3
a =a GS © Fj
a “SY :
Sa bs §
SS

» “5
7p»

CARYBDEIDA. 505

Hydromedusz; for it constitutes the only important point of difference in the velar diaphragms
of the two sorts of animals. The velarium is commonly bound to the wall of the subumbrella
by 4 solid, bracket-like supports, the frenula, one of which is found in each perradius. There
is also a more or less complicated system of entodermal canals or pouches which extend into
the substance of the velarium from the gastrovascular space of the bell.

There has been a tendency among modern authors to regard the velarium as being com-
posed of a series of fused lappets, the pouches being remnants of lappet-pouches. We have
no proof, however, that this is the case, and the “aoa | in the youngest Charybdeidz yet
seen is as entire as in the adult.

There are 4 groups of gastric cirri, or phacelli, in the interradial corners of the central
stomach, at the inner ends of the 4 interradial septa, and this feature alone would distinguish
these forms from the Hydromedusz in which no such structures have been observed. The
gonads, also, are entodermal instead of ectodermal when mature as in Hydromedusz. Both
the gonads and gastric cirri are structures of the subumbrella, as is also the entire muscular
system.

There is a well-developed nerve-ring on the bell-cavity side of the subumbrella near the
margin. This ring forms 4 loops upward to the 4 perradial sense-organs (plate 57, fig. 2).
There are 8 ganglia upon the nerve-ring, 4 perradial and 4 interradial. Each of the perradial
ganglia sends off 2 nerve-roots, which pass through the gelatinous substance of the subumbrella
and fuse as they extend down the inner (centripetal) side of the stalk of the sensory-club.
The 4 interradial ganglia are situated at the bases of the tentacles into which they probably
send nerve-fbers.

The Carybdeide are inhabitants of the warmer waters of all the oceans, and none of
them has been found in the Polar seas. Most of them have been taken in the open ocean,
but they are also found swarming in harbors and other places near land. When young they
appear to be bottom forms, but they usually come to the surface when mature.

But little is known concerning the embryonic development of these forms. Conant, 1897,
found that in Trzpedalia the ovum develops into a free-swimming planula, which soon settles
down upon the bottom and becomes a hydra-like polyp with a mouth and 4 tentacles. Haacke,
1887, found a very young individual of Carybdea rastoni in which a short style canal extended
upward from the upper floor of the central stomach to the aboral apex of the bell, and he
believes it possible that this structure may represent the remnant of some connection between
the young medusa and some form of scyphopolyp nurse, but this is wholly problematical.

Haeckel, 1880, believed that the Carybdeidz were descended from the Stauromeduse,
as, according to him, were also his Periphyllida and the Discomeduse. He believes that
morphologically the Carybdeidze are intermediate between the Periphyllida and the Disco-
medusz. All of this, however, is speculation unsupported by a single fact of any significance.
We must first learn more of the early embryonic stages of the medusa of the Carybdeidze
and Stauromedusz before we venture to state how they may be related in philogeny to other
Scyphomeduse. They have perradial stomach-pouches which are partially separated by
interradial septa as in the Stauromeduse; indeed Goette, 1887, would consider the Caryb-
deidz as an off-shoot of or cousins of the Stauromedusez.

Most of our knowledge of the anatomy of the Carybdeidz is due to the labors of Claus,
1878; Haeckel, 1880; Schewiakoff, 1889; Conant, 1898; and Berger, 1900.

Berger, 1900, reports upon a few physiological experiments upon the reactions of Caryb-
deida (see C. xaymacana). A synopsis of the genera of the Carybdeidz follows:

(?) Procharybdis Haeckel, 1880. 4 simple pedalia. Velarium without velar canals or frenule. This is probably only
a young Charybdea.

Carybdea Péron and Lesueur, 1809. 4 simple pedalia. Velarium with velar canals and with 4 perradial bracket-like
supports (frenula). Stomach small, without hollow bracket-like sides arching over between it and the subumbrella.
Stomach small and simple, with 4 horizontal clusters of gastric cirri.

Tamoya F. Miiller, 1859. Similar to Charybdea, but with a large stomach which is bound to the subumbrella by 4
perradial, hollow arches and with 4 vertical, interradial clusters of gastric cirri.

Tripedalia Conant, 1897. Numerous, 8 to 12 or more, simple unbranched pedalia arranged in 4 interradial clusters.
No hernia-like subumbrella sacs.

Chiropsalmus L. Agassiz, 1862. 4 branched, hand-shaped pedalia with tentacles arising from the tips of the fingers.
With 8 simple, finger-shaped, hernia-like subumbrella gastric sacs. Free-margins of the 8 gonads entire and simple.

Chirodropus Haeckel, 1880. Pedalia hand-shaped as in Chiropsalmus. 8 branched, hernia-like sacs projecting into
the subumbrella from the 4 perradial stomach-pouches. Free-margins of the gonads showing grape-like swellings.

506 MEDUS OF THE WORLD.

Genus (?) PROCHARYBDIS Haeckel, 1880.
Procharybdis, HAEcKEL, 1880, Syst. der Medusen, p. 437.

The type species is P. tetraptera Haeckel from the Sunda Islands, Indian Ocean. It
may prove to be only a species of Carybdea which is regenerating its velum or is immature.

GENERIC CHARACTERS.

Carybdeide with 4 simple, interradial tentacles with pedalia. Velarium simple, without
velar canals or frenule.

P. turricula and P. flagellata of Haeckel, loc cit., p. 438, are both too inadequately known
to be profitably retained in scientific literature. P. cuboides Haeckel, is, I believe, a young
stage of the common Carybdea rastonit. The briefly described Procharybdis securigera
Haeckel, Joc cit., p. 640, from the Pacific coast of Central America, may be C. rastoni, but
it is said to have tentacles which terminate each in a knob. This character is seen in young
individuals of Carybdea.

Procharybdis tetraptera Haeckel.

Procharybdis tetraptera, Harcxe1, 1880, Syst. der Medusen, p. 437, taf. 25, fign. 3, 4

Bell dome-like with thick walls, 30 mm. high, 20 mm. wide. 4 very large, flat, expanded,
simitar-shaped pedalia, with blunt outer ends; these pedalia are nearly as long as the bell-
height and nearly one-third as wide as they are long; they are relatively larger than in any
atier known form of Carybdeide. The 4 perradial sense-clubs are set in niches nearly as
far above the bell-margin as the width of the bases of the pedalia. Each sense-club is small
and apparently contains only a single very large eye and an entodermal concretion. Each
of the flexible, hollow tentacles tapers to a pointed end. They are ringed with nematocysts
and are about 1.5 times as long as bell-height.

The yelarium is without velar canals or bracket-like frenule, and its margin is entire.
It is only about twice as wide as the tentacles and appears to be very rudimentary. It occurs
to me that this velarium may be regenerating after having been lost through accident. Should
the velarium have velar canals and frenulz, the medusa would be one of the genus Carybdea
which it resembles in all other respects.

The stomach is small, wide, flat, with 4 large, pointed lips. There are 4 pairs of gastric
cirri in the interradial corners of the stomach, each consisting of an axial shaft which gives
rise to 20 to 30 filaments on one side. The 8 gonads are developed as in other species of
Carybdeidz on both sides of the 4 interradial septa.

Haeckel describes this medusa from a single preserved specimen found in the Sunda
Archipelago, Indian Ocean.

Genus CARYBDEA Péron and Lesueur, 1809.

Carybdea, Péron et Lesueur, 1809, Tableau des Méduses, Ann. Mus. Hist. Nat., tome 14, p. 332-—Mitne-Epwarps, 1833,
Annal. des Sci. Nat., tome 28, p. 248

Charybdea, Craus, 1878, Arbeit. Zool. Inst. Univ. Wien., Bd. 1, p. 221.—Harcket, 1880, Syst. der Medusen, p. 439; 1881,
Deep-sea Medusew Challenger Expedition, Zool., vol. 4, p. 91 —Conant, 1898, Mem. Biol. Lab. Johns Hopkins Univ.,
vol. 4, No. 1, p. 3.—Haacke, 1887, Jenaische Zeit. fiir Naturwissen., Bd. 20, p. §90.—Bicetow, H. B., 1909, Mem. Mus.
Comp. Zool. at Harvard College, vol. 37, p. 17—Maas, 1909, Abhandl. Aadk. Wissen., Miinchen, Suppl. Bd. 1, Ahandl.
8, p. 40.

The type species is C. marsupralis Péron and Lesueur, of the Mediterranean.

GENERIC CHARACTERS.

Carybdeide with 4 simple, interradial pedalia and tentacles. Velarium supported by 4
perradial, bracket-like frenula which bind it to the subumbrella. Velar canals present. Stom-
ach small, without bracket-like mesenteries joining it to the subumbrella. Development
unknown.

Péron on Lesueur spell this generic name Carybdea; Agassiz, 1862, Cont. Nat. Hist.
U. S., vol. 4, p. 173, changed it to Charybdea.

All hy species of this genus are inhabitants of warm seas. None have been taken north
of Cape Cod on the North American coast, or north of the Mediterranean in Europe. Owing
to the slight differences between them, it is exceedingly difficult to separate the species one

CARYBDEIDA—CARYBDERA. 507

from another. The most characteristic features for s
of oe Bees and the number and character (branched or unbranched) of the velar canals.
_ Cary a ae ated C. rastonit, and C. xaymacana are very closely related, if not mere

i = ~ rr Se : ee . y 4 : . .
varieties, of one and the same medusa. They are found in the Mediterranean, tropical Atlantic,
and Pacific.

_ _C. pyramis, which appears to be identical with Haeckel’s C. obeliscus and C. alata is
distinguished by its long, simitar-shaped pedalia. It comes from the tropical Atlantic.

' C. grandis and C, mosert are closely related tropical P
varieties, one of the other, indeed Bigelow, 1909, believes them to be mere growth stages of
a and the ie me C. moseri being the younger. C. grandts is possibly identical with

ursarius cytheree Lesson (=Tamoya bursaria Haeckel) but the descriptions of the last-
named medusa are so vague that it must, I think, be dropped from our lists.

ie murrayana Haeckel is distinguished by its large number of profusely branched velar
canals. In other respects it closely resembles C. marsuptalis, of which form it may indeed
be a mere variety.

Semper’s Philippine Island Carybdea, called C. philippina by Haeckel, may be identi-
cal with C. moseri, but is too vaguely mentioned to be recognizable.

G verrucosa Hargitt, 1903, is a very young form, too immature for identification.

C. aurifera Mayer is distinguished by its rosin-colored bell, being far darker in color
than in any other species.

F ‘ ue :

Haeckel’s genus Procharybdis appears to be composed of immature or imperfectly known
young specimens of Carybdea. It is not improbable also that future studies will show that
Haeckel’s Procharagma is actually Carybdea.

In some of the species of Carybdea the gastric cirri at the interradial corners of the
stomach are in the form of branched tufts. The mature eggs are set free from the gonads and
float in the gastric pouches, where they undergo a part of their development; but the larval
stages remain practically unstudied.

A few physiological observations are reported of C. xaymacana by Berger and are discussed
under the description of this species.

pecific distinction are the shape and size

acific forms and may be mere local

Carybdea marsupialis Péron and Lesueur.
Plate 58, figs. 1 to 5.
Medusa marsupialis, Lint, 1758, Systema Nature, Ed. 10, p. 660; 1788 (Gmelin) tomus 1, pars 6, p. 3154.
Carybdea marsupialis, Peron ev Lesueur, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 333-
Charybdea marsupialis, Craus, 1878, Arbeit. Zool. Inst. Wien, Bd. 1, p. 221, § taf., fign. 1-48 (anatomy, histology)—Harcket,

1880, Syst. der Medusen, p. 442 (references to literature).

Bell prismatic, 4-sided, somewhat constricted near aboral apex, and with flat top. It
is 40 mm. high and 30 mm. wide at level of velarium. The apex of bell, the pedalia, and
the interradial ridges of the exumbrella are besprinkled with numerous, quite regularly spaced,
oval nematocyst-warts of dull ocher-yellow color. The 4 interradial pedalia are each 17 mm.
long and 10.5 mm. wide at their widest parts. The flexible, lash-like, filamentous parts of
the tentacles are very contractile and range in length from 2 to 12 times as long as bell-height.
These tentacles are hollow and regularly ringed with nematocysts.

The 4 perradially placed sense-clubs are each set in a niche almost covered by a gelat-
inous flap. These sense-clubs are 5 mm. above the velar margin of the bell. Each sense-
club has a median pair of large eyes with a doubly convex lens, and also 4 small, lateral ocelli,
which lack lenses. All 6 eyes are directed inward so as to look into the bell-cavity, and are
of ectodermal structure. Besides the 6 eyes there is a large terminal mass of entodermal
cystalline concretions.

The velarium is supported by 4 bracket-shaped buttresses (frenula) which bind it to the
subumbrella in the radii of the sense-clubs. There are usually 3 (occasionally 4) branched,
blindly-ending centripetal vessels in each octant of the velarium. These vessels branch quite
complexly and irregularly, but do not anastomose. The base of the stomach is wide and
flask-shaped, but the throat-tube is very narrow and with 4 simple, lanceolate lips. Altogether
the manubrium is not more than one-third as long as the depth of the bell-cavity. There
are 4 minute clusters of gastric cirri, one in each interradial corner of the stomach. These

508 MEDUS OF THE WORLD.

Synopsis of the Spectes of Carybdea.*

C. marsu- | C. rastonii= | C. xaymaca- | C. murraya- | C. alata. C. pyramis= C. grandis=| C. moseri=
pialis. | a variety of | na=a variety na=a varie- | a variety of | a variety of | a variety or
C. marsu- | of C. marsu-) ty of C.mar- | C. alata. C. alata. young stage
pialis (?) pialis. | supialis (?) | of C.grandis.
Size of bell in| 4-sided, 4-sided, Truncated | Square- |60 high, 50 | Truncated | Truncated | High-dome-
mm. prismatic. | prismatic. | pyramid | sided, flatly | wide. Sides) pyramid. | pyramid. shaped, thin
40 high, 35 high, above; pris- | dome-like | pyramidal, | 30 high, 230 high, | walled. 80
30 wide. | 30 wide. | matic below.| top. 6ohigh, apex 20 wide. 140 wide. | high, 47
| 23 high, 50 wide. rounded. Thick walls.| wide.
15 wide.
Form and size, Spatula- Small,scalpel- Scalpel-shap- | As in C. ras- | Spatula- Very long, | Wide, flat, | Wide, flat
of pedalia in| shaped, flat.) shaped. One~ ed, flat, wider) tonii. One-| shaped. narrow, spatula- as in C.
terms of One-third | third toone-| than in C. | third long, | One-half curved, shaped. grandis.
height of long, one- fourth long, | rastonii, One-| one-sixth long, one- | scythe- One-sixth | One-third
bell. | fourth wide.| one-fifth to | third to one- | wide. fifth wide. | shaped. long, one- | long, one-
| one-sixth | halflong, one- 1 long, ninth wide. | fifth wide.
| | wide. fourth to one- one-eighth
| third wide. wide.
Length of | 2 to 12 L.5+ | 8+ 2+ 2+ I+ 0.75 to 1 I to 1.5
flexible ten-
tacles in
| terms of
height of bell.)
| Number of | Six. 6 6 6 6 6 1 to 3
eyes in each | 2 median; 1 median; | 2 median;
sense-club. | 4 lateral. 2 lateral. | 2 lateral.
Form and | 3, occasional-| 2 branched |2 asin C. | 6 profusely |3 branched | 3 unbranch- | 3 branched, | 3 unbranch-
number of | ly 4, branch-| but not anas-) rastonii. branched, | complexly, | ed, or only | butnotanas-| ed.
velar canals | ed but not | tomosing. but not anas-| but not anas- some of them) tomosing.
in each oc- | anastomo- tomosing. | tomosing. | branched.
tant of velar-| sing. Non-anas-
ium. | tomosing.
Size and Base wide, | Wide, flat, | Wide, flat, | Wide, flat, | Short with 4 Small, with 4 Small, with | As in C
shape of __ | throat-tube | less than half) with large, | with 4 short} simple lips. | large lips. | 4 simple grandis.
stomach. | slender and | as long as | lanceolate lips. lips.
| small. | bell-cavity. | lips.
Where found., Mediter- Tropical West Indies, | Coast of Tropical Tropical Tropical Tropical
ranean. Pacific. tropical SierraLeone, Pacific. Atlantic. Pacific. Pacific.
Atlantic. West Africa.
Depth of
200 fathoms.

* For C. aurifera see text.

cirri are brush-like, and in each cluster about 8 to 10 primary branches arise, and each gives
off 2 to 3 lateral branches, each of which terminates in a brush of 10 to 13 filaments. There
are thus 100 to 150 of these terminal filaments in each interradial cluster of gastric cirri.
The gonads are 8 leaf-like expansions on both sides of the 4 interradial septa. They extend
not quite to the velar margin or to the interradial edges of the stomach.

Bell and pedalia dull milky-ocher, due to the color of the exumbrella nematocyst-warts.
Flexible parts of tentacles dull pink. Ocelli very dark brown, nearly black; basal branches
of gastric cirri dull horny-brown.

The medusa is common in the Mediterranean, but its development remains unknown.
Claus, 1878, gives a detailed account of its anatomy and histology. When young it apparently
remains in deep water probably at or near the bottom, but when mature it swims upward to
the surface.

Carybdea rastonii Haacke.

Charybdea rastonii, Haacke, 1886, Zool. Anzeiger, Bd. 9, p. 554; 1887, Jena, Zeit. fiir Naturwissen., Bd. 20, p. 591, taf. 35, fign.
1-15; 1888, Biol. Centralblatt, Bd. 8, p.357.—von LenpeEnFexp, 1888, Biolog. Centralblatt, Bd. 8, p.218.—Marer, 1906,
Bull. U.S. Fish Commission, vol. 23, Part 3, 1903, p. 1134, plate 1, figs. 1-1¢.—B1Getow, H. B., 1909, Mem. Mus. Comp.
Zool. at Harvard College, vol. 37, p. 17, plates 1 and 10.—Maas, 1909, Abhandl. Akad. Wissen., Miinchen, Suppl. Bd.
1, Abhandl. 8, p. 41.

Char ybdea arborifera (young medusa), Maas, 1897, Mem. Museum Comp. Zool. at Harvard College, vol.23, p. 86, taf. 14, fign. 7-10.

Procharagma prototypus (young meduse), Harcxet, 1880, Syst. der Medusen, p. 436, taf. 25, fign. 1, 2.—Procharybdis cubotdes,
Ibid., p. 439-

Bell nearly cubical, with flatly rounded top and nearly plane vertical sides. 35 mm. high,
25 to 30 mm. wide. Pedalia small, only one-third to one-fourth as long as bell-height, and

PLATE 57.

Fig. 1. Carybdea xaymacana, young medusa. Nematocyst-warts so promi-
nent in young medusa (fig. 5, plate 56) are beginning to disappear.
Nassau Harbor, Bahamas, June, 1903.

Fig. 2. Tamoya haplonema, natural size. Mature medusa dredged from a
depth of 2 fathoms off Brown’s Point, Great Peconic Bay, Long
Island, New York, September 5, 1902.

Fig. 2’. Tamoya haplonema. Quadrant of bell-margin showing velar canals.

Fig. 2”. Tamoya haplonema. Sense-club showing side that is directed inward
toward the interior of the bell-cavity.

Fig. 2’”. Tamoya haplonema. Side view of sense-club.

Fig. 3. Chiropsalmus quadrumanus, 0.66 natural size. Fish Commission
Laboratory, Beaufort, North Carolina. From the bottom off
Beaufort entrance at a depth of 3 to g fathoms, July, 1904.

Drawn from nature, by the author.

MAYER

PLATE 57

Pha
ANNE i>
RAY MSs Wie
OTS OP
Y ee
RSS ,
io Vv
Aittitts
PNW

SES

Qe

CARYBDEIDA2—CARYBDEA. 509

three-fifths as wide as long. Flexible lashes of tentacles 1.5 times as long as bell-height. 4
sense-clubs in niches 5 mm. above velarium. Each club with 2 large median and 4 small
lateral eyes, and an entodermal lithocyst. Velarium wide with 4 perradial, subumbrella
frenulz and 16 short, branched, non-anastomosing velar canals, 4 in each quadrant. Manu-
brium flat, wide, less than half as long as depth of bell-cavity. 4 simple lips. 4 very small,
interradial tufts of branched gastric cirri as in C. marsupialis. ;

8 leaf-shaped gonads along entire sides of the 4 interradial septa. Each gonad is widest
near the stomach and tapers toward both ends. Flexible parts of tentacles and gastric cirri
dull pink. It swims toward a light at night.

Gulf of St. Vincent, South Australia; Honolulu Harbor, Hawaiian Islands; Subig Bay
and Nasugbu, Luzon, and Mausalay, Mindoro, Philippine Islands, in January. Common
on surface. Probably widely distributed over the tropical Pacific.

The medusa begins to develop its gonads when the bell is only 11 mm. high and they are
large in meduse 15 mm. high. ;

The youngest meduse found by Haacke had a pyramidal bell and an axial-canal above
the stomach-cavity as if it might have been attached at one time to a scyphostoma nurse.
Each sense-club had but 2 eyes, the median ones; and the 16 velar canals were simple and
unbranched. C. arborifera Maas, 1897, from Honolulu, is clearly the young of this species.

This small medusa may be distinguished by its cubical bell and small pedalia. It is
closely related to the Mediterranean C. marsupzalis.

The following are the dimensions (in millimeters) of a specimen obtained by the U. S.
Bureau of Fisheries steamer Albatross in Subig Bay, Luzon, Philippine Islands, on the surface,
January 6, 1908. Height of bell, 34 ; width of bell, 20; length of pedalium, 11; width of
pedalium, 6.5; sense-clubs, 6 above the velar margin; flexible shafts of tentacles, contracted,
30 long. In this specimen the bell was unusually narrow.

Carybdea xaymacana Conant.
Plate 56, figs. 5 to 7; plate 57, fig. 1.

Charybdea xaymacana, Conant, 1897, Johns Hopkins Univ. Circl., No. 132, p. 8, fig. 8; 1898, Mem. Biol. Lab. Johns Hopkins
Univ., vol. 4, No. 1, pp. 4, 7-573 figs. 1-16, 31-34, 36-43, 57-67, 69, 70, 72.—Bercer, E. W., 1898, Jour. Comp. Neurol-
ogy, Granville, vol. 8, p. 223, 5 figs. (structure of eyes); 1900, Mem. Biol. Lab. Johns Hopkins Univ., vol. 4, No. 4,
p- 1-84, 3 plates—Mayer, A. G., 1904, Mem. Nat. Sci. Museum Brooklyn Inst. Arts and Sci., vol. 1, plate 7, fig. 60.

Tamoya punctata (young medusa), Fewxes, J. W., 1883, Bull. Mus. Comp. Zool. at Harvard College, vol. 11, p. 84, plate 1, figs. 4-6.

Bell 18 to 23 mm. high, 15 mm. broad. Sides vertical for two-thirds their height, above
which they slope slightly inward. A slight concavity at top of bell. Pedalia flat and scalpel-
shaped and between one-third and half as long as bell-height. The 4 tentacles are at least 8
times as long as the bell-height. The 4 sensory-clubs are situated each within a niche about
one-seventh or one-eighth the distance from bell-margin to apex. Each club contains an ento-
dermal, crystalline concretion and 6 ectodermal eyes; 2 of these eyes are large and median,
4 are small and lateral. These eyes are all on the centripetal side of the club, so as to look
inward into the bell-cavity. The median eyes are each provided with a prominent lens,
whereas the lateral eyes have no lenses. Velarium about one-seventh as broad as bell-diame-
ter. 16 velar canals, 4 in each quadrant; these canals are forked at their ends, at times with
more than 2 branches. Stomach flat and shallow. The throat-tube, which is well developed.
with 4 large oral lobes, hangs down into bell-cavity a distance between one-third and half
the bell-height; it is very sensitive and contractile and can be inverted into the stomach,
The 4 tufts of gastric cirri are epaulet-shaped and of small size. Each tuft arises from a
stalk-like base which projects from the subumbrella floor of the stomach. There are 8 leaf-
like gonads.

Bell translucent, slightly pink or milky with bluish-purple nettling warts near the aboral
apex of the exumbrella and bluish-purple tentacles. -

This species was found by Conant in Kingston Harbor, Jamaica. I have obtained it in
Nassau Harbor, New Providence Island, Bahamas, in spring and summer.

Good figures of the mature medusa are given by Conant, 1897-98.

Berger, 1900, finds that strong light, or darkness, inhibits the pulsation of this medusa.
A sudden change in the intensity of the light acts as a stimulus. Removal of all 4 sense-

510 MEDUS2Z OF THE WORLD.

clubs causes a short temporary “‘paralysis,” but pulsation is soon resumed. Severance of
the marginal nerve-ring in 8 places, so as to isolate the sense-organs from the tentacles, does
not interfere with pulsation. The operation, however, causes the pedalia to bend inwardly
by contraction. When the pedalia are cut off the medusa swims unnaturally, being unable
to steer itself and turning in circles and somersaults. Removal of the perradial and inter-
radial marginal ganglia produces paralysis of the pulsations.

When young the exumbrella of this medusa is regularly besprinkled with brown-colored,
conspicuous clusters of nematocysts. When the bell is 4 mm. high it is pyriform, thin-walled,
and the pedalia are merely short, flattened, swollen basal bulbs of the ringed tentacles. The
velarium has no yelar canals and there are no gastric cirri. The young medusa is abundant
in Nassau Harbor, Bahamas, during the summer.

Carybdea aurifera Mayer.
Char ybdea aurifera (young medusa), MayeR, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 70, plate 25, figs. 81-83.

A young medusa was described from the Tortugas, Florida, in 1900, but in 1909 a much
larger but yet immature specimen was found. In this large specimen the bell is 7 mm. long,
5 mm. wide at the velar margin, thin walled, and tapering to a blunt apex. The exumbrella

‘< is besprinkled by large, wart-like clusters of nettle-cells. The
4 pedalia of the tentacles are 2 mm. long and the shafts of
the tentacles about the same length. These tentacle-shafts are
each ringed by 7 to g rings of nematocysts. The 4 rhopalia
are set in niches about 1 mm. above the velar margin. Each
club bears 1 large, median and 4 small, lateral eyes. The
velarium is wide, has well-developed, circular muscles and is
supported by 4 perradial frenulea. There are 16 main velar

Fic. 328.—Carybdea aurifera, young medusa. From life, by the author, at Tortugas, Florida, July 11, 1909.
A, side view. B, oral view showing velar canals. C, gastric cirri. D, side view, and inner side of sense-club.

canals; the 8 adjacent to the frenula are narrow and simple, but the 8 adjacent to the
tentacles are wide and each gives off 3 or 4 finger-like processes. The manubrium is wide,
4-sided and nearly half as long as the height of the bell, with 4 simple lips. There are about
100 simple, tapering, unbranched gastric cirri. The most characteristic feature of this species
is its uniform rich rosin or golden-brown color. It is a rare form and has been taken only
twice in ten years in surface tows, in July, at Tortugas, Florida.

Carybdea alata Reynaud.
Carybdea alata, Reynaup, 1830, Lesson’s Centurie Zoologique, p. 95, planche 33, fig. 1—VannGrren, 1908, Deutsche Siidpolar

Expedition, 1901-03, Bd. 10, Zool. 2, p. 34, fign. 3, 4.

See synoptic table of species of Carybdea.

Bell 55 mm. high, pyramidal, with a rounded apex and rounded angles. Bell 42 mm.
wide at margin and 17 mm. wide at base of rounded apex. The 4 pedalia are each 27 mm.
long and 12 mm. wide at widest part. The rhopalia are 13 mm. above bell-margin. When
the bell is 55 mm. high there are 6 wide dichotomously forked velar canals in each quadrant
berween successive pairs of tentacles. In medusw having a bell 60 mm. high, the forking of
these canals becomes more complex and quite irregular, no two quadrants being alike. Thus

CARYBDEIDH®—CARYBDEA. 511

it is probable that C. pyramus from the West Indies, C. obeliscus from the Cape Verde Islands,
C. philippina from the Philippine Islands, and C. grandis from the Paumotos Islands are
only varieties or developmental phases of C. alata, the oldest species.

Carybdea alata var. pyramis Haeckel.

Charybdea pyramis, Harcxket, 1880, Syst. der Medusen, p. 440, taf. 25, fign. 5-8.
(?) Charybdea obeliscus, HarcKxet, Ibid., p. 441.
(?) Carybdea alata, ReyNauD, 1830, Lesson’s Centurie Zoologique, p. 95, planche 33, fig. 1.

Bell about 30 mm. high and 20 mm. broad. A 4-sided truncated pyramid, the upper
part being about one-third and the lower two-thirds as wide as the bell-height. A deep
interradial furrow bordered by a pair of prominent ridges extends down the 4 sides of the
exumbrella. The very long, narrow pedalia are lancet-shaped and about as long as the
bell-height. The 4 perradial sense-clubs have each 6 eyes and are set in deep niches on the
sides of the bell. Bell-margin displays 8 shallow lappets, the clefts being occupied by the
4 frenule and the 4 pedalia. The 4 frenulz which support the wide velarium are 3-cornered
and thick. 24 simple, 3-cornered velar canals, 6 in each quadrant. The stomach is not quite
as wide as the bell-radius, and is shallow and quadratic. Neck large, 4-sided, pyramidal,
and separated from the stomach by a deep stricture.
4 large, 3-cornered lips with folded edges. Gonads 8
wide leaves with irregular, crenulated, free margins. This
form is found in the tropical regions of the Atlantic.

Carybdea alata var. grandis Agassiz and Mayer

Charybdea grandis, Acassiz, A. and Mayer, 1902, Mem. Museum Comp.
Zool. at Harvard College, vol. 26, p. 153, plate 6, figs. 26-31.— BiceLow,
H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p. 17-
(?) Charybdea grandis, Browne, 1905, Report Pearl Oyster Fisheries, Gulf of
Manaar, Suppl. Report No. 27, Roy. Soc. London, p. 157.
?) Bursarius cytheree, Lesson, 1829, Voyage de la Coquille Zoophytes,
B 9, Voyag' q PAY
p- 108, planche r4, fig. 1.
(2) Tamoya bursaria, Maas, 1903, Scyphomedusen der Siboga Exped., Monog.

II, Pp. 4.

This medusa may be identical with Lesson’s Bur-
sarius cytheree but the latter is so vaguely described and
figured that I feel obliged to omit it from serious con-
sideration.

Adult medusa.—Bell high, pyramidal, with blunt

apex. 230 mm. high, and 140 mm. wide. Gelatinous

) & 4, substance quite thick and of remarkably tough consis-

E : tency. There are 4 interradial, wing-like pedalia, the

Fic. 329.—Garybdea alata, var. grandis. Young broad sides of which extend outward ina radial direction.

medusa, after Agassiz and Mayer, in “These pedalia are each about 40 mm. long, and 25 mm.

eae ees award des oA wide canal extends through the substance of

each of them into the flexible portion of the tentacle,

which arises from distal end of pedalium. The flexible part of the tentacle is about 140 mm.

long and its surface is ringed with transverse rows of nematocyst-cells. Each of the 4 per-
radial sense-organs arises from a deep niche about 27 mm. above level of velarium.

The sense-organ is knob-shaped, mounted upon a short stem, and contains from I to 3
ectodermal ocelli and an entodermal concretion. In old medusz there is usually a single,
median ocellus in each sense-organ, but a young specimen 30 mm. in height had a large
median and 2 small lateral ocelli. These ocelli are all directed as if to perceive objects within
the bell-cavity. Velarium well developed and suspended by 4 mesenteries or frenulz in the
radii of the sense-organs. 24 short, tree-like, non-anastomosing velar canals extend centrip-
etally inward into the substance of the velarium. Manubrium short, 4 slightly recurved
lips. There are 4 interradial crescentic areas of numerous short, gastric cirri, the horns of
each crescent pointing centripetally. 4 wide perradial pouches, extending outwardly from
the stomach, are separated one from another by 4 narrow interradial partitions, but are placed
in communication one with another by means of lateral canals leading into the lumen of the

512 MEDUS OF THE WORLD.
pedalia. The gonads consist of 8 leaf-like folds attached to the sides of the interradial septa
and hanging free in the radial pockets.

Gelatinous substance of bell is hyaline; entoderm translucent and milky-white; gastric
cirri and flexible parts of tentacles pink or yellow-pink; sensory knobs of rhopalia dull ocher
in color; ocelli deep brown, almost black.

This species is by far the largest Carybdea known.

Found at Fakarava and at Anaa Islands, Paumotos Islands, South Pacific by the Alba-
tross in October, 1899. A large swarm upon the surface at Anaa Island.

This medusa may be identical with the vaguely described Bursartus cytheree Lesson,
from New Guinea, or with Tamoya bursaria briefly mentioned by Maas, from the Malay
Archipelago. Bigelow, 1909, believes that C. moseri is only the young of this medusa.

Carybdea alata var. moseri Mayer.
(?)— —, Semper, 1860, Zeit. fiir wissen. Zool., Bd. 13, p. 561, taf. 39, fig. 9.
(?) Charybdea philippina, Harcxet, 1880, Syst. der Medusen, p. 440. =
Charybdea moseri, Mayer, 1906, Bull. U.S. Fish Commission for 1903, vol. 23, part 3, p- 1135, plate 1, figs. 2-2c.

This common Hawaiian Island medusa may be only a small variety, or a young stage,
of Carybdea grandis. (See table of synopsis of the species of Carybdea.) It is probably
identical with a medusa briefly mentioned and inadequately figured by Semper, from the
Philippine Islands.

Bell 80 mm. high, 47 mm. wide, dome-shaped, with flat top and thin, uniform walls. Each
sense-club has 2 large median and 2 small lateral eyes. The sensory niches are long, trans-
verse, narrow, and 14 mm. above the velar margin. Pedalia spatula-shaped, flat, expanded,
25 mm. long, 17 mm. wide, 24 simple, unbranched, velar canals. Tentacles ringed, tapering
throughout their length, hollow, and about 1.5 times as long as bell-height. Gonads not quite
as long as the septa on which they are developed. Stomach small, flat, 4 simple lips, gastric
cirri simple and unbranched. Honolulu, Hawaiian Islands.

Carybdea murrayana Haeckel.
Charybdea murrayana, Harcxet, 1880, Syst. der Medusen, p. 442; 1881, Report Deep-sea Meduse Challenger Expedition, Zool.,

vol. 4, p. 93, plate 26, figs. 1-10.

Bell 50 mm. wide, 60 mm. high, quadratic below, with flatly dome-like top. Pedalia
narrow, tapering, flattened laterally, one-third as long as bell-height. Each sense-club has
2 large median and 4 small lateral eyes, as in C. marsupralis. Velarium wide, with 48 pro-
fusely branching, non-anastomosing canals. The 4 clusters of gastric cirri in the interradial
corners of stomach are profusely branched, as in C. marsupialis.

Off coast of Sierra Leone, west coast of Africa. Depth of 200 fathoms.

Distinguished from C. marsupralis by its large number of velar canals.

Genus TAMOYA F. Miiller, 1859.
Tamoya, Miter, F., 1859, Abhandl. Naturf. Ges. Halle, Bd. 5, p. 1.—Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 174.—
Harcxe, 1880, Syst. der Medusen, p. 442.
The type species is T. haplonema F. Miiller of the Atlantic coasts of North and.South
America, south of Cape Cod.
GENERIC CHARACTERS.

Charybdeide with 4 simple, interradial tentacles provided with pedalia. Stomach wide
and deep; its 4 perradial sides flattened so as to present the superficial appearance of mesen-
teries binding the 4-sided cesophagus to the subumbrella. There are 4 vertical, interradial,
thread-like or brush-like bands of gastric cirri in the middle of interradial sides of stomach.

The so-called mesenteries of Haeckel are merely the flattened, perradial sides of the cruci-
form stomach.

Haeckel’s Tamoya bursaria and T. gargantua are too imperfectly known to be retained
in scientific literature.

This genus is very closely related to, if not identical with, Carybdea, being distinguished
only by its large stomach with its perradial mesenteries and its vertical clusters of gastric cirri,
It may eventually prove necessary to unite this genus with Carybdea, for the differences
between them are merely of an intergrading character. ;

CARYBDEID#—TAMOYA, TRIPEDALIA. 513

Tamoya haplonema F. Miiller.
Plate 57, figs. 2 to 2”.

Tamoya haplonema, Miter, 1859, Abhandl. Naturf. Ges. Halle, Bd. 5, p. 1, taf. 1, 2—Acassiz, L., 1862, Cont. Nat. Hist. U.S.,

vol. 4, p. 174-—HaeckeL, 1880, Syst. der Medusen, p. 443.—Brooks, 1882, Studies Johns Hopkins Univ. Biol. Lab., vol. 2,

p- 138.—von LenpeEnFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 245.—Mayer, 1904, Memoirs Nat. Sci.

Museum Brooklyn Inst. Arts and Sci., vol. 1, p. 28, plate 7, figs. 60-64.

Tamoya prismatica, Harcket, Ibid., p. 443-
Carybdea (Tamoya) haplonema, Fewxes, 1889, Report U. S. Commis. Fish and Fisheries for 1886, p. 526.

Bell 90 mm. high, 55 mm. wide, with vertical sides, and relatively flat top. Exumbrella
surface thickly covered with white, wart-like clusters of nematocysts. 4 pedalia, 30 mm.
long, flat, spatula-shaped, and sharp-edged. Tentacles go mm. long, hollow, very flexible
and bearing regularly spaced rings of nematocysts that are capable of inflicting a severe sting
to the hand. The sensory-clubs have 2 large median and 4 small lateral eyes, all being upon
the inner side of the bulb. The large eyes are provided with prominent convex lenses and are
ectodermal. There is a large terminal mass of concretions of entodermal origin.

The velarium is well developed and there are 10 dendritic velar canals in each quadrant,
which terminate in numerous, non-anastomosing branches. The nerve-ring running from
the base of each pedalium to the sensory-clubs is distinctly visible as a white-colored cord.
The stomach extends about a third of the distance from inner apex to level of velarium, and
there are 4 slightly recurved lips. Gastric cirri short and numerous.

The 8 genital organs are curtain-like sheets with frilled edges, which project from the
4 interradial septa into the perradial gastrovascular pouches of bell on either side. In old
specimens the gonads are so large that their free edges overlap beyond the central line of each
perradial stomach-pouch.

Gelatinous substance of bell transparent. The long, flexible tentacles are milky-yellow,
often with a faint purple hue. There are large, white, wart-like clusters of nematocysts over
the pedalia and velarium. The genital organs are milky-yellow, the eyes dark brown.
This medusa is exceedingly active, the gelatinous substance of its bell is tough and rigid.
Tamoya haplonema is widely distributed, having been found on the coast of Brazil, in
the West Indies, at Beaufort in North Carolina, and in Great Peconic Bay, and Branford
Harbor, Long Island Sound, New York, in the autumn. Our figures were obtained from a
specimen captured at the last-named place early in September, 1902. None was found upon
the surface in Great Peconic Bay, but all were brought up in dredges from the bottom at
depths of a fathom or more.

Haeckel’s Tamoya prismatica from the West Indies is apparently identical with 7. hap-
lonema. It is described as follows:

Bell 80 mm. high, 40 mm. broad, pyramidal, and 4-sided. The pedalia are longer and
narrower than in 7. haplonema. They are wedge-shaped, and 3 times as long as broad,
and about half as long as bell-height. In their upper parts they are 3-sided, but below they
are 2-sided with small meridional wings. Velarium very wide, with numerous, narrow,
branching canals. Stomach large, occupying upper third of bell-cavity. Throat-tube about
as large as stomach, 4 prominent lips. Color (?) Marginal sense-organs ( ?)

This form is found in the West Indies. It is probably only a variety of T. haplonema.

l

Genus TRIPEDALIA Conant, 1897.

Tripedalia, Conant, 1897, Johns Hopkins Univ. Circulars, No. 132, p. 9; 1898, Mem. Johns Hopkins Univ. Biol. Lab., vol. 4,
No. I, p. 5-

The type species is T. cystophora, described by Conant from Kingston Harbor, Jamaica.

GENERIC CHARACTERS.

Carybdeide with 4 interradial groups of tentacular pedalia, each tentacle being mounted
upon a separate, unbranched pedalium which arises from the bell-margin. Velarium with
canals and with 4 perradial frenule. No hernia-like sacs project into the bell-cavity from
the main stomach-pouches of umbrella.

This genus is very closely related to Chiropsalmus, but the pedalia themselves do not
branch, but each pedalium of each cluster arises separately from the interradial corner of the

514 MEDUSA OF THE WORLD.

bell-margin. In Chiropsalmus, on the other hand, each of the 4 pedalia gives rise to side
branches which bear tentacles. Moreover, in Tripedalia there are no subumbrella, hernia-
like, gastric diverticula as in Chiropsalmus.

Tripedalia cystophora Conant.
Tripedalia cystophora, Conant, 1897, Johns Hopkins Univ. Circulars, No. 132, p. 9, fig. 9; 1898, Mem. Johns Hopkins Univ.

Biol. Lab., vol. 4, No. 1, pp. 5, 22; figs. 17-30, plates 1, 2, 5, 7, figs. 44, 45, 53-56, 68, 71.

Bell cubical, with edges slightly rounded; 12 mm. high, and of about 15 mm. wide.
There are 4 interradial groups of pedalia, each group consisting of 3 distinct, separate pedalia,
each one of which arises from the bell-margin and gives rise to a single tentacle. The pedalia
are flattened and resemble a slender knife-blade, and
are about half as long as the bell-height. The 12
tentacles are each about 2.5 times as long as the
pedalia. 4 sense-clubs are situated in niches at about
one-fifth or one-fourth the height of bell above mar-
gin. Each sense-club has 2 large, median and 4 small,
lateral eyes and a terminal lithocyst. The median
eyes have doubly convex lenses. Velarium about
one-sixth as broad as bell-diameter. There are 24
simple, unforked velar canals, 6 in each quadrant.
These velar canals are triangular in outline, and the
8 adjacent to the 4 frenule are only half as wide as
the others. Stomach wide and shallow, butthethroat-
tube is long and extends downward in some cases to
bell-margin; cruciform in cross-section, with 4 well-
developed, oral lobes in the radii of the sense-organs.
There are 15 to 21 organs, resembling lithocysts, in
the gelatinous walls of the manubrium; each con-
sists of a round or oyal sac lined with ciliated cells
which keep in motion and bear up an irregular,
coarsely granulated concretion. These organs are
scattered irregularly through the gelatinous substance and are probably of entodermal origin.
The small, tapering, gastric cirri are brush-shaped and spring from 4 short stalks in the inter-
radial corners of the stomach. There are 4 wide, perradially situated, gastrovascular
pouches in the umbrella, which are separated by 4 interradial septa; but these septa are
incomplete in the regions of the pedalia, and thus the 4 stomach-pouches are placed in com-
munication one with another, as in other Charybdeide. The gonads are 8 leaf-like sheets
attached to the sides of the 4 interradial septa and projecting out into the 4 perradial
stomach-pouches. ‘The medusa is light yellowish-brown, the gonads being of the same color.

Figure 330 shows a mature female, 4 times natural size, drawn from nature, by the author.
In order to illustrate their shape, the lips are shown twisted 45° from their natural position.

This species is found in Kingston Harbor, Jamaica, in great abundance during the sum-
mer among the mangrove roots of the islands in a shallow, muddy lagoon on the western side
of the harbor, north of Port Henderson. It disappears in winter.

The dimensions of the mature specimen here figured are as follows: Bell 12 high, 15
mm. wide. Pedalia 5 mm. long, 2.1 mm. wide. Rhopalia 2.25 mm. above velarium margin.
Stomach 5.5 mm. wide, 7 mm. long. The gonads were mature and the gastrovascular space
filled with swimming planula. This medusa was captured on May 24, 1909.

Conant succeeded in obtaining females having embryos within their gastrovascular
pouches. The embryos were thrown out into the water as free-swimming planula, which
settled down on the bottom and sides of the aquarium in a day or two and quickly developed
into small Scyphostoma with mouth and typically with 4 tentacles and 4 taniolz, although
3 and 5 tentacled specimens were not uncommon. In this condition they lived for 3 weeks
without essential change. I find that many, but not all of the planulz, are besprinkled with
dark brown pigment-spots which are scattered over the ectoderm of the narrow posterior end
of the larva.

Fic. 330.—Tripedalia cystophora.

Fig.

Fig.
Fig.
Fig.
Fig.

I.

Pirate 58.
All figures are of Carybdea marsupzalis.

Mature medusa, 1.25 times natural size. Naples Zoological Station,
December 5, 1907.

. Sense-club seen from exumbrella side.

. Side view of sense-club.

. One of the velar canals showing its branches.

. One of the gastric cirri showing its brush-like terminal branches.

Drawn from life, by the author.

MAYER

PLATE 58

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CARYBDEIDA—CHIROPSALMUS, 515

Genus CHIROPSALMUS L. Agassiz, 1862.

Chiropsalmus, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 174.—Harcket, 1880, Syst. der Medusen, p. 446.—von Len-
DENFELD, 1884, Proc. Linnean Soc, New South Wales, vol. 9, p. 247.

GENERIC CHARACTERS.

With 4 interradial, branched pedalia which give rise to a number of tentacles. 4 wide
perradially situated stomach-pockets in the subumbrella, and each of these gives rise to
finger-shaped, unbranched, hernia-like pouches which project into the bell-cavity. Wide,
marginal pouches and numerous canals in the velarium. 8 leaf-shaped gonads.

The type species of this genus is Chiropsalmus quadrumanus, described as Tamoya
quadrumanus by F. Miller, 1859. This species is found in the warmer waters along the
Atlantic coasts of North and South America south of Cape Hatteras.

Chiropsalmus quadrumanus L. Agassiz.
Plate 57, fig. 3.

Tamoya quadrumanus, Mix.er, F., 1859, Abhandl. Naturf. Gesell, Halle, Bd. 5, pp. 1-11, taf. 2, 3, fign. 18-32.
Chiropsalmus quadrumanus, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 174-—Harcxet, 1880, Syst. der Medusen, p.

447—von LeNnDENFELD, 1884, Proc. Linn. Soc. New South Wales, vol. 9, p. 248.—Conanrt, 1898, Mem. Johns Hopkins

Univ. Biol. Lab., vol. 4, No. 1, p. 4

Bell dome-shaped, about 140 mm. wide and 100 mm. high. 4 large, hand-shaped
pedalia, the 7 to g finger-like branches of which give-tise each to a single, long, slender tentacle.
7 to g tentacles thus arise from each pedalium. These tentacles are hollow and flexible, and
are covered with numerous, closely set rings of nematocysts. A large axial-canal extends
through the pedalium and gives off branches, one to each tentacle. The 4 sense-organs are
situated within 4 covered niches upon sides of bell at about one-sixth the distance from margin
to apex. There are 6 ectodermal eyes upon the sense-club, 2 large median and 4 small lateral.
Velarium very wide with 16 large branched pouches which extend into it from the 4 main,
gastrovascular spaces of the bell. The 16 velar pouches give rise in turn to numerous small,
branching canals which ramify through the velarium. Stomach wide and globular, the mouth
surrounded by 4 large, triangular lips. 4 wide, perradial pouches extend from stomach into
wall of bell; each of these pouches gives rise to 2 finger-shaped, hollow, hernia-like sacs
which project from the subumbrella side into the cavity of the bell; these 8 sacs are situated
very near base of stomach. There are 4 interradial, crooked, crescent-shaped rows of gastric
cirri on inner walls of stomach.

This species was found by Miiller at Desterro, Santa Catharina, Brazil, and later it was
obtained by H. V. Wilson at Beaufort, North Carolina, where it is quite abundant on the
sea-bottom, about a mile off shore, though sometimes found within the harbor itself.

Chiropsalmus buitendijki Horst.

Chiropsalmus buitendijki, Horst, 1907, Notes from the Leyden Museum, vol. 29, No. 2, p. tor, plate 2, figs. 1-6.

This species from the harbor of Batavia, Java, is distinguished by its 8 long, simple,
finger-shaped, subumbrella pocket-arms nearly as long as the depth of the bell-cavity, so that
they almost touch the velarium. The pedalia also have 5 or 6 lateral branches arranged,
judging from Horst’s figure, in a linear series on the outer side of the main shaft, the largest
branch being nearest the bell. In other species of Chiropsalmus the side branches of the
pedalia are irregularly arranged.

In other respects the Javanese medusa resembles the other species of Chiropsalmus. Bell
cubical, 65 to 70 mm. high and wide, with a slightly arched apex. Main shafts of pedalia
sickle-shaped, about half as long as bell-height, and laterally compressed. The 5 or more
lateral branches all arise from the abaxial side of the pedalium and form a decreasing series,
the smallest being outermost. The rhopalia are in niches about one-fifth the distance between
velarium and bell-apex. Each club has 2 large median and 4 small lateral eyes. The velari'um
is wide, has 4 frenulz, and 16 dendritically branched velar canals. Bell transparent, flexible
parts of tentacles of a rosy hue. In its simple finger-shaped processes of the subumbrella floor
of the bell this species resembles the American Chiropsalmus quadrumanus to which it is
closely related.

516

MEDUSZ OF THE WORLD.

Chiropsalmus quadrigatus Haeckel.

lripatus, Haecker, 1880, Syst. der Medusen, p. 447-

The following description is based upon a study of six specimens obtained in a seine

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Pe} Wu

by the U. S. Bureau of Fisheries
steamer Albatross among the Phil-
ippine Islands, from January to
June, 1908.

Bell dome-like and 4-sided, with

an evenly rounded, aboral apex. 70
to 100 mm. high, 80 to 100 mm. wide
at level of velarium. There are 4
hand-shaped, interradial pedalia,
laterally flattened, which arise from
the sides of the bell at a short dis-
tance above the velar margin.
These pedalia are about 30 mm.
long and 11 mm. wide, and each
bears 5 to g finger-shaped terminal
projections which give rise to an
equal number of long, flexible, hol-
low tentacles. These tentacles are
thickly but somewhat irregularly
ringed with nematocysts. They
vary in length, but the longest when
contracted are about 150 mm. long.
The pedalia of a medusa 100 mm.
wide were each 47 mm. long and
21 mm. wide.

The 4 perradial sense-clubs are
set within covered niches in thesides
of the bell about 14 mm. above the
velar margin. Each sense-club’con-
tains an entodermal, terminal, abax-
ial mass of concretions and on its
Fic. 331-—Chiropsalmus quadrigatus Haeckel, side view of half-grown inner side are 6 eye-spots. The 2

medusa, half natural size. Drawn by the author, from a preserved median eyes have each a prominent

specimen collected by the Albatross. nv e =
A, enlarged side view of sense-club. B, inner side of sense-club. cook lens, but th i, smaller, lat

C, oral view of stomach showing the 4 lips and 8 gastric sacs. eral eyes are merely pigmented
D, enlarged view of gastric cirri. E, side view of a pedalium ocelli. The eyes are arranged so as

with all but one of the tentacles cut across. to view objects withinthe bell-cavity.

The yelarium is 13 mm. wide and supported by 4 bracket-like perradial frenula. There
are about 50 dendritic, non-anastomosing velar canals, 12 to 15 in each quadrant. The
velarium is diaphragm-like and does not hang downward beyond the velar margin but stretches
flatly across tending to close the opening of the bell-cavity.

The wide central stomach is only about 20 mm. long and there are 4 lanceolate lips
with entire, simple margins. The 4 perradial sides of the cruciform stomach are much flattened
and form the so-called ‘‘mesenteries” of Haeckel, bridging across, bracket-like, between the
subumbrella and the 4-sided cesophagus. 4 pairs of gastric saccules arise from the perradial
sides of the stomach and project downward into the bell-cavity (c. fig. 331). Each saccule is
laterally flattened, cock’s-comb-shaped, with an irregularly notched margin, and is about
20 mm. long and 11 mm. wide. A solid, gelatinous projection extends downward so as to fill
the greater part of the cavity of each gastric saccule.

There are 4 long, interradial bands of gastric cirri marking the borders of the central
stomach. These cirri are fusiform, simple, sharp-pointed, and unbranched and arise in 4 or
5 rows. Each cirrus is about 3 mm. long.

CARYBDEIDA—CHIROPSALMUS. 517

The peripheral, gastrovascular system consists in the usual 4 wide, perradial stomach-
pouches, separated by 4 narrow, interradial septa, which are interrupted at the point of origin
of each pedalium where a canal extends downward from the adjacent stomach-pouches and
branches in finger-like ramuli leading into the tentacles of the pedalium. The 50 or more
velar canals have already been described. The 8 leaf-like gonads are well developed and
resemble those of Carybdea but are not yet mature. They arise on both sides of each inter-
radial septum and project into the-wide, perradial stomach-pouches.

The bell is milky-yellow in formation and there are traces of a dull purple-pink coloration
in the tentacles. The ocelli are dark brown.

I am informed that this medusa is abundant in Subig Bay, Luzon, Philippine Islands,
where it is captured in seines. The natives are said to preserve it in vinegar, and when so
prepared it is sold in the markets for food.

In the young medusa having a bell 18 mm. high and 20 mm. wide, there are only 4 or 5
tentacles on each pedalium and the subumbrella saccules have not begun to develop, nor is
there any visible trace of the gonads. The abaxial finger of each pedalium is longest, and
the 3 others are evidently of more recent development, are very short, and lie nearer the velar
margin. The longest (abaxial) tentacles are about 7 mm. long when contracted, and the
axially placed younger tentacles are shorter. The perradial brackets of the velarium are
beginning to develop and there are about 7 irregularly but simply branched velar canals in
each quadrant.

Haeckel, 1880, describes a somewhat larger young specimen from Rangoon, Indian
Ocean. The medusa is common among the Philippine Islands.

I am told that this medusa grows to be about 150 mm. in diameter, although the largest
specimen in the Albatross collection is only 100 mm. wide, but appears to be mature. It is
found swimming in shallow water near the shore. It is very closely allied to the Atlantic
C. quadrumanus, but may be distinguished by its laterally flattened cock’s-comb-shaped
gastric saccules, those of C. quadrumanus being finger-shaped.

The following is a record of specimens of Chiropsalmus quadrigatus obtained by the
Albatross in 1908 in the Philippine Islands:

Locality. Date. Height of bell} Width of bell | Number of tentacles! Remarks.
in mm. in mm. upon each pedalium,
Cataingan Bay, Masbate, April 18 49 55 6, 6, 7,7 Immature. The 8 gastric
near shore. saccules only beginning to
appear.
Do. April 18 51 51 8, 6, 7) 5 Do.
Do. April 18 40 20 6, 5, 6, 6 Do.
Subig Bay, Luzon. January 7 18 20 4> 49 4) 4 No gastric saccules and no
Caught in a seine. gonads.
Do. January 7 7° 80 9, 5, 8, 8 With well-developed but
See fig. 331 immature gonads. With
large cock’s-comb-shaped
gastric saccules.
Mausalay, Mindanao. June 4 97 100 8, 8, 8, 8 Apparently mature.

Chiropsalmus zygonema Haeckel.
Chiropsalmus xygonema, Haxrcke1, 1880, Syst. der Medusen, p. 641.

Bell 4-sided, pyramidal with a truncated apex, 60 mm. high and 40 mm. wide, including
height of velarium. The 4 pedalia are leaf-shaped, each with only 2 short, asymmetrical,
gelatinous, finger-shaped branches which give rise to long, lash-like tentacles. In all, there-
fore, there are but 8 tentacles. Stomach round and sac-like, cesophagus small with 4 lappets,
though only half as long as stomach. 4 interradial bow-like areas of gastric cirri in stomach
wall. The stomach gives rise to 4 perradially situated pouches, on both sides of the entrance
to each of these pouches near the stomach are 2 very small, oval pouch-arms—8 in all.

This species is found off the Argentine coast, South America. It differs from all other
species of Chiropsalmus in its simple, 2-pronged pedalia, and in the very srall finger-like
pouches of the stomach. The velarium and gonads are similar to those of C. quadrumanus.

518 MEDUSE_OF THE WORLD.

Genus CHIRODROPUS Haeckel, 1880.
Chirodropus, Harcxet, 1880, Syst. der Medusen, p. 447.

The type species is Chirodropus gorilla Haeckel, of the Atlantic coast of equatorial Africa.

MEE GENERIC CHARACTERS.

Carybdeidz with 8 branched, or feathered, sac-like
projections which extend from the 4 perradial stomach-
pouches into the bell-cavity. The free margins of the
gonads show grape-like swellings. With 4 hand-like
pedalia.

This genus resembles Chiropsalmus, but is distin-
guished by its branched hernia-like pouches in the bell-
cavity, whereas in Chiropsalmus these projections are
unbranched; moreover, in Chiropsalmus, the free mar-
gins of the gonads are simple and entire, whereas in
Chirodropus they are broken up into grape-like clusters
of swellings.

Chirodropus gorilla Haeckel.
Chirodropus gorilla, HAecKEL, 1880, Syst. der Medusen, p. 448, taf. 26,

fign. 1-8.

Bell dome-shaped, evenly rounded above and in-
cluding the velarium, which hangs downward, 150 mm.
high and 120 wide. Sides rounded; the sculptured
angles usually seen on surface of exumbrella in Charyb-
deide are very poorly developed.

The 4 interradial pedalia are hand-like, asymmet-
rical and hardly one-fourth as long as bell-height. Each
pedalium gives rise to 9 long, narrow, tapering gelati-
nous finger-like processes, each bearing a long, flexible,
tapering tentacle longer than bell-height. Thus there
are 36 tentacles in all.

The 4 perradial sensory niches are deep and heart-
shaped. They are set in the sides of the bell at a some-
what higher level than the pedalia and are about 60 mm.
If above velar margin. The sense-club is mounted upon

I a slender stalk. Number of eyes (?)
/ Fic. 332-—Chirodropus gorilla, Below the marginal nerve-ring are 16 gelatinous

after Hseckel, im Das Syst Ja ypets which form an integral part of the wide velarium
| which projects downward. The 16 lappets are cleft in
the 4 interradii and in the 8 adradii, and are divided
by the 4 perradial frenule. The 8 lappets flanking the 4 frenulz are about 1.5 times as
long as the 8 which flank the 4 interradial pedalia. These lappets contain diverticula of the
perradial stomach-pouches and each lappet-pouch gives rise to about 6 dendritically branched
velar canals, which only occasionally anastomose and which extend outward nearly to margin
of velarium, running mainly parallel one to another.

Each perradial stomach-pouch gives rise to a pair of elongate, tapering, hollow processes,
which project downward into the bell-cavity and bear numerous finger-like side processes,
all of which arise from inner (axial) side of main process. These processes are thus much
more complex than the simple finger-shaped ones of Chiropsalmus. Central stomach large
and urn-shaped, 4-sided, bound to subumbrella by 4 perradial mesenteries. There are 8
dendritic, digestive glands, a pair on either side of each perradial corner of stomach-cavity.

The 4 pairs of gonads are attached to the subumbrella sides of the 4 interradial septa,
as in other Carybdeida. Their free outer margins bear grape-like clusters of swellings. The
only specimen studied by Haeckel was a male, found at Chenchozo Loango, Lower Guinea,
about 5° S. lat. West coast of Africa. Described in detail by Haeckel who is the only natur-
alist who has seen the medusa.

CARYBDEIDA—CHIRODROPUS. 519

Chirodropus palmatus Haeckel.
Chirodropus palmatus, HAEcKEL, 1880, Syst. der Medusen, p. 448.

This is possibly only the young of a variety of C. gorilla being smaller, and only 70 mm.
wide and 100 mm. high, including the suspended velarium. The 2 pouch-arms which project
into the bell-cavity from each perradial stomach-pouch are fused together in their upper
two-thirds, leaving only their lower thirds free. They bear numerous filaments, as in
C. gorilla. Finally, each pedalium bears 21 fingers instead of 9, as in C. gorilla.

A single specimen is described by Haeckel from preserved material found near St. Helena
off the west coast of Africa.

We can not be sure that this form is distinct from C. gorilla until we know the normal
limits of variability of the latter species, and it seems probable that there is but a single species
which is somewhat variable in the number of tentacles, etc.

Order STAUROMEDUSE.

Lucernaride (in part), JouNsTON, 1847, British Zooph., p. 244, 2d ed.

Stauromedusa, Harcxet, 1880, Syst. der Medusen, p. 363.—Goertre, 1887, Abhandl. zur Entwickelungsgesch. der Thiere, Heft
4, p- 64.—Maas, 1906, Fauna Arctica, Bd. 4, Lfg. 3, p. 499; 1907, Ergeb. Fortschitte der Zoologie, Bd. 1, pp. 194, 198.

Lucernaride, GRoss, 1900, Jena. Zeit. fiir Naturw., Bd. 33, p. 611.—KassIANow, 1901, Zeit. fiir wissen. Zool., Bd. 69, p.287.

Lucernaria, CLark, 1863, Boston Journ. of Nat. Hist., vol. 7, p. 531-

Neglecting for the moment to consider the family Tesseranthine Haeckel, the Stauro-
medusz constitute a unitary group of sessile Scyphomedusz which are attached to objects
by means of an adhesive pad at the extremity of the aboral stalk of the bell. The bell-margin
usually exhibits 8 adradial lobes, the pointed ends of which terminate in clusters of hollow
knobbed tentacles; although in Capria we find that these tentacles are not knobbed, and
in the genus Stenoscyphus there are no marginal lobes, while in Lipkea there are no tentacles
and the lobes are perradial and interradial, not adradial.

In the 8 perradial and interradial notches between the marginal lobes we may find a
knobbed tentacle, which may be metamorphosed into an adhesive organ serving as a sort of
anchor. In the genera Lucernaria, Kishinouyea, Craterlophus, Capria, and Lipkea, however,
these anchors are not found. The central stomach gives rise typically to 4 perradial pouches
which are partially separated one from another by 4 interradial septa, but these pouches
communicate one with another at the margin, thus forming a peripheral ring-sinus. The arms,
or marginal lobes, are hollow, as are also their knobbed tentacles. There are 4 interradial
septa in the central cavity of the aboral stalk or peduncle; and these partitions may fuse in
the center, thus inclosing 4 separate perradial cavities in the stalk as in Haliclystus. The
gonads, which are developed in the entoderm of the subumbrella, are typically interradial and
more or less horse-shoe-shaped with the free ends of the horse-shoe directed outward; but
often the horse-shoe is cleft in the middle, giving 8 adradial, crescentic gonads. There is a mar-
ginal ring-muscle in the subumbrella which may be entire or divided into 8 separate perradial
and interradial sectors. Centripetal to this ring-muscle system are the radial muscles. ‘The
stalk also has a system of 4 interradial, longitudinal muscles. As in scyphostoma larve the
4 interradial septa of Stauromedusz are not simple, solid-walled partitions, but each contains a
funnel-like pit, livid with ectoderm, which dips downward from the subumbrella thus hollowing
each partition. These funnel-cavities contain longitudinal muscles which extend downward
eyen to the aboral end of the stalk itself.

Clark, 1863, and after him Gross, 1900, have made careful studies of the internal anatomy
of the sessile Stauromeduse. They conclude that we may divide these forms into 2 families,
the Eleutherocarpide with 4 simple, perradial stomach-pouches, and the Cleistocarpide in
which the neighboring halves of the adjacent gonads unite at their distal ends in the radii of the
corners of the mouth. Thus the gonads become united by a transverse, circumferential mem-
brane, which divides each of the 4 perradial stomach-pouches into 2 spaces, an outer and an
inner, the oral or inner one of which forms a cul de sac or confined space which contains the
gonads and opens at the axial end into the gastric cavity. The genera Halimocyathus, Cra-
terlophus, and Depastrum are examples of the Cleistocarpida, while Stenoscyphus, Kishinouyea,

520 MEDUS.® OF THE WORLD.

Caria. Lipkea. Lucernaria and Haliclystus represent the more simply organized Eleuthero-
de. A clear understanding of these relations can best be obtained from a study of the
ires Gross, 1900 (Jena. Zeitsch. fiir Naturwissen., Bd. 33, p. 613, taf. 23, 24). Being
rnal characters, | have not made use of them in the classification of the Stauromeduse.

\s one would expect in sessile animals the Stauromedusz show evidences of degeneration,

of marginal lobes or of tentacles; and in none of them do we find the lithocyst-bearing

Ectoderm

Entoden ——

Gelatinous substance

Fic. 3324.—Median longitudinal perradial sections and cross-sections of Stauromeduse. Somewhat diagrammatic, after Gross.
A. The internal anatomy of the Cleistocarpide illustrated by Craterlophus tethys.
B. The internal anatomy of Eleutherocarpida illustrated by Haliclystus. The cross-partition, or claustrum (C), is
found in the Cleistocarpide but is absent in the Eleutherocarpida.
In both figures: C, transverse partition spanning between gonads; ex, outer chamber; fc, funnel-pits containing
longitudinal nucleus; g, gonads; gt, gastric cirri; /m, longitudinal muscles; s, interradial septa.

sense-organs seen in all other orders of Scyphomedusez. Eyes and “‘otocysts”

the Staurome dusz, and pulsation is not exhibited by the sessile forms.

‘There is reason to believe that the Stauromeduse are the most degenerate of all Scypho-
medusz and are to be regarded as sexually mature Scyphostome (see Kassianow, 1901;
Goette, 1887). Their degeneracy is amply accounted for by their sessile mode of life; Hornell,
1883 (Natural Science, London, vol. 3, p. 204), and Hurst (Jbid., p. 209) believe that they are

are absent in

STAUROMEDUS®. 521
descended from some more highly organized forms of Scyphomedusw. Their chief reasons
for this belief lie in the facts that inthe Lucernaride the marginal anchors (colletocystophores)
are highly variable and may therefore be vestigial organs. Moreover, in Lucernaria cam-
panulata the marginal anchors exist in the very young medusa but disappear as development
proceeds and are not found in the adult, This evidence, although meager, supports the

Synopsis of the Genera of Stauromeduse.

Tesseranthine: Pelagic
medusz with solid,
tapering, non-knobbed
tentacles, and without
marginal lobes.

Lucernarine: Attached medusa, usually with hollow, knobbed
tentacles, and typically with marginal lobes.

Tesseraria= Tessera+ Depastrum= Stenoscyphus. Lucernaria. Kishinouyea.
Tesserantha+ Tesseraria| Depastrella+
Haeckel. Depastrum
Haeckel.
Number and char- | 8 solid, tapering, without | Hollow, ending in| An anchor in each| None. None.

acter of perradial
and interradial
tentacles.

Character of adra-
dial tentacles.

Other tentacles.

Cleistocarpide are
indicated by C,
Eleutherocarpide
by E.

Gonads.
Stalk at aboral pole

of bell, serving for
attachment.

Marginal lobes of
bell.

terminal knobs.

Similar to perradial ten-
tacles.

Similar to perradial ten-
tacles.

4 interradial, horse-shoe-
shaped.

An aboral projection
which does not serve for
attachment Medusa
pelagic.

None.

nettling knob.
I to 3 tentacles
in each cluster.

Similar to per-
radial and inter-
radial, but with
more tentacles in
each cluster.

None.

(e

As in Tesseraria.

An aboral stalk
serving to attach
the medusa.

8 adradial sinuos-
ities.

interradius and
perradius.

A cluster of hol-
low, terminally-
knobbed tenta-
cles in each ad-
radius.

None.

E.

Adradial.

A stalk with 4
perradial cham-
bers.

None.

8 clusters of hol-
low, terminally-
knobbed tenta-
cles.

None.

E.

8 adradial (4 cleft
in interradii).

Single-chambered -

8 adradial lobes.

As in Lucernaria.

None.

As in Lucernaria.

With 4 perradial
chambers.

8 adradial.

Lucernarine: Attached medusz, usually with hollow, knobbed tentacles, and typically
with marginal lobes.

Number and charac-
ter of perradial ten-
tacles.

Character of adradial
tentacles.

Other tentacles.

Cleistocarpide are
indicated by C,
Eleutherocarpide by
E.

Gonads.

Stalk at aboral pole of
bell serving for at-
tachment.

Marginal lobes of the
bell.

With 4 perradial
chambers.

8 adradial.

As in Lucernaria.

4 interradial.

With 4 perradial
chambers.

8 adradial.

8 adradial.

With 4 perradial
chambers.

8 adradial.

8 adradial.
Single-chambered.

8 adradial.

Haliclystus. Halimocyathus. | Craterlophus Capria. Lipkea
Halimocyathus ( ?)
8 anchors. 8 anchors. None. None. None.
As in Lucernaria.| As in Lucernaria. | As in Lucernaria. | A row of basally None.
webbed, finger-
shaped, unknobbed
tentacles.
None. None. None. None. None.
E. fer (er E. ES

4 interradial (?)
Single-chambered.

Eight; 4 perradial
and 4 interradial.

522 MEDUS.Z OF THE WORLD.

view that the Stauromedusz are degenerate, and, indeed, degeneracy would very probably
ilt from their se ssile condition.

Che Tesseranthine, known only through the works of Haeckel, who alone has observed
‘ear to be pelagic and are said to differ from other Stauromedusz in having solid,
non-knobbed tentacles, and in lacking marginal lobes. Their relationship to the
Stauromedusz is uncertain. Only 4 specimens have ever been taken and these are described
by Haeckel from preserved material. It will be well to suspend judgment in respect to their
structure, relationship, and development until more specimens have been studied.

The sessile Stauromedusz are creatures of cold seas. None is known from the tropics.
They are littoral forms and attach themselves to Fucus, Zostera, and other seaweeds among
the tidal eddies of the coast. They are generally rare and only locally abundant in any case.

Owing to their rarity many of the species of Stauromeduse are imperfectly described,
and it is probable that some of them should be reduced. Kassianow, 1901, has studied the
nervous system of Lucernaria, Haliclystus, and Craterlophus, and the results of his studies
are here reviewed in the discussion of the genus Lucernaria, wherein I have also reviewed
the brief observations of Fol and of Kowalevsky, 1884, upon the early stages of the develop-
ment of Lucernarta.

The sessile Stauromeduse display no rhythmical pulsation in their bells, but no studies
have been made to discover whether the larve at any stage possess this power. Hornell,
1893, and Browne, 1896, have studied the variations of Haliclystus octoradiatus. The results
of their studies are referred to in the description of this species.

According to A. Meyer, 1865, the Lucernarida have great regenerative power. The bell
may reproduce a new stalk and parts of the medusa may regenerate the whole (see genus
Haliclystus). Kassianow, 1901 (Zeit. fiir wissen. Zool., Bd. 69, p. 371), reports upon some-
what similar experiments upon Craterlophus.

The relationship which may exist between the Stauromeduse and the Carybdeidz is
discussed in the introduction to this volume.

tnem, ap}

tapering,

Genera TESSERA, TESSERANTHA, and TESSERARIA Haeckel, 1880.
Tessera, Tesserantha, HarcKxe, 1880, Syst. der Medusen, pp. 373-3753 1881, Deep-Sea Medusz Challenger Exped., vol. 4, p. 49-
Tesseraria, HaecKet, 1880, Syst. der Medusen, p. 633.

Haeckel describes 4 specimens of these medusz from preserved material, and is the only
naturalist who has seenthem. They are said to differ from other Stauromedusz in their solid,
non-knobbed tentacles, and in their being free-swimming, non-attached forms.

Haeckel states that these medusz have a simple, uncleft, umbrella margin, no marginal
anchors, but 8 to 16 simple, perradial, interradial, and adradial tentacles. The broad marginal
ring-muscle of the subumbrella is entire, not divided into 8 isolated sectors. Centripetal to
the ring-muscle there is a system of radial-muscles. The medusz are pelagic and have no
stalk of attachment, although a hollow apical projection is found at the aboral pole of the bell.
The tentacles are solid and do not terminate in nematocyst-knobs.

The stomach gives rise to 4 wide, perradial, gastric pouches which are possibly divided
one from another in the 4 interradii by 4 short, narrow septa. These septa extend centrifugally
from the 4 gonads, but are so short that they do not reach the bell-margin, and thus there is a
wide, marginal, gastral ring-sinus. There are 4 interradial, horse-shoe-shaped gonads with
their free ends pointing outward. 4 simple lips and 4 interradial areas of gastric filaments in
the stomach.

According to Haeckel these 3 genera are distinguished as follows:

Tessera, with 8 tentacles, 4 perradial and 4 interradial. Bell 4 to 8 mm. wide. .
Tesserantha, with 16 tentacles, 4 perradial, and 4 interradial, and 8 adradial. Bell 6 mm. wide.
Tesseraria, with 32 tentacles. Bell 10 mm. wide.

In all respects these medusa resemble one another so closely that I am lead to suspect
that they may prove to be but stages in the growth of one and the same medusa. It will be
observed that the larger medusw have the greater number of tentacles.

The following is a brief diagnosis of the characters of these medusz, according to Haeckel:

Tessera princeps Haeckel, 1880 (Syst. der Medusen, p. 347, taf. 21, fign. 1-6), is 5 mm.
high and 4 mm. wide, with a pyriform bell and hollow apex. There are 4 perradial tentacles
about as long as the bell height, and 4 shorter (undeveloped ?) interradial tentacles. There

STAUROMEDUS—TESSERA, TESSERANTHA, TESSERARIA, DEPASTRUM. 523

are only 4 simple, interradial, gastric cirri, one in each interradius of the central stomach above
(centripetal to) the 4 horse-shoe-shaped gonads. A single specimen was found by the Chal-
lenger southeast of Kerguelen Island, Antarctic Ocean, on February 19, 1874.

Tessera typus Haeckel, 1880 (Lbid., p. 638), bell 8 mm. wide, 12 mm. high, pyramidal,
4-sided, elongate. 8 tentacles, 4 perradial and 4 interradial, of equal lengths, each nearly twice
as long as the bell-diameter. 6 to 8 gastric filaments in each interradius. 4 horse-shoe-shaped
gonads in proximal half of subumbrella. A single specimen from the Indian Ocean, south of
Madagascar.

T esserantha connectens Haeckel, 1880 (Syst. der Medusen, p. 373; 1881, Deep-sea Medusze
Challenger Expedition, Zool., vol. 4, p. 50, plate 15, figs. 1 to 8) differs from ‘“Tessera
princeps” in being larger and in having 16 tentacles and numerous gastric cirri. The
medusa is 9 mm. high and 6 mm. wide. The tentacles are perradial, interradial, and adra-
dial, the largest being the first named, and the last quite short and apparently immature.
There is an ectodermal pigment spot on the exumbrella side of the base of the 8 perradial
and interradial tentacles. A ridge of nettle cells extends toward the apex of the bell from
the base of each tentacle. These pigment spots and ridges are not seen in Tessera prince ps.
The numerous gastric cirri are arranged on both sides of the 4 interradial partial septa or
teniola of the central stomach. The 4 interradial pits of the subumbrella are deeper
than in Tessera. Altogether all of the differences between T essera and Tesserantha are such
as one would expect to find in one and the same medusa in advancing stages of growth.

Tesserantha connectens was found by the
Challenger near the island of Juan Fernandez,
South Pacific, at a depth of 2,160 fathoms. It is
elaborately described by Haeckel, 1881.

T esseraria scy phomeda Haeckel, 1880 (Lbid., p-
638), has a goblet-shaped bell, 10 mm. wide, 15
mm. high. 32 tentacles equal each to each and not
quite as long as the bell-diameter. 4 simple rows
of gastric cirri. 4 horse-shoe-shaped gonads.

Found in Bass Straits between Australia and
Tasmania. One specimen in Godeftroy Museum.

Genus DEPASTRUM Gosse, 1858.

Depastrum, Gossr, 1858, Annals and Mag. Nat. Hist., vol. 1, ser. 3,
p- 419.

Depastrum+ Depastrella, Harcxet, 1880, Syst. der Medusen, pp.
376, 378.

Depastrum, Dixon, G. Y. and A. F., 1893, Proc. Roy. Dublin Soc.,
vol. 8, p. 180.—Braumont, 1894, Trans. Liverpool Biol. Soc.,
vol. 7, p. 254.—Maas, 1906, Fauna Arctica, Bd. 4, p. 500.

Carduella, AttmMan, 1860, Report British Association, p. 143.—Clark,
1863, Boston Journal Nat. Hist., vol. 7, p. 546.

The type species is Depastrum cyathiforme of
the northern coasts of Europe, first described by M.
Sars, 1846, as Lucernaria cyathiforme. Gosse,
1858, established for it the genus Depastrum.

GENERIC CHARACTERS.

Stauromedusz with 16 clusters of tentacles, 4
perradial, 4 interradial, and 8 adradial, arranged
in one or in several rows around the bell-margin.
Tentacles are all similar each to each, and are hol-
low, and terminate each in a nematocyst-knob.
There may be one or more tentacles in each per-
radius and interradius. No marginal anchors.
With divided stomach-pouches as in the Cleistocarpide. An unbroken marginal, subumbrella
ring-muscle. 4 small, interradial partial septa in the central stomach, leaving a wide,
marginal ring-sinus. 4 interradial horse-shoe-shaped gonads with their convexities inward

Fic. 333.—Tesserantha connectens, after Haeckel in Deep-
sea Meduse of Challenger Expedition.

524 MEDUSZ OF THE WORLD.

and free ends directed centripetally. A 4-sided throat-tube. An aboral stalk serving for
attachment. The adradial lobes are reduced to 8 barely discernible sinuosities.

Depastrella Haeckel, with 16 clusters of marginal tentacles arranged in a single row, is
probably only the young of Depastrum. Depastrella appears to be intermediate between the

Tesseranthinz and the Lucernarine.

Depastrum cyathiforme Gosse.

Lucernaria cyathiforme, Sars, M., 1846, Fauna Littoral, Noweg., fasc. 1, p. 26, taf. 3, fign. 8-13.

athiforme, Gosse, 1858, Annal. and Mag. Nat. Hist., vol. 1, p. 419; 1860, Ibid., vol. 5, p. 481, figs. 1-3.

siformis, Kererstein, 1862, Zeit. fiir wissen. Zool., Bd 12, p. 24.

yathiformis, ALLMAN, 1860, Trans. Microscop. Soc. London, vol. 8, p. 125, plate 5.—Clark, 1863, Boston Journal Nat.
Hist., vol. 7, p. 546. :

Calicinaria cyathiformis, Mune-Epwarps, 1860, Hist. Nat. der Corall., tome 3, p. 459, Paris.

Depastrum cyathiforme, Harcxer, 1880, Syst. der Medusen, p. 378 (literature); D. polare, p. 639, and Depastrella carduella,
p- 376. ; §
Depastrum cyathiforme, Dixon, G. Y.and A. F., 1893, Proc. Roy. Dublin Soc., vol. 8, p. 180.—Braumonr, 1894, Trans. Liverpool
Biol. Soc., vol. 7, p- 254-—Russext, 1904, Annals and Mag. Nat. Hist., ser. 7, vol. 13, p. 62, plate 5 (references to localities
on the British coast)—Browne, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 774 (found in the Firth of Clyde at Little

Cumbre and near Keppel pier)—Maas, 1906, Fauna Arctica, Bd. 4, p. 500.

Medusa urn-shaped, about 6 to 10 mm. wide and of somewhat greater height. Stalk
about as long as bell-height, flexible, contractile, and with an irregularly expanded adhesive
foot. Bell-margin sinuous, subumbrella deeply concave. 36 to 100 tentacles arranged in 4
perradial and 4 interradial clusters of 1 to 3 tentacles each, and 8 adradial clusters, each con-
sisting of about g tentacles. The tentacles bear each a terminal knob in adult specimens and
are hollow and not retractile. Mouth 4-sided, cruciform, with 4 perradial buttresses, between
which there are 4 interradial funnel-like pits inthe floor of the subumbrella extending down-
ward into the tissue of the 4 septa. The 4 gonads form each an interradial horse-shoe, the
outer points of which do not extend to the bell-margin.

According to Clark, 1863, the perradial stomach-pouches are each bridged across by a
cross-partition or claustrum extending over from the sides of adjacent gonads. Thus the gonads
are confined within the 4 axial chambers adjacent to the mouth and are separated by cross-
partitions from the outer parts of the perradial pouches.

Color is dirty chocolate-brown, the stalk paler.

This form grows permanently attached to rocks between tide-limits and does not reattach
itself if torn from its anchorage.

Northern coasts of Europe. It is generally rare, and is found only locally in such places as
the Firth of Clyde, Orkney Islands, near Bergen, Norway, Weymouth, England, etc.

Beaumont, 1894, Maas, 1906, and other recent observers have come to the conclusion that
Depastrella carduella Haeckel, 1880 (p. 376, taf. 21, fign. 5 to 12), is only the young or an
undeveloped stage of the Depastrum cyathiforme. Also Depastrella allmant, from Handa Island
and the Orkneys, and D. polare from Spitzbergen, described by Haeckel, 1880, p. 639, appear
to me to be identical with D. cyathiforme.

When young the tentacles are arranged in a single row around the margin and there is
but one tentacle in each perradius and interradius, but when older the perradial and inter-
radial tentacles become three times as many, and the adradial ones increase so as to be arranged
in several rows.

It appears to me to be fruitless to attempt to separate species upon the length of th
peduncle, its winged or unwinged (contracted or expanded) condition, as has been done by
Haeckel, and until more detailed studies of living specimens have proven the contrary to be the
case we had better venture to assume that all of the so-called Depastrellas and other forms of
the North Atlantic are synonymous with De pastrella cyathiforme.

Genus STENOSCYPHUS Kishinouye, 1902.

Stenoscyphus, Kisuinouye, 1902, Journal College Science, Tokyo, vol. 17, art. 7, p. 2, figs. 1, 2.
Stenoscyphus (?) Brocu, 1907, Hydroiden und Medusen, Report Second Arctic Exped. Fram, No. 12, p. 9.

The type species is Stenoscyphus inabat Kishinouye, of Japan, which has 8 marginal
anchors, 8 clusters of adradial tentacles, and a 4-chambered peduncle.

STAUROMEDUS4—STENOSCYPHUS., 5

GENERIC CHARACTERS.

Stauromeduse with simple, uncleft bell-margin without adradial lobes. With 8 (or 12?)
perradial and interradial marginal “anchors,” 8 (or 12 ?) adradial clusters of knobbed tentacles.
With a ring-shaped, entire, coronal muscle. Stalk of attachment is 4 (or 6?) chambered.
Stomach 4-chambered as in the Eleutherocarpida. Gonads interradial or on both sides of
the interradil.

Kishinouye would constitute for S. nabai a new family, the Stenoscyphidz, which would
be intermediate between Haeckel’s Tesseridz and Lucernaride. 8. inabar has a 4-chambered
stalk and 8 separate, adradial gonads.

Broch, 1907, describes a medusa which may belong to the genus Stenoscyphus but which
has 12 (6 perradial and 6 interradial) marginal anchors and 12 adradial clusters of tentacles.
The mouth is 6-rayed instead of 4-rayed, as in S. inabai. A variation in the number of rays
may be expected in these degenerate medusz and it seems inexpedient at present to sepa-
rate the 6-rayed from the 4-rayed form.

Stenoscyphus inabai Kishinouye.
Depastrum inabai, Kisu1nouye, 1893, Zool. Mag. Tokyo, vol. 5, p. 416.
Stenoscyphus inabai, Kisu1nouye, 1902, Journal College Science, Tokyo, vol. 17, art. 7, p. 2, plate 1, figs. 1, 2.

About 25 mm. long, elongate, narrow, funnel-shaped, and quadratic in cross-section.
Stalk with 4 interradial grooves. Exumbrella smooth; subumbrella beset with large, urn-
shaped groups of nematocysts. The 8 principal tentacles are large, round, adhesive bodies
(anchors). Secondary tentacles are short and knobbed, and arranged in 8 adradial clusters
of about 25 each. There are neither arms nor lobes to the subumbrella. The ring-like coronal
muscle is well developed and entire. 4 long, deep, interradial, gastrogenital pits extending
to the aboral end of the bell. 8 gonads, in form of 4 pairs of bands along the interradial
muscles. Each gonad composed of 40 sacs in 2 rows.

Color dark brown flecked with white, anchors red, gonads brown.
Subumbrella pale-green, manubrium yellowish.

Kata-ura, Kii Province, and Misaki, Japan. In winter. Rare.
Attached to Sargassum. The animal can detach itself from the Sar-
eit, gassum “at will’? and can crawl about, using its oral
str a, “~~ and aboral adhesive organs, thus effecting a locomo-
tion resembling that of a leech. It can not swim by
pulsations of the bell.

Stenoscyphus (?) hexaradiatus Broch.

Stenoscyphus (?) hexaradiatus, Brocu, 1907, Hydroiden und Medusen,
Report Second Norwegian Arctic Expedition in the Fram, No. 12,
p- 9, taf. 2, fign. 3-6.

Bell-shaped, with a cylindri-
cal peduncle somewhat longer
than height of bell. Bell about
6 mm. wide, total height of stalk
together with bell 1omm. Throat-
tube short, 6-sided. Peduncle pro-
ya vided with 6 longitudinal muscles.
Fic. 334.—Stenoscyphus inabia, after Kishinouye, in Jour. College of Science 129 marginal anchors, 6 radial and

Tokyo, Japan. 6 interradial. These are shaped
Fic. 335.—Stenoscyphus hexaradiatus, after Broch, in Report of the Second very muchas ordinary tentacles,

Norwegian Arctic Expedition in the “Fram.” ak Leas le adie Cee ae
like form,and each provided with anabaxial cushion. ‘Tentacle-clusters arranged in 12 adradial
groups, each composed of 7 to 10 tentacles which arise in several rows from the bell-margin.
Among each cluster of suctorial tentacles there are one or more tentacles which resemble the
marginal anchors. 6 horse-shoe-shaped, folded gonads with their convex sides abaxial. The
two wings of each horse-shoe do not extend to the circular muscle. Color (?)

526 MEDUS OF THE WORLD.

This medusa is distinguished by its 6-rayed structure, whereas in 8. inabai the peduncle
is 4-sided and there are 8 adradial tentacle-clusters anktead of 12 asin S. hexaradiatus. It is
possible, as Broch states, that S$. hexaradiatus may be the type of a new genus, but we must
await results of future studies before deciding this point. Unfortunately there is only one
specimen, obtained by the Fram off Fosheims Peak, Arctic Ocean, on the second expedition.

A closely allied form from the Kurile Islands is described by Kishinouye, 1909, under
the name Thaumatoscyphus distinctus. (See Appendix to this volume).

Genus LUCERNARIA O. F. Miiller, 1776.

Lucernaria, Miter, O.F., 1776, Prodromus Zoolog. Dan., p. 227.—Sars, 1846, Fauna littoral. Norveg., fasc. 1, p.20.—AGass1Z,
L., 2, Cont. Nat. Hist. U. S., vol. 4, p. 175.—Crark, H. J., 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 551-—
Acassiz, A., 1865, North age hee oes Jena. Zeit. fiir Naturwissen., Bd.7, p. 487.—Harcket, 1880, Syst.
der Medusen, p. 389; 1881, Deep-Sea Medusx, Challenger Report, Zool.,vol. 4, p. 53 Snore ELL, 1893, Natural Science, vol.
3, p- 208.—Hurst, Ibid., p. 209.

Lucernosa, Haecket, 1881, Ibid., p. 62.

Lucernaria, Kowavevsxy, 1884, Zool. Anzeiger, Bd. 7, p. 712.—Kasstanow, 1901, Zeit. fiir wissen. Zool., Bd. 69, p. 287, 11
fign., taf. 22-25.—Maas, 1906, Arktischen Medusen, Fauna Arctica, Bd. 4, Lfg. 3, p- 499-

Lucernaria+ Lucernosa, ANTIPA, 1892, Zoolog. Jahrb., Abth. Syst., Bd. 6, pp. 378, 391-

The type species of this genus is Lucernaria quadricornis from the northern Atlantic
coast of Europe, Greenland, and America.

GENERIC CHARACTERS.

Stauromeduse with 4 simple perradial stomach-pouches asin Eleutherocarpida. With-
out marginal anchors or marginal papilla. Peduncle single-chambered with 4 separate
teniola or partial septa. 8 adradial lobes which bear tentacles.

Antipa, 1892, would restrict Lucernarra to include forms with gonads of simple structure,
whereas “ Lucernosa” he would institute to define forms having complex gonads made up of
numerous more or less separated sacs set side by side in a series of follicular ridges. In this
respect he follows the suggestion of Haeckel, 1881, p. 62.

In all young medusx of Lucernaria the gonads are simple, flat, leaf-like expansions in the
entoderm of the subumbrella, and in many of the species they develop transverse ridges, and
these may become so marked as to appear as deep foldings across the gonad, thus dividing it into
a series of more or less sac-like corrugated pouches. Ghia’ is, however, a relative matter and can
not be safely used as a generic distinction; moreover, I think we should, if possible, avoid found-
ing genera upon details of internal anatomy requiring dissection to defeoniee their condition.

Kassianow, 1901, finds that in Lucernaria, Craterlophus, and Haliclystus there is a plexus
of bipolar ganglion cells in the ectoderm of the exumbrella. These ganglion cells have each
two nuclei. There is a motor center at the point of each arm, and the nervous epithelium
spreads out from the bases of the tentacles. The perradial and interradial anchors are not
motor centers as one would expect them to be were they derived from rhopalia.

The early development of Lucernaria has been studied by Kowalevsky, 1884 (Zool.
Anzeiger, Jahrg. 7, p - 712), who found that the eggs and sperm were discharged in the Bay
of Sebastopol near evening in August. The segmentation is total and equal, and the entoderm
is apparently formed by aeaninar A single, central, entoderm cell was seen, however,
with a slender prolongation extending from between the ectoderm cells, and thus it is possible
that the entoderm may originate from one of the ectoderm cells which withdraws into the
interior. The larva elongates, the entoderm becoming a single linear row of cells, and the
ectoderm becoming very thin at the ends. The ectoderm is not ciliated, but the larva creeps
about. On the fourth day the larve attach themselves and become flat and rounded, and
the entoderm forms a mass instead of a single layer, as previously. The larva then encyst
themselves in a hard, secreted covering, within which they remain for about two weeks; after
escaping they became lost to observation. Fol, 1873, found also that in Lucernaria the seg-
mentation is complete and results in the formation of a single-layered blastosphere, which

elongates, becomes ciliated and 2-layered. After this it becomes attached. 8 small, tentacle-
like, marginal bodies, 4 perradial and 4 interradial, appear, but soon degenerate and disappear.
Thus apparently Lucernaria is descended from a Haliclystus-like ancestor. Hornell, 1893,
states that 8 marginal anchors are found in the young medusa, but disappear in the adult.

STAUROMEDUS

Synopsis of the Spectes of Lucernaria.

LUCERNARIA,

L. kukenthali

L. quadricornis | L. pyramidalis L. walteri Antipa. L. haeckeli Antipa.
Miller. Haeckel. Antipa. |
Width of bell in mm.| 50 to 60 40 to 50 55 to 60 55 to 60 27
Height of bell without} 25 to 30 50+ Jo+ 80+ 45 to 50
peduncle in mm.
Length of peduncle in| 25 to 40 40 to 50 80+ Jot 15
mm.

Angular distance
between arms.

Number of tentacles
on each arm.

Form and position of
gonads.

Color.
Where found.

The 4 perradial

As in L. quadri-

notches twice as|_cornis. cornis.
wide and deep as

the interradial.

100 to 120 130 to 140 700 to 750

8 from beginning
of peduncle to
ends of arms.

arm.

Very variable. 2

North Atlantic
coasts of Europe
and America.

8 from base of
peduncle only to
crotch of each

Labrador coast.
Is this not L.
quadricornus
contracted
through preser-
vation in alcohol ?

of 8 arms.

Light brown.

As in L. quadri-

8 wide lancet-shaped'
extending to ends

East Spitzbergeno,
Arctic Ocean.

The 4 perradial

800 to 850

8 small, lancet-
shaped, cross-
folded bands
reaching to ends
8 arms.

>

Arctic Ocean.

notches 3 times as
wide and deep as
the 4 interradial.

East Spitzbergen,

The 4 perradial
notches only little
wider and deeper |
than the 4 inter-
radial

80 to go

8 very wide, over-
lapping; extending}
| not quite to bases
of| of 8 arms.

>

East Spitzbergen,
Arctic Ocean.

L. infundibulum

L. campanulata

L. bathyphila

L. australis

Haeckel. Lamouroux. Haeckel. Vanhoffen
(immature).
Width of bell in mm. 24 20 to 30 50 to 60 9
Height of bell without | 13 20 to 25 55 to 70 10
peduncle in mm.
Length of peduncle in | 7 10 to 15 5 to 10 Absent or undeveloped.

mm.

Angular distance be-
tween arms.

Number of tentacles on
each arm.

Form and position of
gonads.

Color.
Where found.

As in L. haeckeli.
60 to 80

From middle of pedun-
cle to bases of arms.

?

Spitzbergen, Arctic
Ocean.

45° apart. Similar each
to each.

30 to 40

From point of juncture
between bell and
peduncle to tips of 8
arms.

Very variable.

Black Sea and Medi-
terranean to English
coast. New Zea-
land (?)

As in L. kiikenthali.
80 to 120

Broad and short. Neither
reaching to base of
peduncle nor ends of
arms. Each gonad
has 200 sac-like fold-
ings.

?

Between Farée and
Shetland Islands,
North Atlantic. From
a depth of 540
fathoms.

As in L. quadricornis.
25 to 30

Not developed.

?

Gauss Station, Ant-
arctic Continent.
From depth of 192
fathoms. November 3,
1902.

Lucernaria quadricornis O. F. Miiller.

Lucernaria quadricornis, Miter, O. F., 1776, Prodrom. Zool. Dan., p. 227; 1788, Zool. Dan., vol. 1, p. 51, plate 39, figs. 1-6 —
Lamarck, 1816, Syst. Anim. sans Vert., tom. 2, p. 354.—Cuvier, 1817, Régne Animal, tom. 4, p. 53-—Sars, M., 1829,

Bidragtil Sodyrenes Natur., fasc. 1, p. 43, tab. 4, figs. 14-18; 1846, Fauna littoral. Norveg., fasc. 1, p. 20, plate

3» figs.

I-7.—Stimpson, 1853, Smithsonian Cont. to Knowledge, Marine Invert. Grand Manan, p. 8.—Carus, 1857, Icon.,
Zootom, taf. 4, fign. 1, 2—Crarx, H. J., 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 552, (good list of literature).—
Acassiz, A. 1865, North Amer. Acal., p. 62.—Haercket, 1880, Syst. der Medusen, p. 390.—KEFersTEIN, 1863, Zeit. fiir
wissen. Zool., Bd. 12, p. 20.—TascHENBERG, 1877, Halle, Zeit. ges. Naturw., Bd. 49, p. 82, taf. 2, fig. 2.

Disk about 50 to 60 mm. in diameter and, together with the peduncle, 50 to 70 mm. in
height. The bell is of the shape of a 4-sided funnel and about twice as wide as high. Peduncle

MEDUSA OF THE WORLD.

528

somewhat longer than the bell-height and with 4 longitudinal, interradial strands of muscle-

fibers. Bell-margin divided by 8 clefts or notches, the 4 perradial ones being about twice as

wide and deep as the 4 interradial; thus the 8 arms are brought quite close together in 4
Each arm bears 100 to 120 tentacles. The stomach gives rise to 4 wide,

separate pairs.
Found on the northern coasts of Europe, on the Greenland coast, and on the coast of

perradial pouches, which are lined on their edges by the 8 gonads.
Color variable, being either gray, green, yellow-brown, red-brown, or very dark brown.
It has never been taken south of Massachusetts Bay. Very

America, north of Cape Cod.
rare on American coast.
Complete descriptions and good figures of this medusa have been given by Sars, 1846;

Carus, 1857; and Taschenberg, 1877.

Lucernaria “‘pyramidalis” Haeckel =L. quadricornis (?)
Lucernaria pyramidalis, Harcxet, 1880, Syst. der Medusen, p. 391, taf. 22, 10 fign.

Bell 4-sided and pyramidal, about 40 to 50 mm. wide and (with the style) go to 100 mm.
high. The 8 arms grouped in 4 pairs, the 4 perradial notches of the bell-margin being twice

as wide and deep as the 4 interradial. Each arm bears a large cluster of 130 to 140 tentacles.

moo»

Mii

NRT

\Y

Lucernaria “ pyramidalis,” after Haeckel, in Das System der Medusen

Fic. 336.
Peduncle about 0.25 to 0.33 longer than bell-height. 4 short, blunt taniola at its free distal
end and a sharply marked pyloric stricture at its proximal end at base of bell. 4 linear, inter-
radial strands of muscle-fibers traverse the length of the peduncle. Mouth cruciform, with
4 short, folded lips. The gonads are 8 blindly ending, transversely folded bands, grouped
into 4 interradial pairs. They begin on the floor of the subumbrella at a short distance above
the pyloric stricture and extend only to the crotch of the bell-arms. They diverge centrif-

STAUROMEDUSA2—LUCERNARIA. 529

ugally outward, and the components of each pair are separated one from another by the
4 interradial septa. The gastric filaments in the regions of the gonads are small but very
numerous.

This form is found on the Labrador coast. A good description of it is given by Haeckel,
1880. It is certainly closely related to, if not identical with L. quadricornis, but Haeckel states
that the peduncle is separated from the bell by a pyloric stricture, or ring-furrow, which is
not the casein L. guadricornis. Haeckel studied only preserved specimens and I strongly sus-
pect that this so-called pyloric constriction may have been caused by unnatural contraction.

Lucernaria walteri Antipa.

Lucernosa walteri, ANtIPA, 1892, Zoolog. Jahrb., Abth. Syst., Bd. 6, p. 379, taf. 17, fign. 1-9.

150 to 160 mm. high and 55 to 60 mm. across the bell. Bell goblet-shaped; stalk round,
single-chambered, somewhat higher than the bell. Stalk with 4 well-developed, linear, inter-
radial,longitudinal muscles. 8 arms arranged in pairs with the 4 perradial concavities between
the arms twice as wide at the 4 interradial notches of the margin. Each arm with a terminal,
ball-like cluster of 700 to 750 short, knobbed tentacles. 8 wide, lancet-shaped, adradial
gonads, extending to ends of the 8 arms; they are folded, band-like, and lie in the subumbrella
wall of the perradial stomach-pouches. The cavity of the stalk extends directly into that of
the bell, without a pyloric stricture. Color light brown. East Spitzbergen, Arctic Ocean.
This is one of the largest known Lucernarians. Special description given by Antipa.

Lucernaria kiikenthali Antipa.

Lucernosa kiikenthali, ANtipa, 1892, Zoolog. Jahrb., Abth. Syst., Bd. 6, p. 386, taf. 18, fign. 10, 11.

More than 150 to 160 mm. high, 55 to 60 mm. wide across the bell. Bell goblet-shaped,
somewhat higher than wide. Stalk not quite as long as the bell itself, with no constriction
or other sharp distinction between stalk and bell. Stalk round, single-chambered, with 4
well-developed, linear, longitudinal muscles. 8 arms arranged in pairs. The 4 perradial
notches of the bell-margin are 3 times as wide and 3 times as deep as are the interradial. Each
arm has a terminal, ball-like cluster of 800 to 850 short, knobbed tentacles. Gonads are 8
small, lancet-shaped, cross-folded bands extending to ends of the 8 arms. Color (?) East
Spitzbergen, Arctic Ocean. Described in detail by Antipa, 1892. I am inclined to suspect
that this is only a variety of, if not identical with, Lucernaria walteri. It appears to be dis-
tinguished only by its narrow lanceolate gonads and slightly wider perradial notches.

Lucernaria haeckeli Antipa.
Lucernosa haeckeli, ANtira, 1892, Zoolog. Jahrb., Abth. Syst., Bd. 6, p. 388, taf. 18, fign. 12-14.

60 to 65 mm. high and 27 mm. wide at widest part, which is below bell-margin. Bell
oval, goblet-shaped, widest near middle. Stalk conical, wide, and hardly one-third as long
as bell. Stalk single-chambered with 4 interradial, longitudinal muscles. 8 short arms
arranged in pairs. The 4 perradial notches of the margin only a little wider and deeper than
the 4 interradial notches. Each arm terminates in a ball-like cluster of 80 to go knobbed
tentacles. 8 very wide gonads, so wide that they overlap one another. The gonads extend
not quite to bases of arms, and quite fill the lower parts of the radial chambers of bell.

This species is distinguished from L. bathyphila by its short bell-stalk and the position
and shape of its gonads. Color (?) East Spitzbergen, Arctic Ocean.

Lucernaria infundibulum Haeckel.

Lucernaria infundibulum, Harcxet, 1880, Syst. der Medusen, pp. 392, 385.

Bell funnel-shaped, flat, not quite twice as wide as high, 24 mm. wide. Peduncle some-
what less than 10 mm. long, 4-sided, pyramidal, and single-chambered. The medusa is
distinguished by having 4 hollow, interradial tamiole each with a funnel-like, central cavity
extending to lower end of peduncle, recalling the condition seen in the aboral end of bell in
Periphylla. Each hollow septum bears along the entire length of its side walls a pair of well-
developed longitudinal muscles and 2 rows of gastric filaments. The 8 umbrella-arms are
arranged in 4 pairs, the 4 perradial notches being wider than the 4 interradial. Each arm has

53k MEDUS® OF THE WORLD.

So tentacles. 8 gonads extending from middle of peduncle to base of arm. Found at
Described by Haeckel from a preserved specimen.

Lucernaria campanulata Lamouroux.

Lucernaria campanulata, Lamouroux, 1815, Mém. du Museum Hist. Nat., tome 2, p. 472, planche 16, figs. 1-7.—K EFERSTEIN,
1862, Zeit. fiir wissen. Zool., Bd. 12, p. 23, taf. 1, fig. 4 —Harcxet, 1880, Syst. der Medusen, p. 392 (list of literature).—
Houtron, 1880, Trans. New Zealand Institute, vol. 12, p. 275.—Ctaus, 1883, Untersuch. ber Organisation und Entwick.
ler Medusen, p. 35.—GrAxrFFE, 1884, Arbeit. Zool. Inst. Wien., Bd. 5, p- 344-—Hornext, 1893, Nat. Sci., London, vol. 3,

». 208.—Braumont, 1900, Proc. Roy. Irish Acad. Dublin, Ser. 3, vol. 5, p. 811.—Kassianow, 1901, Zeit. fiir wissen. Zool.,

Bd. 69, p. 370, fig. 11
ria convoloulus, JouNSTON, 1835, London, Mag. Nat. Hist., vol. 8, p. 59, fig. 3-
ria, Kowarevsxy, 1884, Zool. Anzeiger, Jahrg. 7, p- 712 (embryology).

This medusa is at once recognized by its symmetrically octagonal disk with its 8 arms
45° apart and with equally developed notches between them. There are no longitudinal
muscles in the 4 interradial longitudinal ridges of the peduncle. The medusa is 20 to 30 mm.
wide and 30 to 40 mm. high, including the peduncle. Color very variable, being yellowish,
red, brownish, ete. Found along European coasts from the Black Sea and Mediterranean
‘o southern England, Ireland, and Wales. Graeffe states that it is found only locally at Trieste,
Adriatic Sea, in May and June, becoming mature at the end of the latter month. Hutton
records it from Brighton near Dunedin, New Zealand, but he gives no description.

For an account of Kowalevsky’s observations upon the early stages of the larva see
genus Lucernarta.

Hornell, 1893 (Nat. Sci., vol. 3, p. 208), states that 8 marginal anchors are found in the
young medusa, but they soon disappear and are not found in the adult. This leads one to con-
clude that Lucernartais derived directly
from Haliclystus. Indeed Horst, 1893,
finds that the variations in number and
development of the marginal anchors is
so great that he is inclined to consider
Lucernaria to be identical with Hal-
iclystus. Kassianow, 1901, records the
capture of an abnormal specimen having
2 medusa bells arising from one stalk.

Lucernaria bathyphila Haeckel.

Lucernaria bathyphila, Harcxer, 1880, Syst. der
Medusen, p. 640; 1881, Report Deep-Sea
Meduse Challenger Expedition, Zool., vol. 4,
p- 54, plates 16, 17, 21 figs.

Lucernosa bathyphila, ANtira, 1892, Zoolog. Jahrb.,
Abth. Syst., Bd. 6, p. 379-

Fic. 337-—Lucernaria bathyphila, after Haeckel, in
Deep-sea Meduse Challenger Expedition.

For description, see synopsis of
the species of Lucernaria. Haeckel,
1881, gives a very detailed and fully illustrated description of this species.

Lucernaria australis Vanhoffen.
Lucernaria australis, VANuOrrEN, 1908, Deutsche Siidpolar-Exped., 1901 to 1903, Bd. 10, Zool. 2, p. 32, fign. 1, 2.

Described by Vanhéffen from an immature specimen which had neither gonads nor
peduncle.

Bell 10 mm. high, 9 mm. wide, thimble-shaped, with sloping sides. No basal stalk, but
with a weakly developed ring-furrow at aboral end of bell, and an indication of the beginning
of a single-chambered peduncle. 8 short arms, 2 to 2.5 mm. long, grouped in pairs somewhat
closer in the interradii than in the perradii, and with the perradial concavities of margin
deeper than the interradial. 25 to 30 short tentacles, with small terminal knobs on each arm.
There were 7 very small, tentacle-like marginal bodies somewhat asymmetrically placed
near the 4 perradial and 3 of the interradial points of the bell-margin. It is therefore possible
that this medusa may be a young Haliclystus, but if the marginal bodies degenerate it is a
Lucernaria, for such a course of development is known according to Hornell, 1893, in the
European species of Lucernarta. The 4 perradial lips of the cruciform mouth are folded and
are at about half the distance between the depth of bell-cavity and margin.

STAUROMEDUSA—KISHINOUYEA, HALICLYSTUS. 531

Wide ring muscle at margin of subumbrella and 8 strands of radial-muscles extending
outward to the tentacles. These radial strands are one-third wider than the ring muscle,
They inclose 4 narrow, triangular areas in the interradii and 4 wider, rectangular spaces in
the perradii. These subumbrella areas between the muscle strands exhibit many large nettle-
cells. There are about 12 simple, unbranched, gastric filaments in each of the 8 rows. Found
at Gauss Station, Kaiser Wilhelm II Land, Antarctic Continent, at a depth of 192 fathoms,
in November, 1902.

Genus KISHINOUYEA, nom. nov.
Schizodiscus, preoccupied by Kittl, 1891.
Schizodiscus, KisH1Nouye, 1902, Jour. College Sci., Tokyo, vol. 17, art. 7, p. 5.
The type species is K. nagatensis of Japan, first described by Oka under the name Lucer-
naria nagatensts.

GENERIC CHARACTERS.

Stauromedusz with 4-chambered stomach as in the Eleutherocarpide, and without adhe-
sive anchors. Umbrella deeply notched, with 8 adradial lobes. 8 adradial clusters of
knobbed tentacles. Peduncle 4-chambered without muscle-fbers in the taniola. Gonads
8 adradial bands of laterally oblong sacs.

This genus is very closely allied to Lucernarra and is distinguished only by its 4-chambered
aboral stalk. In young medusz, however, it is single-chambered, as in Lucernaria, but the
4 interradial septa, or teeniola, unite near the pyloric region as growth proceeds, and thus the
peduncle comes to have 4 perradial, separate chambers.

Kishinouyea nagatensis.

Lucernaria nagatensis, Oxa, 1897, Zool. Mag. Tokyo, vol. 9, p. 67, plate 1. Also; Annot. Zool. Jappon. Tokyo, vol. 1, p. 141,
ee ae KisHiNovuye, 1902, Journ. Science College, Tokyo, vol. 17, art. 7, p. 6, plate 1, figs. 3-6.

The 8 adradial lobes are united in pairs, so that the 4 perradial notches are about twice
as deep as the 4 interradial. These
8 adradial lobes are bent at right
angles to the oral side. The disk
has the shape of a Greek cross.
Peduncle 4-chambered in adult.
Well-developed, interradial, longi-
tudinal muscles in the subumbrella.
Marginal muscle divided into 8 U-
shaped pieces. No primary tenta-
cles. Adradial tentacles short, adhe-

Fic. 338.—Kishinouyea nagatensis, after Kishinouye, sive, in clusters of GS: Gastral fila-

in Jour. Col. of Science, Tokyo. ie , : a

ments branched, few in number.

Gonads, 8 broad, adradial bands of laterally oblong sacs. Color variable, matching its
surroundings. Japan.

Genus HALICLYSTUS Clark, 1863.

Haliclystus, CLtarx, 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 559; 1878, Smithsonian Contributions to Knowledge, Article
2, Part. 2—Acassiz, A., 1865, North Amer. Acal., p. 63.—Harcker, 1880, Syst. der Medusen, p. 387.—Kisninouye,
188g, Proc. U.S. National Museum, vol. 22, pp. 125, 129.—Maas, 1904, Résult Camp. Sci. Prince de Monaco, fasc. 28,
P- 43; 1906, Fauna Arctica, Bd. 4, Lfg. 3, p. 499—Gross, 1900, Jena. Zeit. fiir Naturwissen, Bd. 33, p. 614 (anatomy).
Lucernaria, Bercu, 1888, Vidensk. Meddel. Naturhist. For. Kjébenhavn, p. 214 (embryology).

The type species of this genus is H. auricula from the Atlantic coasts of Europe and of
New England, United States.

GENERIC CHARACTERS.

Stauromedusze similar to Lucernaria but with 8 perradial and interradial marginal
anchors, and with a 4-chambered, aboral peduncle. The embryology has been studied by
Bergh, 1888, and is similar to that of Lucernaria, excepting that in Lucernaria the 8 anchors
become lost as growth proceeds.

bo

MEDUS OF THE WORLD.

The regeneration of Haliclystus has been studied by A. Meyer, 1865 (40 ste Versamml.
deutsch. Naturforscher Aerzte, Hannover, p. 217), who gives a brief account of his experiments.
It appears that the medusa possesses considerable regenerative capacity. The style is regen-
erated if it be removed, and if only the lower end of the style be cut off the medusa usually
regenerates a new aboral end; but occasionally a bell is regenerated, thus giving an animal

with 2 bells.

Characteristics of the So-called Spectes of Haltclystus.

Somewhat similar results were attained by Kassianow, 1901, on Craterlophus.

H. octoradiatus. | H. salpinx. | H. stejnegeri. H. auricula. H. antarcticus. H. kerguelensis.
Umbrella. | Conical, surface | Pyramidal, Conical, surface | Pyramidal, Flat, twice as wide) As in H. antarc-
flat, 2 to 3 times | octangular, flat, a little octangular, as high, conical.| ticus.
as broad ashigh.| much broader | broader than almost as broad
than high. high. as high.
| Peduncle. | Cylindrical, al- Quadrangular, | About half aslong} Almost aslong as | 4-sided, prismatic, Twice as high as
most as long as | _ prismatic, as height of um-| height of um- half to two- | umbrella.
height of um- considerably brella, with 4 brella, with 4 thirds aslong as | Prismatic,
brella; no inter-| longer than interradial longi-| deep longitudi- | height of um- 4-sided.

Eight arms.

radial longitudi- |
nal grooves.

45° apart. Each
arm with 30 to 60)
tentacles.

height of um-
brella.

45° apart.

Each arm with |

60 to 70 tenta-
cles.

tudinal grooves.

45° apart. Each
arm with 70 to
100 tentacles.

nal grooves.

United in pairs.
Each arm with
100 to 120 tenta-
cles. Interradial
clefts only half

brella. With 4
longitudinal
grooves (con-
tracted ?).

45° apart and
similar each to
each. Each arm
with, more than
100 tentacles.

As in H. ant-
arcticus, but
with not more
than so tenta-
cles on each

or two-thirds as arm.
wide as perradial,

Eight Large, egg-shaped) Vasey large, as | Large,egg-shaped, Large, shaped like) Large, biscuit- Small, oval, only
marginal or nearly club- icug as arms,| half as long as | coffee-beans, shaped, about as} one-third as
anchors. shaped, one- obliquely breadth of as long as long as width of | wide as width

fourth as long as| trumpet- peduncle. breadth of peduncle. of peduncle.
breadth of shaped. peduncle.
peduncle.
Gonads. In each gonad 20) In each gonad | In each gonad 100) In each gonad 100) 8 gonads, widely | 8 widely
to 30 large sacs | 40 to 50 sacs | to15osacs,6to| to 150 sacs in 6 | separated one separated,
in2longitudinal,| in 4 longitudi- | 8 sacs abreast in| to 8 radial rows. | from another, broad, lancet-
alternate rows. nal rows. the broadest 100 to 150 sacs | shaped gonads.
part. in 6 to 8 radial

rows in each

gonad.

Where North Atlantic. Coast of Maine,| Bering Island, North Atlantic. Island of South | Kerguelen
found. United States. | Commander Coasts of Europe} Georgia, Ant- Island, Ant-

Islands, North and America. arctic Ocean. arctic Ocean
Pacific.

Haliclystus auricula Clark.

Lucernaria auricula, Ratuxr, 1806, Miiller’s Zool. Dan., Bd. 4, p. 35 (exclus. synon).
Haliclystus auricula, CLark, H. J., 1863, Jour. Boston Soc. Nat. Hist., vol. 7,p. §59-—Acassiz, A., 1865, North Amer.Acal., p.63,
figs. 88-go.—Ciark, S. F., 1876, Exploration of Alaska, vol. 1, p. 235.—Ctark, H. J., 1878, Smithsonian Contributions
to Knowledge, pp. 1-130, plates 1-9, 145 figs.—HarckeEL, 1879, Syst. der Medusen, p. 389.—Scutater, 1891, Zeit. wiss.
Zool., Bd. 52, p. 580; Revue Sci. Nat. St. Pétersbourg, Anné. 2, p. 176 (sense-organs, structure of).—Braumonr, 1893,
Trans. Liverpool Biol. Soc., vol. 7, p. 259; 1900, Proc. Roy. Irish Acad. Dublin, ser. 3, vol. 5, p. 806—Bicrtow, H.
B., 1909, Proc. U. S. National Museum, Washington, vol. 37, p. 316.
(?) Haliclystus tenuis, Kisuinovye, 1910, Journal College Sci., Tokyo, vol. 27, art. 9, p. 4, plate 1, fig. 3.

Disk about 20 to 30 mm. wide and (with the style) 20 to 30 mm. high. Bell-margin

8-sided. The 4 interradial notches between the arms being only half to two-thirds as wide
as the 4 perradial clefts. Each arm terminates in a large cluster of 100 to 120 tentacles. These
tentacles are each about quarter as long as bell-diameter. They are hollow and terminate in
a globular tip thickly covered with nematocysts. There are 8 large, perradial and interradial,
marginal anchors (colletocystophores), which are coffee-bean-shaped and mounted, each one,
upon a short cylindrical base. These organs are much larger than in the European H. octo-
radiatus. The aboral peduncle, or stalk, is about as long as bell-height. It is cruciform in
cross-section, there being 4 deep, interradial, longitudinal furrows beneath which are 4 well-

STAUROMEDUSAS—HALICLYSTUS.

developed strands of longitudinal muscle-fibers. Alternating with these are 4 perradially
situated, longitudinal chambers which communicate with the central stomach in the umbrella.
There are 8 broad, adradial gonads, grouped into 4 interradial pairs. Each gonad is wide
and triangular. These gonads begin a little above the point of junction of the stalk with the
disk and extend to the ends of the 8 arms. Each gonad contains 100 to 150 out-folded sacs

arranged in 6 to 8 radiating, longitudinal rows. Gastric cirri numerous.

(>

Oe,

aaca

XY

8,
>

Fic. 339-—Haliclystus auricula, after Clark, in Smithsonian Contributions to Knowledge. The smaller

figures are of natural size and illustrate characteristic attitudes.

Color very variable, but each medusa is commonly of one color, parti-colored individuals
being very rare. Individuals are either blue, green, yellow, olive, orange, or very rarely red,

pink, or violet. Medusz of brownish and purple hues are also common.

MEDUS OF THE WORLD.

~

This species is found in Massachusetts Bay and off the northern coasts of Europe from
England to Norway. Clark reports it from Norton Sound, Alaska, and Bigelow from Labra-
dor and Newfoundland. It is only locally common on the New England coast.

It may be distinguished from H. octoradzatus by its more slender bell and stalk, its large
marginal anchors, and the greater number and smaller size of the genital sacs upon the gonads.
Kishinouye, 1910, records a form from Japan which resembles H. auricula excepting that

each gonad consists of only two rows of saccules. He calls this medusa H. tenurs.

Haliclystus octoradiatus Clark.

Lucernaria auricula, Montacu, 1808, Trans. Linn. Soc. London, vol. 9, p. 113, plate 7, fig. 5.—Sars, M., 1829, Bidragtil
Sédyr. Naturhist., p. 34, taf. 4, fign. 1 to 13.—JounsTon, 1838, Hist. British Zoophytes, p. 229, fig. 35.—AGassiz, 1862,
Cont. Nat. Hist. U. S., vol. 4, p. 176.

Lucernaria octoradiata, LAMARCK, 1816, Hist. anim. sans vert., tom. 2, p. 474.—STEENSTRUP, 1859, Vidensk. Meddel. Nat. Foren.
Kjoébenhavn, p. 108.—Sars, M., 1860, Forhandl. Vid. Selsk. Christiania, p. 145.—Kerersrein, 1863, Zeit. wissen. Zool.,
Bd. 12, p. 22, taf. 1, fign. 1-3.—Tascnenpere, 1877, Zeit. Ges. Naturw., Halle, p. 91, taf. 2, fig. 4.—Bercu, 1888,
Vidensk. Meddel. Nat. For. Kj6benhavn, p. 214, fign. 1-3.

Haliclystus octoradiatus, Ctarx, H. J., 1863, Jour. Boston Soc. Nat. Hist., vol. 7, p. §65.—Harcket, 1880, Syst. der Medusen,
p- 388.—Levinsen, 1893, Vid. Meddel. Nat. Foren. Kjébenhayn, (5), Bd. 4, p. 146—Browne, 1896, Quart. Journ. Mic.
Sci., vol. 38, part. 1, pp. 1-8, plate 1, 22 figs Hornet, 1893, Natural Science, London, vol. 3, p. 33 (abnormalities) —

Browne, "1895, Quart. Journ. aeaae Sci., ser. 2, vol. 38, p. 1, plate 1 (variations)—Gross, 1900, Jena. Zeit. Naturw.,

Bd. 33, p- 611, taf. 23, 24 (anatomical ).—Braumont, 1900, Proc. Roy. Irish Acad., Dublin, ser. 3, vol. 5, p. 808.—Maas,

1904, Résult. Camp. Sci. Prince de Monaco, fasc. 28, p. 44—WietTrzyKowskl, 1909, Comptes Rendus, Paris, tome 149,

p- 746.

Disk 20 to 30 mm. wide and, with the stalk, 20 to 30 mm. high. Disk flat and about
2 to 3 times as broad as high. The 8 adradial arms are 45° apart, not grouped in 4 more
or less approximated pairs, as in H. auricula. Arms very wide, concavities of bell-margin
shallow. Each arm bears a terminal cluster of 30 to 60 tentacles, instead of 100 or more,
as in H. auricula. The 8 perradial and interradial marginal anchors (colletocystophores) are
egg-shaped, and about one-fourth as long as diameter of stalk. Stalk cylindrical, without
longitudinal furrows, 4-chambered, and with 4 interradial, longitudinal strands of muscle-
fibers. There are 8 separate gonads which do not extend quite to the end of the arms or to the
aboral septa. Each gonad contains only 2 rows of alternately arranged, large, genital sacs.
Color quite variable, being either grayish-yellow, brownish-yellow, or grayish-brown.
Found on the North Atlantic coasts of Europe, on the Greenland coast, and at Spitzbergen.
The most complete descriptions of this species are given by Sars, 1829; Keferstein, 1863;
Browne, 1895; and Gross, 1g00. Its variations have been studied by Hornell, 1893, and
by Browne, 1896. The abnormal forms are very irregular, symmetrical variations rarely
appearing. The medusa is one of the most variable known. At Jersey, England, according
to Hornell, 66 per cent of the specimens were abnormal in some respect; but at Plymouth,
according to Browne, only 34 per cent were abnormal and the aberrations were quite different
from those found at Jersey. We are unable to determine whether this difference is fostered by
isolation or is due to the effect of local influences in the environments of Plymouth and Jersey.
It may also be due to a difference in variative tendency in the medusz of the two places.
The development of H. octoradiatus has been studied by Bergh, 1888. The egg is fertilized
after being discharged into the water and then it retracts somewhat from the vitelline mem-
brane. 2 polar bodies are found; the segmentation is total and equal, and there is no cleavage
cavity. The entoderm appears to be formed by polar ingression of cells into the center of the
solid morula which is at first spherical but afterwards it elongates into a rod-like form, which
becomes so long and narrow that the entodermal cells come to be arranged one after another
in a single row as in the planula of Solmundella. The planula of H. octoradrata is not ciliated,
however, but creeps about by means of worm-like movements. It then attaches itself by, the
anterior a as do other planule of Scyphomedusz. At first the tentacles are not united into
definite clusters but are distributed around the bell-margin, but 8 tentacles are more or less
isolated and lie in the perradial and interradial radii. These form the marginal anchors.
The best description of the development of the planula is given by Wietrzykowski, 1909
(see Appendix to this Volume).
Bergh, 1888, describes an abnormal specimen of H. octoradiatus with a small bud
arising from the side of its bell.

STAUROMEDUS—HALICLYSTUS. 535

Haliclystus salpinx Clark.
Plate 56, figs. 1 to 4.
Haliclystus salpinx, Crarx, H. J., 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 563.—Acassiz, A., 1865, North Amer. Acal.,

p- 64.—HaeckeL, 1879, Syst. der Medusen, p. 388.

(?) Lucernaria salpinx, Grarrre, 1884, Arbeit. Zool. Inst. Wien., Bd. 5, p. 344-

Disk 25 mm. wide and (with the stalk) 20 mm. high. The 8 adradial arms are 45° apart,
and their ends are rounded. Each arm bears 60 to 70 very slender tentacles, the globose tips
of which are smaller than in H. auricula. The 8 marginal anchors are as long as the arms,
slender, and obliquely trumpet-shaped. The edge of the trumpet is considerably thickened
except at a narrow space on the proximal side, by the development of adhesive cells. The
center of this terminal expansion is occupied by a single tentacular remnant about as long as
half the breadth of the trumpet. Aboral stalk relatively longer and more slender than in /7.
aurtcula, 4-sided in cross-section, with 4 longitudinal rows of interradial muscle-fibers. The
stalk is 4-chambered and about 12 mm. long. The 8 genital organs are not so widely separated
as in H. auricula; their broader ends project only about half-way into the arms. The genital
sacs of each gonad are arranged in 4 radiating rows, the marginal rows being shorter than
the 2 middle rows. All of these sacs are of the same size and there are 40 to 45 of them in
each row.

This species was obtained by Stimpson at Mount Desert Island, Maine. The best
description of it is that of Clark, 1863. Our figures are drawn from specimens obtained upon
eelgrass on the inner side of Ram Island near Manchester, Massachusetts, on September 7,
1905. Graeffe, 1884, appears to have found this species at Trieste, Adriatic Sea, in June and
July.

Haliclystus stejnegeri Kishinouye.

Haliclystus stejnegeri, Kisuinouye, 1899, Proc. U. S. National Museum, vol. 22, p. 126, figs. 1-3.

Bell conical, funnel-shaped, 1.33 to 1.5 times as broad as high. 18 mm. wide. Peduncle
nearly quadrate in cross-section and about half as long as umbrella; its 4 interradial, longi-
tudinal grooves are formed by the attachment of the taniola. These septa meet at the longi-
tudinal axis and divide the internal space of the peduncle into 4 perradial chambers which
are continuous with the 4 perradial stomach-pouches. The surface of the exumbrella is smooth
and the line of demarcation between the stalk and the umbrella is distinct, although there is
no constriction at this point. There are a few small clusters of nematocysts at the radial
sinuses of the umbrella margin. The radial muscle plates are, as in other species of Hali-
clystus, developed in the perradii and interradii of the subumbrella. The margin of the
umbrella displays 8 equally spaced, adradial arms, all of the same size. The 8 incisions are
about as deep as the width of the arms themselves.

There are 8 large, egg-shaped, perradial and interradial “anchors,” which are about
half as long as diameter of peduncle; these are situated in the concavities of the clefts, alter-
nating with the tips of the 8 adradial arm-lobes. There are 8 adradial clusters of knobbed
tentacles, one at the end of each of the 8 arms. Each cluster contains 70 to roo tentacles of
various sizes.

Manubrium short and quadrangular, the lips reflected outward. The 8 rows of well-
developed gastric filaments extend from the base of the throat-tube to the proximal ends of
the 8 gonads. The 8 gonads are broad, leaf-shaped, tapering at both ends, and touch each
other along their proximal halves, so that the surface of the subumbrella is almost entirely
occupied by them. There are 100 to 150 round sacs in each gonad; these sacs are not arranged
in rows and those nearest to the 4 principal radii are the largest. There are 6 to 8 sacs abreast
at the broadest part of each gonad. Each gonad is turned over in the 4 principal radii and is
continuous with the mesentery.

Preserved specimens are grayish or pale brown, semi-transparent, with a dark-brown
or nearly black streak at bell-margin.

A number of specimens were found at Bering Island, one of the Commander Islands,
North Pacific, in summer.

The species is well described and figured by Kishinouye (see text-figure 340).

536 MEDUS OF THE WORLD.

Haliclystus antarcticus Pfeffer.

Haliclystus antarcticus, Prerrer, 1889, Mittheil. Naturhist. Museum Hamburg, Jahrg. 6, p. 16.

Bell flat, 17 mm. wide to bases and 27.5 mm. wide to ends of arms. Height 11.5 mm. to
subumbrella disk and 15 mm. to ends of arms. The stalk is 8 mm. long, flexible, and when
expanded it is about two-thirds as long as bell-height. It has 4 interradial, longitudinal
muscles and is 4-sided in cross-section, the longitudinal muscles being in the 4 flat or grooved
sides. The attached end of the stalk is swollen. There are 8 adradial arms 45° apart with
the 8 clefts all of equal depth, and each arm has more than roo tentacles; 8 large biscuit-
shaped, marginal anchors about as long as width of stalk; 8 wide, lancet-shaped gonads,
widely separated one from another and extending to ends of arms. The number of sacs in
the gonads i is not clearly defined in Pfeffer’s specimens, but there appear to be at least 100 to
150 in 6 to 8 longitudinal rows. Stalk single-chambered thus illustrating the close relation-
ship between Haliclystus and Lucernaria.

The medusa is a beautiful blue-violet in color, with lighter, somewhat reddish anchors
and tentacles. Found at South Georgia, Antarctic Ocean.

Haliclystus kerguelensis Vanhéffen.

Haliclystus kerguelensis, VANHOFFEN, 1908, Deutsche Siidpolar-Expedition, 1901-1903, Bd. 10, Zool. 2, p. 31, taf. 2, fig. 1.

Bell 27 mm. wide across the outstretched arms exclusive of the tentacles, and 10 mm.
high. Arms 45° apart, the concavities between them all similar each to each as in H. antarc-
ticus. Peduncle prismatic, 20 mm. long and 3 mm. wide. Not quite 50 tentacles in each
adradial cluster. Marginal anchors only one-third as wide as the peduncle in its expanded
state. 8 wide, lancet-shaped gonads.

Fic. 340.—Haliclystus stejnegeri, after Kishinouye, in Proc. U. S. Nat. Mus.
Fic. 341.—Haliclystus kerguelensis, after Vanhoffen in Deutsch. Sudpolar Expedition.

Bell and peduncle sandy-brown with a play of green over the surface. Gonads dark
olive-brown and plainly visible through the walls of the lighter colored bell. Terminal knobs
of the tentacles rose-red.

Found at Observatory Bay, Kerguelen Island, Antarctic Ocean; in July, growing on
the stems of Macrocystis.

This form is closely related to H. antarcticus, but has fewer tentacles, smaller anchors,
and apparently a longer peduncle, although the stalk of H. antarcticus was probably con-
tracted in Pfeffer’s preserved specimens. There are also color differences between the two
forms.

Genus HALIMOCYATHUS Clark, 1863.

Halimocyathus, Crarx, 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 536.—Acassiz, A., 1865, North Amer. Acal., p. 61.
Halicyathus, Hacker, 1880, Syst. der Medusen, p. 393-
Manania, Crark, 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 541.

The type species of this genus is H. platypus Clark, from Massachusetts Bay.

STAUROMEDUSA—HALIMOCYATHUS. 537

GENERIC CHARACTERS.

Stauromedusz with 4 perradial, gastrogenital pockets in the subumbrella wall of the 4
stomach-pouches, as in Cleistocarpida. With 8 marginal anchors (4 perradial and 4 inter-
radial). 8 adradial clusters of terminally knobbed tentacles. 4 interradial, horse-shoe-shaped
gonads. 8 adradial arms.

This genus is very closely related to Haliclystus, but is distinguished by the partitions
across its 4 perradial stomach- -pouches, such as are found in all Cleistocarpida.

Halimocyathus platypus Clark.

Halimocyathus platypus, Crarx, H. J., 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 537.—Acassiz, A., 1865, North Amer.
Acal., p. 61.—Haercket, 1880, Syst. der Medusen, p. 393.
Lucernaria platypus, TASCHENBERG, 1877, Zeit. Ges. Naturw., Halle, p. 92.

Disk deep funnel-shaped, about 6 mm. wide, and (with the peduncle) 10 mm. in height.
The 8 arms are nearly twice as long as broad and one-third as long as bell-height from peduncle
to margin. Each arm bears 17 to 20 thick, pistilliform tentacles, about as long as greatest
breadth of arms; the tentacles upon each arm are arranged in 5 rows, there being about 7
tentacles in the middle row, 4 in each row on either side of the latter, a one in bach of the
outermost positions. The 8 marginal anchors are small and reverted, being only one-third as
long as shortest tentacles, but proportionally broader; their length is a little less than 3 times
their radial diameter. The peduncle 1 is about half as high as the disk; at the narrowest part,
where it joins the disk, its diameter is nearly half its length, and from there it broadens into a
wide base having a width equal to the length of the peduncle; it is round, or very slightly
furrowed at 4 points opposite the 4 interradial muscles. The 4 separate, longitudinal chambers
of the peduncle are very voluminous and close together, but the longitudinal septa between
them are complete. There are 4 horse-shoe-shaped gonads; the centripetal parts of each
horse-shoe are united across the inner ends of the 4 interradial parts while the free, outer parts
extend to the neighborhood of the marginal anchors. Each arm of the horse-shoe contains
15 to 17 genital sacs. The edges of adjacent gonads are joined by a cross partition as in
other Cleistocarpida.

A single specimen of this species was found by Clark at Chelsea Beach, Massachusetts,
where it was found attached to Zostera, along with H. auricula. It has not been seen since
Clark’s day, and indeed the contamination of the sea-water in this region has destroyed the
Stauromeduse which once abounded there, and which are now exceedingly rare along the entire
New England coast.

Halimocyathus lagena Haeckel.

Holothuria lagenam referens, Miitrer, O. F., 1776, Prodromus Zool. Danica, p. 232.

Lucernaria auricula, Fawricius, 1780, Fauna Grenlandica, p. 341.—Mitne-Epwarps, 1849, Cuvier, Régne Animal. Zoophytes,
planche 63, 9 figs—Sars, M., 1860, Forhandl. Vid. Selsk., Christiania, p. 145.—KerersTeIn, 1863, Zeit. fiir wissen.
Zool., Bd. 12, p. 21.—TascHENBERG, 1877, Zeit. Ges. Naturw., Halle, p. 89.

Lucernaria fabricii, AGassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 176.

Lucernaria typica, GREENE, 1858, Nat. Hist. Review, p. 132.

Manania auricula, Ciarx, H. J., 1863, Journ. Boston Soc. Nat. Hist., vol. 7, p. 542.—Acassiz, A., 1865, North Amer. Acal.,
p- 62.

Halicyathus lagena, Harcxet, 1880, Syst. der Medusen, p. 394.—Levinsen, 1893, Vid. Meddel. Nat. For. Kjébenhavn (5),
Bd. 4, p. 147.

Bell urn-shaped, much deeper than broad. It passes from a rounded base abruptly into
the peduncle. Bell about 5 to 7 mm. wide, and (including the style) about 20 to 30 mm. long.
A cluster of 60 to 70 slender tentacles upon the end of each of the 8 arms. Arms about as
long as broad and grouped into 4 interradial pairs. 8 marginal anchors have the same form
as the knobbed tentacles, but are somewhat smaller. 4 horse-shoe- shaped gonads, the distal
ends of which extend radially outwards are separated from the bell-margin by a wide space.
Each horn of the gonads exhibits 12 to 14 genital sacs.

Color black or dark brown, rarely reddish: brown or yellowish-brown.

Found upon the northern Atlantic coasts of Europe and upon the coast of Greenland.
It occurs on the New England coast north of Cape Cod, but is very rare.

538 MEDUS OF THE WORLD.

(?) Genus CRATERLOPHUS Clark, 1863.

Crarx, 1863, Journal Boston Soc. Nat. Hist., vol. 7, p. 539.—HaeckeL, 1880, Syst. der Medusen, p. 394.—Gross,
Jena Zeit. fiir Naturwissen, Bd. 33, p. 614. —Maas, 1906, Fauna Arctica, Arktischen Medusen, Bd. 4, Lfg.3, p. 500.

1g0o,

The type species is Craterlophus tethys of Helgoland, German Ocean.

GENERIC CHARACTERS.

Stauromedusz with 8 adradial lobes and with 4 perradial gastrogenital pouches in the
subumbrella wall of the 4 perradial stomach-pouches as in the Cleistocarpida. Without per-
radial or interradial marginal anchors or marginal papilla. ‘The peduncle is 4-chambered.

According to Antipa, and Gross, this medusa may sometimes have 8 small tentacles, 4 per-
radial and 4 interradial, in the places of the anchors of other Stauromeduse. It is probable,
therefore, that Craterlophus is actually identical with Halimocyathus.

Craterlophus tethys Clark.

Lucernaria sp., Merrenurimer, 1854, Abhandl. Senckenberg, Naturf. Ges. Frankfurt, p. 15, taf. 1, fign. 5-11.

Craterlophus tethys, CLarx, H. J., 1863, Journal Boston Soc. Nat. Hist., vol. 7, p. 540-—Kuine, 1879, Morpholog. Jahrb., Bd. 5,
p. 141, taf. g-11.—Herrwic, 1879, Jena. Zeitschr. fiir Naturw., Bd. 13, p. 613, taf. 9, fign. 7-12.—HaeckEL, 1880, Syst. der
Medusen, p. 395.—Craus, 1883, Untersuch, tiber Organisation und Entwick. Medusen, p. 35.—AntTiPa, 1892, Zool.
Jahrb., Abth. Syst., Bd. 6, p. 392 (aboral educa —Gross, 1900, Jena. Zeit. fiir Naturw., Bd. 33, p. 614, taf. 23, 24
(anatomy ).—Kass1anow, 1901, Zeit. fiir wissen. Zool., Bd. 69, pp. 299, 372, taf. 22, 24, 25.

Bell deep goblet-shaped, higher than wide, 15 to 25 mm. wide, and 25 to 30 mm. high,
including peduncle. Peduncle short, 4-sided, prismatic, and 4-chambered, one-fourth to one-
third as long as bell-height. Peduncle without longitudinal muscles. The 8 adradial arms
are short, wide, and 45° apart. 60 to 80 knobbed tentacles upon each arm. There are normally
no marginal anchors, although Antipa, 1892, and Gross, 1g00, record abnormal specimens
with 8 small tentacles, 4 perradial and 4 interradial.

The 8 gonads present the appearance of a 4-leaved cross, in the axes of which lie the 4
perradial, mesogonial pouches. The 8 arms of the gonads approach pair-wise and extend
under the subumbrella from base of throat-tube to bell-margin with their proximal ends
nearly touching. Each arm of the gonads has 10 to 16 feathery sinuosities and very numerous
saccules.

Color variable, as in most of the Stauromedusz, being olive-green, yellowish, reddish-
brown, or dark brown.

This form is found at Helgoland, German Ocean, where it lives upon the west coast of
the island upon Ulva, Chorda, or Fucus. Gross, 1900, gives the best description of its inter-
nal anatomy. Kassianow, Igor (p. 371), finds that if Craterlophus tethys be cut longitudin-
ally from the oral pole to the middle of the peduncle, each half regenerates a new individual.
If, however, the cut be not so deep the edges grow together and restore the former individual
although the scar remains as a constriction upon the bell and pharynx. He also reports the
finding of specimens of this medusa with more than 8 marginal lobes.

Craterlophus is imperfectly separated from Halimocyathus, bearing the same relation to it
that Lucernarta does to Haliclystus. We may, however, retain these generic names mainly
as a matter of convenience. In both Craterlophus and Lucernaria the perradial and interra-
dial tentacles or anchors are commonly absent, but occasionally they appear as an abnormality
and in such cases the medusa can not be separated from Halrmocyathus and Halliclystus
respectively.

Craterlophus macrocystis von Lendenfeld.

Craterlophus macrocystis, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 165; 1887, Australian Museum
Descript. Catalogue Medusx Australian Seas, part 1, p. 13.

Umbrella deep and bell-shaped, 12 mm. high, 6 mm. wide. Stalk 8 mm. high and 3 mm.
wide when extended. 8 short arms 45° apart, each with a cluster of about 30 tentacles. Gonads
feathery, as in C. tethys. Color dark olive-green. East coast of New Zealand on Macrocystis.

Rare.

STAUROMEDUSA]—CRATERLOPHUS, CAPRIA, LIPKEA. 539

The stalk of C. macroscystis is two-thirds as long as the bell, whereas in C. teth ys it 1s
only one-fourth to one-third of this length. In pene respects it resembles the very ‘closely
allied C. tethys with which it may prove to be identical.

Genus CAPRIA Antipa, 1893.
Capria, Antira, 1893, Mitth. Zool. Sta. Neapel, Bd. 10, p. 628.

The type species is Capria sturdzit Antipa, from the Island of Capri, Bay of Naples, Italy.
GENERIC CHARACTERS.

Stauromedusz with 8 adradial, lappet-like arms which lack knobbed tentacles, but are
each provided with a row of short, webbed, tooth-like or finger-shaped tentacles. Wart-like
clusters of nematocysts on subumbrella side of each of the 8 arms. No perradial or inter-
radial tentacles or “‘anchors.”” The circular muscle of the subumbrella is entire, not divided
into 8 isolated marginal muscles. The radial-muscle is funnel-shaped and spreads over the
entire surface of the subumbrella. The 4 septal edges of the perradial stomach-pouches
extend nearly to the bell-margin, where they are pierced by the ring-canal. There is a long
throat-tube, 8 adradial gonads, and an aboral stalk to the bell serving for attachment.

The genus Capria is the only representative of the family Capriide of Antipa, 1893,
which may be defined as Stauromeduse with 8 adradial arms which lack knobbed tentacles,
but have each a row of rudimentary tentacles joined by a web, one to the other. No anchors.
Ring-muscles of the subumbrella entire, not separated into 8 isolated sectors. Longitudinal
muscles equally developed over the entire subumbrella. Bell provided with an aboral stalk
for attachment. Stomach with 4 simple, perradial pouches as in Eleutherocarpide.

Capria sturdzii Antipa.

Capria sturdzit, ANtIpA, 1893, Mitth. Zool. Sta. Neapel, Bd. ro, p. 168, taf. 40, fign. 1-18.

Body 9 mm. long and 5.5 to 6 mm. wide; globular with a short, broad basal stalk or
peduncle about as long as the bell-portion itself. Basal plate of peduncle or stalk of bell
broad and flat and single-chambered, resembling a suctorial disk. There appear to be
normally 8 short, thick, adradial, paddle-like arms, although the single specimen found had
10 arms; these are hollow and devoid of knobbed tentacles, but are provided with a row of
16 to 20 tooth-like, or short finger-shaped, rudimentary tentacles which are fused one to
another by a web, thus giving the appearance of a multi-toed, bird-like web-foot. There
are 5 to 8 large clusters of nematocysts on the subumbrella side of each of the 8 arms.
There are no ‘“‘anchors” or suctorial tentacles, but the spaces around the bell-margin between
the 8 arms are open and somewhat less in width than are the arms themselves. The circular
muscle of the subumbrella is entire, not cut into 8 isolated muscles, as in certain other
Stauromeduse. There are 4 rows of gastric filaments along the 4 interradial taniola from
the middle of the central stomach nearly to the foot-plate. Mouth-tube long, prismatic,
4-sided, with 4 interradial, longitudinal furrows. Mouth-opening cruciform and quadratic.
8 band-shaped gonads, above (aboral in reference to) the taniola. Color yellowish-white.

Found attached to a Serpula tube dredged from a depth of 40 fathoms near the Blue
Grotto, Island of Capri, Bay of Naples. It is described in detail by Antipa.

Genus LIPKEA Vogt, 1886.
Lipkea, Vocr, 1886, Archiv. Sci. Phys. et Nat. Généve, sér. 3, tome 16, p. 356.
The type species is Lipkea ruspoliana Vogt, from the coast of Sardinia, Mediterranean.
GENERIC CHARACTERS.

Stauromeduse with 8 hollow (4 perradial and 4 interradial) arms. With a continuous
circular muscle. Neither tentacles nor “‘anchors.”” There are well-developed mucous glands
upon the subumbrella. The bell is attached by a sucker.

540 MEDUS OF THE WORLD.

The only known species is described from a single specimen found attached to a Gorgonian
at a depth of 50 fathoms off the Sardinian coast, Mediterranean. It is the only Stauromedusa
having perradial and interradial marginal lobes, and Vogt places it in a new family, the
Lipkeidz.

Lipkea ruspoliana Vogt.
Lipkea ruspoliana, Vocr, 1887, Mém. Inst. Nat. Génévois, tome 17, 53 pp., plates 10, 11, figs. 1-17; 1886, Arch. Sci. Physique

et Naturelles, Généve, sér. 3, tome 16, p. 356.

Bell flat and soup-tureen-shaped with a very short basal stalk, by means of which the
animal is attached. Bell 7 to 8 mm. wide, 4 mm. high, the basal stalk only 1.5 mm. long and
4mm. wide. 8 short, blunt, hollow, marginal lappets, 4 perradial and 4 interradial in position.
These lappets have plain, evenly rounded margins, and on their inner (centripetal) sides are
about 15 to 20 large, oval mucous glands (containing nematocysts ?), the openings of which
are scattered over the inner surface of each lappet. Subumbrella concave, but the central
mouth is elevated and bordered by 4 cruciform lips. The 4 deep, conical (subgenital ?) ostia
are interradial and alternate with the lips in position. Thus 4 of the $ lappets are in the
radii of the lips and 4 others are in the radi of the (subgenital ?) ostia. Ring-muscle entire,
not divided into sectors, and extends around margin of subumbrella at bases of the 8 lappets.
Longitudinal muscle-fibers extend radially outward from this powerful ring-muscle along the
subumbrella faces of the 8 lappets. There are also a few weak, longitudinal muscles in the
exumbrella near the stalk. There are neither tentacles, “‘anchors,” nor other marginal
appendages.

There are conspicuous clusters of gland cells in the ectoderm of the subumbrella. 4 large
clusters are perradial in position and lie at the base of the 4 angles of the cruciform mouth-
tube. There are also 8 linear clusters of these glands at the bases of the 8 lappets on the inner
side of the ring-muscle. Central stomach divided by 4 interradial septa into 4 perradial
chambers. These septa do not extend into the cavities of the 4 interradial lappets, so that
the perradial chambers of the stomach communicate one with another through the cavities
of these lappets. Stalk single-chambered.

No genital products were observed in the folded, membranous floors of the 4 (subgenital ?)
interradial ostia, but 4 pairs of branched, gastric filaments arise from the edges of the 4 inter-
radial septa at base of cesophagus. It would seem that the medusa was immature and that
the folded, follicular organs under the 4 ostia in the 4 interradii of the stomach are destined to
develop the sexual products.

Medusa translucent to milky in color and the clusters of nematocysts on the subumbrella
are yellow.

A single (immature ?) specimen was found by Vogt attached to a Gorgonian at a depth
of 50 fathoms at Alghero on the northeast coast of Sardinia, Mediterranean. Vogt describes
the specimen in detail.

CORONATAE—-PERICOLPA. 541

Order CORONATZ Vanhoffen, 1892.

Discomeduse (in part), Hacker, 1866, Generelle Morphologie, Bd. 2, p- 60; 1880, Syst. der Medusen, p. 450.

Coronate, VANHOFFEN, 1892, Ergeb. der Plankton Exped., Bd. 2, K. c., p. 21; 1902, Wissen. Ergeb. deutsch. Tiefsee Exped.,
Dampfer Valdivia, Bd. 3, Lief. 1, p. 51; 1906, Nordisches Plankton, Acraspede Medusen, Nr. 11, p. 41.

Coronata, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 5; 1907, Ergeb. Fortschritte der Zool., Bd. 1, pp. 191,
199.—Bicetow, H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p+ 20.

CHARACTERS OF THE CORONATZ.

Scyphomedusze with marginal tentacles, a single central mouth-opening, and with the
bell-margin cleft into lappets. The rhopalia arise from clefts between these lappets, and
their entodermal cores contain a terminal mass of crystalline concretions of entodermal origin.
Ocelli may or may not be present. The medusz are free-swimming and have no aboral
stalk for attachment.

With a circular, or coronal, furrow in the exumbrella, and peripheral to this there is a zone
of gelatinous thickenings in the radii of the tentacles and sense-organs. These thickenings,
or pedalia, are divided one from another by radiating clefts which alternate in position with
the marginal sense-organs and the tentacles, and are in the middle (axial) lines of the marginal
lappets. These marginal lappets project beyond the zone of the pedalia. The tentacles are
solid, or not hollow, throughout their lengths. The throat-tube is simple, short, and provided
with simple lips, without curtain-like appendages.

The families of the Coronatz are as follows:

Periphyllide Craus, 1886. 4 interradial rhopalia and 4 or more tentacles.
Paraphyllinide Maas, 1903. 4 perradial rhopalia and 4 or more tentacles.
Ephyropside Craus, 1883. 8 rhopalia (4 perradial and 4 interradial) and 8 or more tentacles.

Collaspide Harcxet, 1880. Numerous rhopalia, alternating with an equal number of tentacles.
Atorellide VANHOFFEN, 1902. With 6 rhopalia and 6 tentacles.

Family PERIPHYLLID& sensu Claus, 1886.

Peromeduse (in part), HAEcKEL, 1880, Syst. der Medusen, p. 396.

Periphyllide, Craus, 1886, Class. der Medusen, Arbeit. Zool. Inst. Univ. Wien, Bd. 7, pp. 97-110.—VANHOFFEN, 1892, Akalephen
der Plankton Expedition, Bd.2, K.d., p.21.—Maas, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, No.1, p.28;
1906, Fauna Arctica, Bd. 4, Lfg. 3, p. so1.—VANHOFFEN, 1906, Nordisches Plankton, No. 11, Acraspede Medusen, p. 41.—
Bicetow, H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p. 23.

Peromeduse, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 166.

FAMILY CHARACTERS.

Coronate with 4 interradial pedalia which bear marginal sense-clubs, and with 4, or
more, pedalia which bear tentacles.

The genera of the Periphyllide are as follows:

Pericolpa Harcket, 1880, sensu Vanhoffen, 1902. With 4 perradial tentacles, 8 adradial lappets, 8 gonads.
Periphylla SteenstRuP, 1837. With (4 X 3) 12 tentacles, 4 perradial, 8 adradial. 16 lappets, 8 gonads.
Periphyllopsis VANHOFFEN, 1900. With (4 X 5) 20 tentacles, 24 lappets.

(?) Nauphantopsis Fewxes, 1885; sensu Vanhoffen, 1902. With (4 X 7) 28 tentacles, 32 lappets.

Genus PERICOLPA Haeckel sensu Vanhoffen.

Pericolpa+ Pericrypta, HAECKEL, 1880, Syst. der Medusen, pp. 413, 414, 640.
Pericol pa, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Valdivia, Bd. 3, Lfg. 1, p. 5o.—Maas, 1903, Scyphomedusen
der Siboga Exped., Monog. 11, p. 12; 1906, Die Arktischen Medusen, Fauna Arctica, Bd. 4, p. 502.

GENERIC CHARACTERS.

Periphyllide with 4 interradial rhopalia, 4 perradial tentacles, 8 adradial lappets. The
8 gonads are adradial or on both sides of the 4 interradi.

Haeckel’s Pericrypta is doubtless only a more advanced stage in the growth of Pericolpa.
The interradial taniolz are solid in the young, but become hollowed by the development of
4 pits in the floor of the subumbrella, and the gastral filaments increase with age. In the
structure of the gastrovascular system this genus resembles Per:phylla, and were it not for
the probability that the number of metameres apparently does not increase in the free ephyra

542 MEDUS OF THE WORLD.

of the Coronatz we would be inclined to regard Pericol pa as being only the young of Perzphylla;
for the development of 8 adradial lappets and 8 corresponding tentacles would change the
medusa to Periphylla.

The species founded by Haeckel are separated upon slight distinctions, some of which
represent mere stages in growth. Probably there are but two forms, P. quadrigata with an
elongate, pointed bell and P. campana with a flat, dome-like bell; but even this can not now
be determined with any degree of certainty.

This genus includes the simplest and possibly most primitive of the Periphyllide.

Pericolpa quadrigata Haeckel.
Pericol pa quadri gata, HAECKEL, 1880, Syst. der Medusen, p. 413, taf. 23, fign. 1-12; P. galea, Ibid., p. 414; (?) P.tetralina, p.640.—

Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 12.

Bell 40 mm. high and 30 mm. wide. The pointed, dome-like apical half of the exumbrella
above the ring-furrow is as high as the width of the pedal and lappet-zone below. The 4
perradial tentacular pedalia are somewhat wider than the 4 interradial, rhopalar ones and
somewhat longer than the 8 marginal lappets. The 4 tentacles are about as long as bell-
height. Throat-tube and stomach wide, filling the greater part of bell-cavity. The basal
part of the stomach leads into the gastrovascular space of bell by 4 perradial ostia, which
are lined with gastral filaments. These ostia lead into a wide sinus which is interrupted by
4 short, partial septa in the interradii. Peripheral to these septa is another wide ring-sinus
which sends out 8 canals in the radii of the sense-organs and tentacles. These radiating canals
fork at their ends and extend around the edges of the lappets, forming a marginal ring-canal.
The 8 gonads are grouped in 4 pairs on either sides of the interradu. Their inner ends are
close together but they diverge outwardly. Color (?)

Found in the Antarctic, southeast of Kerguelen Island by the Challenger, and apparently
identical with P. galia from the east coast of Australia.

Pericolpa campana Maas.
Pericrypta campana, Harcket, 1880, Syst. der Medusen, p. 414.
Pericolpa campana, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 13, taf. 3, fign. 19-22.

Bell about 15 mm. high and 15 mm. wide, with thick gelatinous walls. A deep ring-
furrow around exumbrella separates the evenly rounded dome-like center of bell from marginal
zone of lappets. The 8 clefts between the 8 pedalia are also very deep. The 4 tentacular
pedalia are nearly similar in size to the 4 pedalia of the sense-organs. The 4 tentacles are
tapering and shorter than bell-radius. They are solid and their axial cores project into the
gelatinous substance. These tentacles are situated in the perradi. Each of the 4 interradial
marginal sense-clubs contains a terminal entodermal concretion and a ventral bulbular
swelling, but no ocellus. The 8 marginal lappets are semicircular.

Stomach wide at base, and there are 4 interradial clusters, each with at least 30 gastric
cirri. The stomach is connected with the gastrovascular space of the bell by 4 perradial
openings. Peripheral to these there is a wide ring-sinus and this in turn gives rise to 8 radiating
canals in the radii of the tentacles and sense-organs, and these fork and communicate one
with another at their outer ends, forming a marginal ring-canal. The circular muscles of the
subumbrella are well developed and there are radial-muscle strands near the bases of the
tentacles. There appear to be 8 gonads. Maas records 7 of them irregularly arranged in a
zone at middle of subumbrella, and Haeckel’s specimen was so poorly preserved that he does
not record the character of the gonads. Color (?)

Haeckel’s specimen came from the region of New Zealand, while Maas records one from
the Malay Archipelago, where it was obtained in a vertical net hauled from a depth of 500
fathoms in 17.6’ S. lat., 129° 14.5’ E. long.

This medusa is distinguished by its oval gonads.

Pericolpa tetralina Haeckel.

Pericolpa tetralina, Harcker, 1880, Syst. der Medusen, p. 640.

This medusa is probably identical with P. quadrigata, but both ends of the 8 gonads
diverge from the 4 interradii, while in P. guadrigata only the outer ends diverge and the inner

CORONAT—PERICOLPA, PERIPHYLLA. 543
ends approach one another closely. The medusa is very briefly mentioned by Haeckel and
appears to be immature, being only 20 mm. high and 16 mm. wide. Found off the south coast
of Australia. Described by Haeckel from a single preserved specimen.

Genus PERIPHYLLA Steenstrup, 1837.

Periph ylla, SteenstRuP, 1837, Acta et Cat. Mus. Hafniensis——Haecket, 1880, Syst. der Medusen, p. 418; 1881, Deep-sea Medusx
Challenger Report, Zool., vol. 4, p- 63.—Craus, 1886, Arbeit. Zool. Inst. Univ. Wien, Bd. 7, P- 99-—VANHOFFEN, 1892,
Ergeb. der Plankton Exped., Bd. 2, K.d., pp. 4, 6, 21-—von LenDENFELD, 1884, Proc. Linnean Soc, New South Wales,
vol. 9, p. 168.—Maas, 1897, Mem. Mus. Cann Zool. at Harvard College, vol. 23, No. 1, pp. 28-64; 1904, Résult. Camp.
Sci. Prince de Monaco, fasc. 28, p. 443 1903, Scyphomedusen der Sihoga Expedition, Monog. 11, p. §; 1907, Ergeb. Fort-
schritte der Zool., Bd. 1, pp. 199, 219, etc.—VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer
Valdivia, Bd. 3, Lfg. 1, p. 21; 1906, Nordisches Plankton, Nr. 11, p. 41-—Bicetow, H. B., 1999, Mem. Mus. Comp. Zool.
at Harvard College, vol. 37, p. 24.

Periphema, Harcket, 1881, Deep-sea Meduse Challenger Exped., p. 84.

GENERIC CHARACTERS.

Periphyllidae with 4 interradial rhopalia, 12 tentacles, 4 perradial and 8 adradial. 167
marginal lappets grouped into 4 pairs of rhopalar and 4 pairs of tentacular lappets. A deep
annular furrow separates the dome-like apex of the exumbrella from marginal zone of bell.
Between this ring-furrow and the lappets is a zone of 16 pedalia, 12 in the tentacular and 4 in
the rhopalar radii, and these are separated one from another by 16 deep, radiating clefts,
which extend down the mid-axial lines of the lappets. There are 4 deep, interradial subgenital
pits in the floor of the subumbrella, lined along their edges by rows of internal gastric cirri.
The large central stomach extends peripherally outward into the subumbrella in the 4 perradit.
These 4 openings lead into a wide ring-sinus in the subumbrella which in turn sends out a
radiating vessel in the radius of each tentacle and rhopalium, 16 in all. These vessels fork
before reaching the tentacles or rhopalia, and their diverging ends curve around the edges of
the lappets and form a marginal ring-canal.

The 4 interradial septa are bordered by lines of gastric cirri and there are 8 U-shaped
adradial gonads, one on either side of each septum. anh free ends of each gonad are directed
inward toward the stomach, and their convexities point outward toward the bell-margin.
The rhopalia have no ocelli, but contain a proximal mass of entodermal pigment and a distal
concretion.

The medusz of Perrphylla are deep-sea forms of very wide distribution. The so-called
species are not well separated, being based on the relative height and width of the bell and
on slight color distinctions. The bell appears to become relatively flatter as growth proceeds,
and therefore its proportions would seem to afford a poor criterion for specific distinctions.
Both Vanhoffen, 1902, and Maas, 1904, recognize but 3 species as follows:

P. hyacinthina with high, narrow bell, its height being to width as 44 to 23. The lappet pouches are dark-brown, so that
the gonads can not be seen through them from the outside.

P. dodecabostrycha with bell flat, bluntly pointed, its height being to width as 27 to 18. Gonads visible through the lappets
from the outside.

P. regina with dome-like rounded bell and light red-brown color.

Periphylla “ mirabilis,’ Haeckel, appears to be identical with P. regina; and P. “humilis,’ Fewkes, appears to be an
imperfect P. dodecabostrycha. P. peronit, Haeckel=Charybdea periphylla, Péron et Lesueur, 1809, p. 332, is too
imperfectly known to be classified.

I think it probable that there is but a single species of Perrphylla (P. hyacinthina), and
that P. dodecabostrycha and P. regina are only varieties or local races. It is impracticable
to attempt to separate species upon slight differences of form or proportions in their bells
especially when such characters are subject to constant changes due to growth or to state of
contraction. Bigelow, 1909, who has studied many specimens of these meduse, concludes
that P. hyacinthina and P. dodecabostrycha are identical.

Peripalma corona of Haeckel, 1880 (Sitzungsber. Jena. Gesell. fiir Med. und Naturw.
Jahrg. 1880, Feb. 20; Syst. der Medusen, p. 418), is probably a young Perrphylla. Haeckel
states that the genus is characterized by the absence of interradial pits or perradial concavities
in the subumbrella, and that the 4 interradial teniol of the basal stomach are solid ligaments
without gastral filaments.

544 MEDUSA OF THE WORLD.

A single specimen was found by Haeckel at Algeciras in the Straits of Gibraltar. He
states that owing to the general transparency of the specimen and to its smallness he is able
to describe it only through reference to the larger species of Pertphylla found by the Challenger
expedition. He states that the bell is 30 mm. high, 20 mm. wide, helmet-shaped. The pedal
zone nearly as wide as the lappet-zone. The 16 pedalia are of not quite equal size each to
each. The 16 lappets tongue-shaped, sharp-pointed, the 8 tentacular more projecting than
the 8 rhopalar lappets. 12 tentacles as long as the bell-height and one-third as wide as the
lappets at their bases. Basal, central, and buccal stomachs of nearly equal length.

Color violet, bell more red, tentacles and throat-tube more blue, gonads dark-red.

Periphylla hyacinthina Steenstrup.

Periphylla hyacinthina, Sreenstrur, 1837, Acta Mus. Hafniensis.—Harcket, 1880, Syst. der Medusen, p. 419, taf. 24, fign. 11-
16.—Fewkes, 1886, Report Commiss. Fish and Fisheries U.S. A., for 1884, p. 930.—Craus, 1886, Arbeit. Zool. Inst. der
Univ. Wien., Bd. 7, p. 99, fig. 1, p. 100.—AGassiz, A., 1888, Bull. Mus. Comp. Zool. at Harvard College, vol. 15, p. 131,
fig. 426.—Vannorren, 1891, Zool. Anzeiger, Jahrg. 14, p. 38; 1908, Deutsch. Siidpolar Expedition, 1g01-03, Bd. 10,
Zool. 2, p. 36; 1892, Ergeb. der Plankton Expedition, Bd. 2, K. d., p. 6, taf. 1, fign. 1-10; taf.2, fign. 3-8; taf. 3, fign. 1-7;
1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Dampfer Valdivia, Bd. 3, Lfg. 1, p. 23; 1906, Nordisches Plankton, Nr. 11,
p. 42, fig. 1 Browne, 1903, Bergens Museums Aarbog, No. 4, p. 30-—Maas, 1904, Résult. Camp. Sci. Prince de Monaco,
fasc. 28, p. 47, plate 5, fig. 35; planche 6, figs. 45-46; 19c6, Fauna Arctica, Bd. 4, Lfg. 3, pp. 502, 511 (review of literature ).—
Bicetow, H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p. 26, plates 1 and 9.

Charybdea hyacinthina, Sreenstrur, 1842, Acta Mus. Hafniensis.

The following description applies to the typical P. hyacinthina.

This so-called species is distinguished by its high, narrow bell, the ratio of height to
width being usually about as 44 is to 23. The lappet-pouches are so densely pigmented with
dark purple-brown that the gonads can not be seen through them from the outside.

Bell 80 mm. high and about 42 mm. wide, but it may be much flatter and wider (see fig.
343). The upper, or aboral, half of the exumbrella i is dome-shaped or pyriform with a smooth
external surface. Just below this dome is a deep horizontal circular constriction which is
occupied by circular muscle-fibers. Below this constriction the exumbrella flares slightly
outward, and in this region we may distinguish an upper pedal-zone and a lower lappet-
zone. The pedal-zone is divided by 16 deep, longitudinal furrows into the same number of
projecting, wedge-like thickenings, the pedalia. The 4 interradial pedalia are smaller than
the others and lie in the radii of the 4 marginal sense-organs; while the 12 perradial and
adradial pedalia are situated in the radii of the tentacles. .

There are 16 well-developed marginal lobes, arranged in 8 pairs. 16 deep longitudinal
furrows lie in the mid-radial lines of the lappets and separate the 16 pedalia, so that a furrow
extends about two-thirds the distance down the exumbrella surface of each marginal lappet.
The 4 interradial rhopalar clefts between the lobes are only about two-thirds as deep as the
12 tentacular clefts. Each sense-organ contains a proximal mass of dark-brown entodermal
pigment and a distal concretion, which is protected by an aboral fold forming a niche for its
protection.

The 12 tentacles are equally developed and are each a little longer than the bell-height.
They are solid and are provided with well-developed, longitudinal muscle-fibers. In the
cavity of the subumbrella 4 long, funnel-shaped, interradial pits extend inwardly along the
sides of the central stomach nearly to the aboral apex, where their points do not quite touch.
There is a well-developed zone of circular muscle-fibers in the subumbrella just above the
bases of the tentacles and sense-organs. This zone is divided by 16 longitudinal selvages
into 16 distinct parts; each selvage extends down the middle of a lappet, and thus the adjoin-
ing halves of each pair of adjacent lappets are connected by the circular muscles. Besides
the circular muscles, 8 well-developed, longitudinal muscle-strands lie in the inner, or proximal,
part of the subumbrella wall; 4 of these are perradial and 4 interradial. They are deltoid
in shape, their broad bases extending out into the distal part of the subumbrella, to the upper
part of the zone of circular muscle-fibers. There are 8 U-shaped gonads which are on both
sides of the 4 interradial septa, with their convexities directed outward; thus they appear
to be adradial in position and alternate with the 8 longitudinal muscles of the inner part of
the subumbrella.

The stomach extends from the inner apex of the subumbrella to about the level of bell-
margin. Its cavity may be divided into 3 regions, which we may designate, respectively

CORONAT AR

PERIPHYLLA. 545

as the basal, central, and buccal stomach. The central stomach is a 4-sided prism, the sides
being interradial and the angles perradial in position. The lower or buccal part of the stomach
hangs freely in the cavity of the bell, being joined to the subumbrella at 4 perradially situated
points at its inner end. There are 4 longitudinal, interradial, thickened regions in the wall
of the buccal stomach, which extend downward to the mouth-opening. ‘The central stomach
is a wide cayity which communicates by 4 perradial openings with the gastrovascular space
of the subumbrella of the medusa. These 4 openings are narrow, elongate, longitudinal
clefts, and their edges are lined with numerous gastric cirri which project into the space of

Periphylla hyactnthina.

Fic. 342.—After Vanhéffen, in Valdivia Expedition.
Fic. 343.—After Vanhéffen, in Nordisches Plankton.

Showing variation in shape of bell.

the stomach. The basal stomach is a 4-sided pyramid and 8 rows of gastric cirri extend
up the 4 sides near the angles to the apex of the pyramid where they meet. These rows of
cirri are continuous with those surrounding the 4 perradial side-openings of the central stomach.
The gastrovascular coronal sinus of the subumbrella is thus connected with that of the stomach
by the 4 perradially situated ostia of the central stomach. These open into this wide annular
cavity which occupies the mid-region of the subumbrella above the zone of circular muscles.
Below these, however, it appears as a broad, simple, annular space, which extends outward

546 MEDUSE_OF THE WORLD.

into the 16 marginal lappets. A partial septum extends, however, down the midline of each
lappet, and the gastrovascular space forms a marginal ring-canal around these septa. Diver-
ticula of the annular space also extend for a short distance into the bases of the tentacles,
but the main entodermal cores of the tentacles are solid. A good idea of the structure of the
gastrovascular cavity of the bell may be obtained from Vanhoffen’s figure 3, taf. 1, in “‘ Ergeb.
der Plankton Expedition,” Bd. 2, K. d.

The inner surface of the subumbrella is purple, while the pedalia are copper-brown
colored and the tentacles and lappets are of a milky-blue translucency. The gonads can
not be seen through the densely pigmented bell-walls.

This deep-sea medusa is occasionally found upon the surface in the colder parts of the
North Atlantic. It has been taken in the Bay of Biscay; at the Azores; off Cape Hatteras,
North Carolina; at Martha’s Vineyard; at Spitzbergen, and still more commonly off the
coast of Greenland, but its true habitat is undoubtedly in the deep sea, at or near the bottom.
Nearly all recent deep-sea expeditions record it, and the Valdivia found it in both the Atlantic
and Indian Oceans. The best descriptions are those of Vanh6ffen and Maas.

Haeckel states that the bell of this medusa may become 160 mm. high and 120 mm.
wide, but later observers have not seen specimens of such great size.

Periphylla hyacinthina forma dodecabostrycha.

Chrysaora (dodecabostrycha) dubia, BranpvT, 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 387, taf. 29, 30.

Periphylla dodecabostrycha, Harcke, 1880, Syst. der Medusen, p. 421.—VANuGFFEN, 1892, Ergeb. der Plankton Exped., Bd. 2,
K.d., taf. 2, fign. 1,2; 1908, deutsche Siidpolar Expedition 1901-1903, Bd. 10, Zool. 2, p. 35; 1902, Wissen. Ergeb. deutsch.
Tiefsee Exped., Valdivia, Bd. 3, pp- 21, 23—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 6, taf. 2,
fig. 15; taf. 12, fig. 107 (figure of rhopalium); 1904, Résult. Camp. Sci. Prince de Monaco, fasc. 28, p. 47, planche 5, figs.
36, 37; 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. 64, taf. 11, fig. 1—Mayer, 1906, Bull. U. S. Fish
Commiss. for 1903, vol. 23, part 3, p. 1136, plate 3, figs. 5, 6.

non Periphylla dodecabostrycha, LoBiaNnco, 1903, Mitth. Zool. Sta. Neapel, Bd. 16, p. 219, taf. 7, fig. 3 (this is a Paraphyllina).

(2) Periphylla humilis, Fewxes, 1886, Report Commiss. Fish and Fisheries U. S. A., for 1884, p. 931-

Bell higher than wide in young, wider than high in well-grown meduse. Thus when the
medusa is 50 mm. wide the bell is 55 mm. high; when 45 mm. wide it is 45 high; and when
100 mm. wide it is 70 mm. high.

The gonads may usually be seen through the gelatinous walls of the bell, but not so
clearly in old as in young specimens. Thus the bell is said to be lower, flatter, and less pointed
than in P. hyacinthina, and the bell-walls are more transparent.

Widely distributed over the floor of the
great oceans, and especially in tropical parts of
the Pacific, west coast of Mexico, coast of Chile,
Hawaiian Islands, Philippine Islands, Indian
Ocean, Malay Archipelago, Mediterranean,
Guinea Stream off Atlantic coast of Africa.

It is probably only a variety of, or even
only a growth-stage of, P. hyacinthina.

Periphylla hyacinthina forma regina.

Periphylla regina, Harcker, 1880, Syst. der Medusen, p. 421;
1881, Deep-sea Meduse Challenger Exped. Report, Zool.,
vol. 4, p-85, plates 24, 25.—Maas, 1897, Mem. Museum.
Comp. Zool. at Harvard College, vol. 21, pp. 29, 64, taf. 10,
1 fig.; 1903, Scyphomedusen Siboga Exped., p. 6.—VaANn-
HOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped.,
Valdivia, Bd. 3, Lief. 1, pp. 21, 23; 1908, deutsche Siid-
polar Expedition, 1901-1903, Bd. 10, Zool. 2, p. 36.

Periphylla mirabilis, HAecKEL, 1880, Syst. der Medusen, p. 442;
1881, Challenger Exped. Report, Zool., vol. 4, p. 64, plates
18-23, 40 figs.

This appears to be merely a light violet or
rusty-reddish variety of P. hyacinthina. Bell
usually low, dome-like, about 150 mm. wide, and

Fic. 344.—Pertphylla regina, after Vanhoffen, in : = 5
Tiefsee Expedition Valdivia. : nearly 1.5 times as wide as high, but Haeckel

CORONATA:—PERIPHYLLA, PERIPHYLLOPSIS. 547

records a specimen fully as high as wide. According to him, the pedalia are rectangular and

longer than wide, but according to Maas, 1897, they are nearly circular in outline. The

tentacles are said to be short and thick, and the stomach very large and wide, but these points

as well as the proportions of the bell, are probably affected largely by growth and contraction.
Found on the bottom of the Pacific and Atlantic from the Antarctic regions to the tropics.
It is the largest form of Periphylla and may become 200 mm. wide.

Genus PERIPHYLLOPSIS Vanhiffen, 1900.
Peri phyllopsis, VANHOFFEN, 1900, Zool. Anzeiger, Bd. 23, p. 278; 1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Valdivia, Bd. 3,

Lfg. 1, p. 27—Maas, 1907, Ergeb. und Fortschritte der Zool., Bd. 1, p- 195.—Bicetow, H. B., 1909, Mem. Mus. Comp.
Zool. at Harvard College, vol. 37, p. 27.

The type species is Periphyllopsis brauert Vanhoffen, of the Indian Ocean; from a
depth of 1,200 fathoms.
GENERIC CHARACTERS.

4 interradial rhopalia, 4% 5 (20) tentacles, 46 (24) lappets.

Periphyllopsis braueri Vanhdffen.

Periphyllopsis braueri, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, P- 27,
taf. 2, fig. 7.—Bicrtow, H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p. 28, plates 9 and 12.

Vanhoffen’s single specimen was so imperfect that he could not venture to present a
detailed description of it. Recently, however, Bigelow describes a more nearly perfect specimen

345:

Periphyllopsts braueri.

Fic. 345.—After Vanhoffen, in Valdivia Expedition. Aboral view of bell.
Fic. 346.—With the walls of the stomach torn away leaving only its base with the gastric cirri.

After H. B. Bigelow, in Mem. Museum Comp. Zool. at Harvard College, 1909.

from the collections of the Albatross, and the account here given is mainly derived from his
description.

Bell flattened, 60 mm. wide and 25 mm. high, resembling an A4tolla in shape. Central
disk about 50 mm. wide; ring-furrow deep. 4 interradial thopalia, 24 (46) ovate marginal
lappets, and 20 (45) tapering tentacles, about as long as the bell-diameter. Thus the radial
arrangement of the various organs is the same as is seen in Periphylla, only the numbers of
lappets and tentacles being greater. The ring-muscle of the subumbrella is very weak as in

548 MEDUS2 OF THE WORLD.

Atorella. The peripheral canal-system resembles that of Per:phylla. The central stomach
opens into a wide ring-sinus by 4 perradial ostia separated by 4 interradial septa. The ring-
sinus is about 10 mm. broad and is at the zone of the gonads. On its outer side it gives off 24
broad, spoke-like radial-canals in the radii of the rhopalia and tentacles, and these canals are
connected one with another by a festoon ring-canal at the margin, extending in loops around the
lappet margins. There are 8 oval gonads, adradial in position and equidistant one from
another. The mouth parts were lost in both Vanhdffen’s and Bigelow’s specimens, but there
are about 80 to 100 simple gastric cirri arranged in a single linear row. There are 4 inter-
radial ostia in the subumbrella. The entire entodermal system is chocolate-red.

The Valdivia specimen was dredged from between 1,200 fathoms and the surface in the
Indian Ocean between New Amsterdam and Cocos Islands, and the d/batross specimen which
was studied, while yet alive, by Bigelow was obtained in the Humboldt current off the coast of
Peru between 400 fathoms and the surface.

Genus (?) NAUPHANTOPSIS Fewkes, 1885.

Nauphantopsis, Fewxes, 1885, Report Commiss. Fish and Fisheries U. S. A. for 1883, p. 596; 1886, Report Commiss. Fish and
Fisheries U. S. A. for 1884, p. 944.—VANHOFFEN, 1892, Ergeb. der Plankton Expedition, p. 21; 1902, Wissen. Ergeb.
deutsch. Tiefsee Exped. Valdivia, Bd. 3, Lfg. 1, p. §1.—Hareitr, 1904, Bulletin U. S. Bureau of Fisheries, vol. 24, p. 66.—
Maas, 1907, Ergeb. und Fortschritte der Zool., Bd. 1, p. 199.

GENERIC CHARACTERS.

Coronatz with 32 marginal lappets, 4 interradial sense-organs, and 28 tentacles. With
an annular furrow and 32 radial ridges (or pedalia) upon the exumbrella in the radii of the
tentacles and sense-organs.

Owing to the incompleteness of our knowledge of the only known species of this genus it
must remain problematical.

Nauphantopsis diomedee Fewkes.

Nau phantopsis diemedea, Fewxes, 1885, Report Commiss. Fish and Fisheries U.S. A. for 1883, p. 596; 1886, Report Commiss.

Fish and Fisheries U.S. A. for 1884, p. 946, plate 6, figs. 1, 2; 1888, Amer. Journ. Sci., ser. 3, vol. 35, p- 173; Ann. and

Mag. Nat. Hist., ser. 6, vol. 1, p. 255 —Harairt, C. W., 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 66.

Fewkes gives an unsatisfactory account of this medusa owing to the poor preservation
of his material. We are uncertain whether there are 4 or 8 marginal sense-organs.

Disk quite flat, 70 mm. in diameter. Central part of exumbrella flat and surrounded
by an annular furrow; diameter of this region about 35 mm. Centrifugally from the annular
furrow there is a zone about 10 mm. wide consisting of 32 radial elevations separated by 32
deep radial furrows; these elevations lie in the radii of the tentacles and sense-organs and
alternate with the lappets. Each elevated ridge is bifurcated at its outer end by a deep median
cleft. The 32 lappets are long and rectangular with rounded outer edges. They are each
about 10 mm. long and 8 mm. wide. There are 4 or 8 (?) marginal sense-organs and 24 ( ?)
or 28 tentacles. “The sense-organs were not observed in the specimen studied by Fewkes.
Tentacles slender and flexible and about 80 mm. in length. Subumbrella (?) Mouth (?)
Gonads (?) Color (?)

A single specimen was dredged from a depth of 2,033 fathoms in N. lat. 38° 30’, W. long.
69° 8’.

Family PARAPHYLLINIDZ Maas, 1903.
Paraphyllinide, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 6.

FAMILY CHARACTERS.
Meduse Coronate with 4 perradial rhopalia and 4 or more tentacles.

This family differs from the Periphyllide only in having the marginal sense-organs
perradial instead of interradial.

There is only one known genus among existing medusz, this being Paraphyllina; but
‘this is very closely related, if not identical, with Paraphyllites, a fossil medusa of the litho-
graphic slate of Kelheim.

Maas records a Paraphyllina from the Malay Archipelago, and another specimen was
taken by the Krupp yacht Puritan at a depth of 500 fathoms, near Capri, Bay of Naples.

CORONATAS—PARAPHYLLINA. 549

Genus PARAPHYLLINA Maas, 1903.
Paraphyllina, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p.6; 1907, Ergeb. und Fortschritte der Zool., Bd. 1,
p- 195-
The type species is Paraphyllina intermedia Maas, 1903; from the Malay Archipelago,
between 100 fathoms and the surface.

GENERIC CHARACTERS.

Coronatz with 12 tentacles, 4 perradial marginal sense-organs, and 16 lappets. The 12
tentacles are interradial and adradial with reference to the stomach and lips. There are
4 pairs (8) of interradial gonads. The marginal sense-organs have a terminal lithocyst-sac
and a yentral bulb-like swelling, just beyond which is an eye with ectodermal lens and ecto-~
dermal pigment.

The short 4-sided throat-tube and flat disk-like bell resemble the condition noted in
Nausithoé, but in its 4 marginal sense-organs and 12 tentacles it recalls the condition seen in
Periphylla, with the important Aincrences that in Pertphylla the sense-organs are interradial,
whereas in Paraphyllina they are perradial in position.

Maas places this genus in a new family which he calls the Paraphyllinidez. The characters
of this family are hose of its only genus, Paraphyllina. It is closely related to the fossil genus
Paraphyllites, Maas, 1906 (Neuen Jahrbiich. fiir Mineralogie, Geol. und Palaontol., Bd. 12,
p- 90, 4 fign.). This fossil medusa differs from recent Paraphyllina only in that its 8 gonads are
adradial and placed 45° apart, whereas in Paraphyllina they are grouped in pairs on both
sides of the 4 interradii.

Paraphyllites distinctus is described by Maas from a well-preserved specimen. It is
15 mm. wide and has a well-developed coronal furrow, 16 pedalia, 4 perradial marginal
sense-organs, 12 tentacles, and 8 adradial gonads 45° apart. It is from the lithographic slate
of Kelheim. The latest reference to this fossil is that of von Ammon, 1908, Geonostischen
Jahrespeften, Jahrg. 19, p. 170.

Paraphyllina intermedia Maas.
Paraphyllina intermedia, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 8, taf. 2, fign. 10-14; taf. 11, fig. 106.
Periphylla dodecabostrycha, Lowianco, 1903, Mitth. Zool. Sta. Neapel, Bd. 16, p. 219, taf. 7, fig. 3. See also Maas, 1904, Résult.

Camp. Sci. Prince de Monaco, fasc. 28, p. 48, foot-note.

Bell flatly rounded, 8 mm. high and 15 mm. wide, without pointed apex. The coronal
furrow around the exumbrella is deep and separates the smooth, central, flatly-rounded apex
of exumbrella from the marginal zone of lappets. Lappet-zone about as wide as radius of
central disk of exumbrella. The 16 pedalia of the marginal zone are rectangular with rounded
angles and are separated one from another by deep ‘radiating furrows. blinese pedalia are
in the radii of the tentacles and sense-organs and alternate with the lappets. The 12 pedalia
of the tentacles are of uniform width, while the 4 pedalia of the marginal sense-organs are
only about half as wide as the former. The 16 marginal lappets are oval and bluntly pointed,
and the 8 lappets flanking the 4 sense-organs are somewhat narrower than the others. The
12 tapering, hollow tentacles are all of equal length and are not quite as long as the diameter
of the bell.

The 4 marginal sense-organs are each covered by a hood-like fold of the exumbrella.
Each sense-club contains a small terminal sac-shaped entodermal concretion. On the ventral
(subumbrella) side of the sense-club is a large ectodermal eye with a cup-like mass of pig-
ment and a spherical lens. On the inner side of the eye and upon the ventral side of the
sense-club is a bulb-like swelling. Altogether the sense-clubs resemble those of Nausithoe,
but the eye is larger.

The throat-tube is short and 4-sided and the mouth is a simple cruciform opening. There
are 4 interradial rows of gastric filaments. The coronal ring-canal gives rise to 16 peripheral
pouches in the radii of the sense-organs and tentacles, and these are put into communication
one with another by means of a marginal ring sinus, as in Nausithoé.

The gonads resemble those of Palephy yra and are intermediate in character between
those of Periphylla and those of Nausithoé. They consist of 4 pairs of bean- shaped or egg-
shaped sacs. These 8 sacs project from the subumbrella floor on both sides of the 4 inter-
radii near the sides of the throat-tube and centripetal to the ring-muscle.

550 MEDUS OF THE WORLD.

There is a well-developed ring of circular muscle-fibers in the subumbrella, beyond
the gonads and inside of the insertions of the tentacles. This zone of ring-muscles appears
broken in the mid-radii of the lappets, thus forming 16 trapezoids in the radii of the sense-
organs and tentacles. Radial-muscle strands extend outward in the tentacular radii and con-
verge at the bases of the tentacles.

The medusa is colorless save for the pigment of the eyes in the marginal sense-organs
and for 4 interradial masses of red-brown pigment in the central stomach, leaving a clear
cruciform space between them.

Two specimens were taken by the S:boga in the Malay Archipelago, in vertical nets,
from a depth of 100 fathoms, at 6° 2’ S. lat., 123° 57.7’ E. long.

Dr. Lobianco describes a similar medusa in the collection made by the Krupp yacht
Puritan, froma depth of 500 fathoms, near Capri, Bay of Naples. He generously permitted
me to study the very well-preserved specimen of the medusa in the Zoological Station at
Naples. It resembles Maas’s specimens from the Malay Archipelago except that the gonads

Fic. 347.—Paraphyllina intermedia. From a specimen in the
Naples Zoological Station.
(female) are more slender and their outer ends somewhat longer than in the Malayan speci-
mens. There are 4 deep interradial, crescentic pits in the floor of the subumbrella between
the gonads and the gastric cirri. The medusa was 15 mm. wide and in all respects (save
in the minor details mentioned above) it appears to be identical with the East Indian medusa.
A figure of Dr. Lobianco’s medusa drawn by me from his specimen is presented in fig. 347.

Family EPHYROPSID Claus, 1883.

Ephyropsida, Cravs, 1883, Organisation und Entwick. Medusen, pp. 23, 24; 1886, Arbeit. Zool. Inst. Univ. Wien., Bd. 7, pp. 99,
110.—VANHOFFEN, 1892, Ergeb. der Plankton Expedition, Bd. 2, K. d., p.21.—Maas, 1897, Mem. Mus. Comp. Zool. at
Harvard College, vol. 23, p. 65; 1907, Ergeb. und Fortschritte der Zool., Bd. 1, p. 211.—Bigelow, H. B., 1909, Mem.
Mus. Comp. Zool. at Harvard College, vol. 37, pp- 21, 33-

Ephyrida+ Linergide, Harcxet, 1880, Syst. der Medusen, pp. 476, 490.

FAMILY CHARACTERS.

Coronatz with 8 rhopalia (4 perradial and 4 interradial). 8 or more tentacles and 16 or
more lappets. Mouth surrounded by 4 short, simple lips. 4 perradial ostia connect the central
stomach with a wide ring-sinus. Peripherally this ring-sinus of the subumbrella gives rise to

CORONAT#—PARAPHYLLINA, PALEPHYRA. 551

16 radiating pouches in the radii of the sense-organs and tentacles; these pouches are separated
one from another by 16 septa in the radii of the lappets. These septa may be complete or
incomplete; and when incomplete there is a communication between adjacent pouches at the
bell-margin forming a peripheral ring-canal.

The Ephyropsidz are creatures of the open sea and are very widely distributed, but
are especially abundant in the tropics. In the case of Nausithoé the scyphostoma larva
bears a superficial resemblance to a branching hydroid, and it infests sponges. The ephyra
is produced by strobilization.

The genera of the Ephyropsidz are as follows:

Palephyra HarcKet, 1880 (sens. ampl.)= Ephyrat+ Palephyra+ Zonephyra Haeckel. 8 adradial tentacles, 16 lappets,
4 interradial gonads.

Nausithoé Kouuxer, 1853= Nausicaat+ Nausithoé+ Nauphanta Haeckel. 8 adradial tentacles, 16 lappets, 8 adradial
gonads. No subumbrella saccules.

Linuche Escuscuortz, 1829= Linerges+Liniscus+ Linuche Haeckel. Similar to Nausithoé, but with sac-like gastric
pouches upon the subumbrella.

Genus PALEPHYRA Haeckel, 1880.

Ephyra+ Palephyra+ Zonephyra, Harcket, 1880, Syst. der Medusen, pp. 482-484, 641.
Palephyra, VANHOFFEN, 1902, Wissen. Ergeb, deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 31; 1908,
deutsche Siidpolar Expedition, Bd. 10, Zool. 2, p. 38.

The type species is Palephyra antigua Haeckel, from the Red Sea.

GENERIC CHARACTERS.

Coronate, with 8 adradial tentacles, 8 (4 perradial and 4 interradial) marginal sense-
organs, 16 lappets, and 4 interradial gonads.

This genus is related to Nausithoé, but is more prim-
itive in that there are but 4 interradial instead of 8 adradial
genital organs as in Nausithoé. Indeed, in Nausithoé
itself the gonads begin to develop in the 4 interradii, but
later they divide and migrate into the 8 adradii.

Haeckel distinguished three genera of medusze with
8 tentacles, 8 sense-organs, 16 lappets and 4 interradial
gonads: (1) Ephyra, without lappet-pouches; (2) Pale-
phyra, with 8 cleft lappet-pouches in the ocular radii;
(3) Zonephyra, with 16 cleft lappet-pouches in the rho-
palar and tentacular radii. Haeckel, however, cut no
sections and his ideas of the structure of the gastric

cavity are probably erroneous. His E phyra is apparently
Fic. 348.—“Ephyra prometor.” After only an immature stage of Palephyra, which is in turn
Haeckel, in Das Syst. der Medusen. : : :
identical with Zonephyra.

“Zonephyra corona” Agassiz and Mayer, 1902 (Mem. Museum Comp. Zool. at Harvard

College, vol. 26, p. 157), is apparently a young Pelagia.

Palephyra antiqua Haeckel.

Ephyra prometor (young medusa), Harcket, 1880, Syst. der Medusen, p. 482, taf. 27, fign. 1, 2.
Palephyra primigenia (half-grown medusa), HAEcKr1, Tbid., p. 483, taf. 27, fign. 3-6.
Pelephyra antiqua (adult [?] medusa), Harcxet, Ibid., p. 484.

Bell 20 mm. wide, 8 mm. high. Coronal furrow and pedalia (?), 8 adradial tentacles
about half as long as bell-radius. 8 sense-organs; 16 spatula-shaped, sharply pointed lappets,
half as long as bell-radius. A long 4-sided cesophagus with folded, recurved lips. 4 inter-
radial gonads divided in the 4 perradii; each gonad crescent-shaped with the horns recurved.
6 to 8 slender gastric cirri in each interradius.

Tropical Indian Ocean near Madagascar.

Haeckel describes that which I take to be the young of this medusa as Ephyra (Archephyra)
prometor from the coast of Australia. It is only 8 mm. wide and has 4 simple, interradial,

552 MEDUS OF THE WORLD.
crescent-shaped gonads, each with only one gastric filament. His Palephyra primigenia
appears to be the half-grown medusa. There are only 2 gastric cirri in each interradius, and
the gonads and tentacles are said to be light-reddish, nearly colorless. It comes from the Red
Sea, near Tur.

Palephyra pelagica.
Zonephyra zonaria (young medusa), Harcxet, 1880, Syst.

der Medusen, p. 484, taf. 27, fign. 7, 8.
Zonephyra pelagica, Harcxet, Ibid., p. 485.

Bell 12 mm. wide, 2 mm. high, with

a coronal furrow. Mouth-tube wide and
short, hardly one-third as long as bell-
diameter (contracted [). 16 spatula-
shaped, pointed lappets, half as long as
bell-radius. 8 adradial tentacles not quite
half as long as bell-radius. 4 half-moon-
shaped interradial gonads with ends of
crescent pointing outward. Each gonad
consists of 3 swellings; the middle part
being hardly half as largeas the two lateral
ones. 10 to 12 short, gastric cirri in each
interradius. Color (?) Coast of Japan.
Haeckel describes that which may be

a young stage of this medusa under the
name of Zonephyra zonarta. It is only 8
mm. wide. The mouth tube and tentacles
are longer than in P. pelagica, but this
BiGsa4gr— Zenephyra lita tea eo may be due to conditions of contraction.

a ea The median and terminal swellings of

each gonad are all of the same size. Found off the coast of China. Haeckel’s Zonephyra
connectens (System der Medusen, p. 641), from the tropical Pacific is said to differ from those
described above in having each gonad composed of 2 swollen regions instead of 3 as in his

351-

Fic. 350.—Palephyra “ primigenia,” after Haeckel in Das Syst. der Medusen.
Fic. 351.—Palephyra indica, after Vanhoffen, in Valdivia Expedition.

CORONATH—PALEPHYRA, NAUSITHON. 553

Z. pelagica. here are also 4 lanceolate, complexly folded lips. The medusa is only 10
mm. wide and is probably an immature stage.

Palephyra indica Vanhoffen.

Pale phyra indica, VANUGFFEN, E., 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, p.32, taf. 3, fig. 10.

Bell 12 to 16 mm. wide. 8 adradial tentacles, 8 marginal sense-organs, 16 marginal
lappets. 4 large, bean-shaped interradial gonads wider than the spaces between them. About
80 simple gastric flaments. Bell white, with faint brown, radial punctations in the radii of
the sense-organs and axial lines of the lappets. Stomach brown. Gulf of Aden from a depth
of about 500 fathoms.

Vanhoffen presents a beautiful figure of this medusa, drawn from life.

Genus NAUSITHOE Kolliker, 1853.

Nausithoé, KOii1KkeR, 1853, Zeit. fiir wissen. Zool., Bd. 4, p. 323-—Gecenzaur, 1856, Zeit. fiir wissen. Zool., Bd. 8, p. 211.—
Hertwic, O., und R., 1878, Nervensyst. und Sinnesorg. der Medusen, p. 105.—Ctaus, 1883, Organisation Entwick. Medu-
sen, p. 24.—VANHOFFEN, 1892, Ergeb. der Plankton Expedition, p. 21; 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition,
Dampfer Valdivia, Bd. 3, Lfg. 1, pp. 28, 30; 1906, Nordisches Plankton, Nr. 11, p. 43.—Maas, 1904, Résult. Camp. Sci.
Prince de Monaco, fasc. 28, p. §3; 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 18.—Bicretow, H. B., 1909,
Mem. Mus. Comp, Zool. at Harvard College, vol. 37, p. 33-

Octogonia, Miter, J., 1854, Miiller’s Archiv. fiir Anat., etc., p. 97.

Ephyropsis, GEGENBAUR, 1856, Miiller’s Archiv. fiir Anat., etc., p. 239.

Nausicaa+ Nausithoé+ Nauphanta, Harcket, 1880, Syst. der Medusen, pp. 486, 487.

Scyphostoma larva:

Spongicola fistularis, Scuurtze, F. E., 1877, Archiv. fiir Mikroscop. Anatomie, Bd. 13, p. 795-
Stephanoscyphus mirabilis, ALLMAN, 1874, Annals and Mag. Nat. Hist., ser. 4, vol. 14, p- 237-
Nausithoé, Lozianco, S., und Mayer, P., 1890, Zool. Anzeiger, Jahrg. 13, p. 687.

The type species is N. punctata of the Mediterranean, Atlantic, Pacific, Indian, and
Arctic Oceans.

GENERIC CHARACTERS.

Ephyropsidz with 8 (4 perradial and 4 interradial) marginal sense-organs and 8 adradial
tentacles. There are 16 marginal lappets and 8 separate adradial gonads, the gonads isolated,
not grouped in pairs. The central stomach communicates with a wide ring-sinus in the bell
by means of 4 perradially situated ostia; the 4 interradial septa between these openings give
rise to the entodermal gastric cirri. Peripherally, the ring-sinus gives forth 16 simple, unrami-
fied pouches in the radii of the sense-organs and tentacles. The partitions between these
pouches are not complete, for there is a marginal ring-canal. No saccules on the subumbrella.

We may possibly distinguish the following “species” of Nausithoé:

N. punctata, with finely punctured, central disk without radiating furrows, large gonads, gastral filaments not grouped
into clusters. All oceans.

N. clausi, with smooth central disk, small gonads. Caroline Islands, Pacific.

N. challengeri, central disk with radiating furrows. Tristan d’ Acunha, South Atlantic.

N. albatrossi, smooth central disk, long, narrow marginal lappets. Gastral filaments grouped into clusters, with a number
of separate clusters in each interradius. Gulf of Panama, Pacific Ocean.

N. rubra, red color. Pitted central disk. Simple gastral filaments which are not grouped into clusters. Indian and
South Atlantic Oceans=N. punctata (?)

N. picta, similar to N. punctata, but with chocolate-brown or carmine gonads and blue gastric cirri= N. punctata (?).

It is apparent that there are only 4 well-marked forms of Nausithée: (1) the punctata,
rubra, picta group with pitted central lens which lacks radiating furrows, and with gastric
filaments arising singly, not in clusters; (2) N. claus: with smooth central lens; (3) N.
albatross: with gastric filaments grouped in clusters; (4) N. challengeri with radial furrows
upon the central lens.

The scyphostoma larva of Nausithoé infests sponges and bears a superficial resemblance
to a branching hydroid. See N. punctata. The medusa of Nausithoé is peculiar in having
clusters of small crystals scattered at intervals within the ectoderm of its umbrella.

{ MEDUSA OF THE WORLD.

Nausithoé punctata Kolliker.
Plate 60, figs. 4 and 5.
itho® punctata, KOuuxKeER, 1853, Zeit. fiir wissen. Zool., Bd. 4, p. 323-—Kererstein und Euxers, 1861, Zool. Beitr. Neapel,
Messina, p. 80, taf. 13, fign. 1-3—Harckex, 1880, Syst. der Medusen, p. 486.—Craus, 1883, Organ. und Entwick. der
Medusen, pp- 24, 41, taf. 6, fign. 44-46b; taf. 7, fign. 47-53; taf. 8, fign. 54-55d— Hamann, 1883, Zeit. fiir wissen. Zool.,
Bd. 38, p. 420, taf. 23 —Grarre, 1884, Arbeit. Zool. Inst. Wien., Bd. 5, p. 342 (at Trieste, Adriatic, June to Oct., rare).—
Vannorren, 1892, Ergeb. der Plankton Expedition, Bd. II, K.d., p. 13, taf. 3, fign. 8, 9; 1892, Ergeb. der Plankton Expe-
dition, Bd. 2, K. c., Nachtrag; 1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Bd. 3, Lfg. 1, p. 29; 1906, Nordisches
Plankton, Nr. 11, p. 43, fign. 2, 3 (North Atlantic 59° 39’ N. lat.); 1908, deutsche Siidpolar Expedition, Bd. 10, Zool. 2,
p- 37-—Mayer, 1goo, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 67, figs. 67, 86; plate 23, figs. 87,88; plate
26.—Acassiz, A. and Mayer, 1902, Mem. Museum Comp. Zool. at Harvard College, vol. 26, p. 155, plate 7, fig. 32.—
Bicrtow, H. B., 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 263, plate 6, fig. 21 (Maldive Islands,
Indian Ocean).—Maas, 1904, Résult. Camp. Sci. Prince de Monaco, fasc. 28, p. 54.—Browne, 1905, Report Pearl
Oyster Fisheries, Gulf of Manaar, p. 157.—Bicetow, H. B., 1909, Mem. Museum Comp. Zool. at Harvard College, vol. 37,
D. 35, plate 12.
Nau re Sinida, Gecenpaur, 1856, Zeit. fiir wissen. Zool., Bd. 8, p. 211.—Carus, 1857, Icones Zootom., taf. II, fign. 17, 22, 23.—
Herrwic, O. und R., 1878, Nervensyst. und Sinnesorgane der Medusen, p. 105, taf. 9, figs. 2, 5, 10-13; taf. 10, fig. 17.
Nausitho® punctata, var. polaris, Maas, 1906, Fauna Arctica, Bd.4, Lfg.3, p.511= Nau phanta polaris Fewkes (review of literature).
Nau phanta vettoris pisant, VANHOFFEN, 1892, Ergeb. der Plankton Expedition, Bd. 2, K. d., p. 15, taf. 3, fig. 10.
Nau phanta polaris, Fewxers, 1888, Report Lady Franklin Bay Expedition, p. 40, plate 1, figs. 1, 2, Annals and Mag. Nat. Hist.,
ser. 6, vol. 1, p. 255.
Nausithoé marginata, MetscunikorF, 1886, Embryologische Studien an Medusen, Wien, p.23 (egg); p- 37 (segmentation); p. 66
(formation of entoderm); taf. 10, fign. 1-22.
Larva SraGe:
Nausithoé punctata= S pongicola fistularis=Stephanoscyphus mirabilis, Lopianco, S., AND Mayen, P., 1890, Zool. Anzeiger,
Jahrg. 13, p. 687.
Spongicola fistularis, Scuuize, F. E., 1877, Archiv. fiir Mikroscop. Anat., Bd. 13, p. 795, taf. 45-47 (larval stage).
Stephanoscyphus mirabilis, ALLMAN, 1874, Nature, vol. 10, 251; Annals and Mag. Nat. Hist., ser. 4, vol. 14, p. 237-
Parasitic scyphostoma upon Esperia, Kowarevsky, A., 1873, Mem. Imperial Soc. Lovers of Natural History, Moscow,
vol. 10, part 2, p. 7 (Russian).

Adult medusa.—The umbrella is discoidal, flatter than a hemisphere, quite thick, and
g to 15 mm. wide. Central disk of exumbrella thick, raised, and lenticular with a finely
punctate surface, not quite half as wide as the
medusa itself and separated from the peripheral i
zone of pedalia by an annular furrow. This outer .
zone of pedalia is composed of 16 radial thicken-
ings, 8 in the radii of the tentacles and 8 in the

Nau

352- 353:

Fic. 352.—Nausithoé punctata, from life, by the author, at Naples Zoological Station, Jan. 15, 1908.
Fic. 353.— Nausicaa phaacum,” after Haeckel, in Das Syst. der Medusen.

radii of the sense-organs. They thus alternate in position with the 16 marginal lappets and
are separated by deep radial clefts in the mid-axial radii of the lappets (fig. 5, plate 60). The
8 tentacles are adradial and arise from deep clefts between the lappets. The bases of these
tentacles are set in thick, socket-like pedalia and arise from the subumbrella side of the bell.
The tentacles are about three-fourths as long as bell-diameter. The entodermal core of
each tentacle is solid and composed of highly vacuolated cells. There are 8 marginal sense-
organs, 4 radial and 4 interradial; these sense-organs are set at the bottom of 8 clefts between
the lappets, but the clefts of the sense-organs are not quite as deep as those of the tentacles.
The Hertwigs (1878, fig. 2, plate g) and Claus (1883, fig. 47, plate 7) have shown that each
sense-organ contains a distal entodermal mass of cystalline concretions, and a ventral proxi-
mal, ectodermal eye provided with lens, retina, and nerve-fibers. The 16 marginal lappets are
long, flexible, and spatula-shaped. The mouth is a simple, cruciform opening at center of

~

CORONATA—NAUSITHOB. 555

subumbrella; the 4 lips are without prominent oral appendages or palps. The central

stomach is connected with a wide annular sinus in the disk by means of 4 wide, perradial ostia,

alternating with which there are 4 short interradial septa (see Claus, 1883, p. 27, taf. 7, fig. 48).

The broad ring-sinus is interrupted near the bell-margin by 16 septa in the radii of the mid-

axial lines of the lappets. These septa are not complete, however, but leave a marginal

ring-canal. Thus the 16 peripheral stomach-pouches are in the radii of the sense-organs and
tentacles, and are joined by the marginal ring-canal in the axial line of each lappet.

Four groups of simple, unbranched, gastric cirri are upon the interradial septa of the
central stomach and project centripetally inward into the central stomach; altogether there
are about 28 gastric cirri, about 7 in each group. The 8 adradial gonads occur in the 8 ten-
tacular radu Bid are upon the floor of the subumbrella in the zone of the wide, inner ring-
sinus of the bell. Each gonad is large and globular and consists of a pocket-like fold of the
entoderm of the subumbrella (see Claus, 1883, p. 31). A zone of well-developed circular
muscle-fibers is on the subumbrella between, and centrifugal to, the gonads. Centripetal
to this, powerful strands of radiating muscle-fibers extend outward to the tentacles and mar-
ginal lappets; also, 8 poorly developed strands of radiating muscles extend outward in radial
aad interradial positions from the base of the cesophagus to the zone of circular muscles.

The color of this medusa 1s quite variable. The gelatinous substance of the bell 1s usually
translucent-milky, greenish, or light brownish. The gonads are brownish or red or, in the case
of the males, bright yellow. Rosin-colored spots in the ectoderm of the exumbrella, especially
upon the lappets, are due to clusters of small crystals (see Claus, 1883, fig. 44, taf. 6).

A young ephyra of this species was found by us near Flamingo Key, Bahama Islands,
February 9, 1893. It was 2 mm. in diameter, and there were as yet no marginal tentacles
and only 4 gastric cirri. A slightly older ephyra fas been figured
by Claus, 1883, fig. 48, taf. 7.

This males ¢ is a surface form, and is common in the Med-
iterranean, Atlantic, Pacific, and Indian Oceans, and in all trop-
ical or warm seas. Nausithoé polaris (Nauphanta polaris
Fewkes) fromthe Arctic Ocean, appears to be identical with
N. punctata, and if this be true Vanhoffen is right in stating
that Nausithoé punctata is found in all oceans. It is abundant
in summer at Tortugas, Florida, and in the Bahamas, but has
not been found on the Atlantic coast of the United States north
of the Carolinas. Vanhoffen, 1906, describes a specimen 12 mm.

., _ wide from N. lat. 59° 39’, W. long. 8° 49’.

(Ne Wi Cg Hamann, 1883, studied the development of the ephyra of
after Maas,in Mem. Mus.Comp. ‘ uf
Zoology at Harvard College. this species and finds that the gonads first appear as 4 inter-

radial entodermal swellings in the subumbrella wall of the
stomach, at a time when the ephyra has but one gastral filament in each interradius. Later
the 4 original gonads divide and migrate so as to become 8 in the adradii of the subumbrella
wall of the Gite The genital products originate in the entoderm and migrate into a gelat-
inous space between two layers of entoderm. Pie spermaries appear as a series of follicles
in this space.

According to Metschnikoff, 1886, who has studied the early development of Nausithoé
“marginata” (which is apparently identical with N. punctata) the eg gg is citron-yellow, 0.23
mm. in diameter, and is laid in mid-day in December; segmentation is total but somewhat
unequal, the cells 6 the v egetativ e pole being largest. A aides central, cleavage cavity is formed
and the gastrula results from invagination at the hinder end of the larva. The blastopore
then closes over and the entoderm is entirely inclosed by a layer of ciliated ectoderm, and
the free-swimming planula is thus formed.

The remarkable scyphostoma larva of Nausithoé punctata bears a striking superficial
resemblance to a hydroid and it lives commensal within sponges such as Suberites, Myxilla,
Reneira, Esperia, etc. It is especially abundant at Trieste and Naples. This hydroid-like
larva forms a branching tree-like growth within the body of the sponge, the polyp- -mouths
and their tentacles projecting out of the oscula of the sponge. The branching, tree-like stock
of the larva is covered with an irregularly annulated, chitinous perisarc, which terminates
at a short distance below the zone of tentacles of each polypite. The mouth of the polypite

556 MEDUSA OF THE WORLD.

is at the extremity of a short, blunt-conical hypostome, which is surrounded by an annulus
of about 40 solid filiform tentacles. 4 longitudinal partitions lined by entoderm extend
throughout the cavity of the stem. These do not meet in the center, but form only partial
septa, comparable with the mesenterial partitions of other scyphostome of Scyphomeduse.
There is no marginal ring-canal. There are external, longitudinal and internal (meso-
dermal) circular muscles. The polypites are translucent-white.

Lobianco and Paul Mayer, 1890, found that ephyra of Nausithoé arise by strobilization
from this larva. The young ephyra has only 4 gastric filaments and no tentacles. Kowal-
eyski, 1873, also observed the giving off of the ephyra, but did not determine that they were
Nausithoé.

It is not surprising that this peculiar larva should have received various names: Allman
calls it Stephanoscyphus mirabilis; F. E. Schulze describes it in detail under the name S pong-
icola jistularis; but its true nature was discovered by Lobianco and Paul Mayer, 1890.

Haeckel’s Nausicaa pheacum from Corfu, Mediterranean, may be identical with N.
punctata, but the 8 gonads tend to be grouped in 4 interradial pairs, forming a broken crescent
in each interradius, with a wide separation between the outwardly directed horns of each
crescent. It may have been described
from an abnormal or young specimen
of N. punctata (See Haeckel, 1880, Sit-
zungsber. Jena. Gesell. fiir Med. und
Natur., Jahrg. 1880, Feb. 20.)

Nausithoé clausi Vanhdffen.

Nausithoé clausi, VaANHOFFEN, 1892, Ergeb. der Plank-
ton Expedition, Bd. 2, K. d., p. 14, taf. 4, fign.
Is 25

Disk about 9 mm. wide; central
lens-shaped dome of exumbrella flat,
smooth, unpitted, and without radial fur-
rows; 5 mm. wide, 16 well-developed
marginal pedalia. Medusa 3 times as
wide as high. 16 very blunt, 3-cornered
marginal lappets, three times as wide as
long and hardly one-ninth as long as
bell-radius. 8 adradial tentacles with
well-developed, conical bases. Tentacles
as long as bell-radius. 8 marginal sense-
organs alternating with tentacles. 8 gonads in the tentacular radii, very small, spherical,
only 1.3 as wide as the pedalia. Ring-muscle of subumbrella one-third as wide as _bell-
radius. Numerous, small, simple gastric cirri arising in a linear row in each interradius.
Color (?)

Pacific Ocean east of the Caroline Islands. A single specimen appears to be N. punctata
with poorly developed marginal-lappets and small gonads.

Fic. 355.—Nausithoé rubra, after Vanhoffen, in Valdivia Expedition.

Nausithoé challengeri Vanhiffen.

Nau phanta challengeri, Harcket, 1880, Syst. der Medusen, p. 487; 1881, Report Deep-sea Medusex, Challenger Exped., Zool.,
vol. 4, p. 103, plates 27, 28, 20 figs. 4
Nausithoé challengeri, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped. Valdivia, Bd. 3, Lfg. 1, pp. 28, 31.

Bell 12 mm. wide. Central lens of exumbrella separated by a deep annular furrow from
zone of pedalia. Somewhat less in diameter than bell-radius, its margin cleft by 16 radiating
furrows which do not extend to center of exumbrella. Marginal zone of pedalia well developed,
the 8 ocular being narrower than the 8 tentacular. Tapering tentacles somewhat longer than
bell-radius. The 8 large gonads are twice as long as wide and are elongated outwardly.
They are somewhat wider than the intervals between them. 4 interradial clusters of simple
gastric cirri which arise at equal spaces in a single row ir each cluster and are not grouped
into brushes as in N. albatrossi. Each cluster has about 24 cirri.

CORONATH—NAUSITHOE, LINUCHE. 557

Found near the island of Tristan d’Acunha, South Atlantic, at a depth of 1,425 fathoms,
in an open net, on March 16, 1876.

Nausithoé albatrossii Vanhoffen.
Nauphanta albatrosii, Maas, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. 83, taf. 14, fign. 1-3
Nausithoé albatrisst, VANEOFF EN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Bd. : Lig. I, pp- 28, 30.

Disk 35 to 40 mm. wide, 35 mm. high. The marginal lappets are narrow and elongate
with nearly parallel sides and rounded ends; not heart-shaped, as in Nausithoé challengert or
N. vettoris prsant (=N. punctata). Central disk smooth and without a notched margin, such
as is seen in N. challenger. Stomach, especially the lips, much shorter than in either NV. chal-
lengeri or N. vettoris pisant. 8 gonads, elongate, oval, bladder-like organs. The gastric fila-
ments arise in a row in each interradius, each row being composed of about 4 clusters, of 5
filaments each. All 5 filaments of each cluster arise close together, and with wide, free intervals
between the groups. This species is remarkable for its large size.

Gulf of Panama, Pacific coast of Central America.

Nausithoé rubra Vanhoffen.
Nausithoe rubra, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Bd. 3, Lfg. 1, p. 39, taf. 1, fign. 4, 5 -BicrLow,

H. B., 1909, Mem. Museum Comp. Zool. at Harvard College, vol. 37, p. 36, plate 12, fig. 6.

Bell 15 mm. in diameter. With large pits over the exumbrella surface of the central
disk. Simple gastral filaments arise separately, not in clusters. § large gonads. 16 short,
pointed, marginal lappets. 8 tentacles longer than the bell-radius. Tentacles and bell red-
purple, stomach blue. Indian and South Atlantic Oceans. It appears to be distinguished
from N. picta by its deep color and large pits over the exumbrella surface of the central disk;
moreover, fees to Bigelow, 1909, the rhopalia in N. rubra lack ocelli, while there is a
ventral ocellus in N . punctata. The septal nodes of N. rubra are broadly triangular and the
4 perradial gastric ostia are narrow. The marginal canal-system is as in N. punctata.

Nausithoé “picta” Agassiz and Mayer—N. punctata (?)
Nausithoé picta, Acassiz, A., and Mayer, 1902, Mem. Museum Comparative Zool. at Harvard College, vol. 26, p. 154, plate 7,

fig. 33.—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 19, taf. 1, fign. 5-8

Bell 15 to 22 mm. wide, somewhat flatter than a hemisphere. Coronal furrow of exum-
brella well-developed, central disk of exumbrella smooth-edged. 16 marginal lappets, wide
and pointed. 8 tapering adradial tentacles, with wide bases, are somewhat shorter than
bell-radius. 8 marginal sense-organs, radial and interradial in position with reference to the
lips. 8 large, egg-shaped or oval gonads project from middle zone of subumbrella in the
tentacular radii. Mouth cruciform, the lips being in the radii of 4 of the marginal sense-
organs. 4 clusters of gastric cirri in the interradii, each cluster consisting of about 12 cirri.

The gonads are chocolate-brown to carmine and the gastric cirri are blue. The ocelli
of the 8 sense-organs are dark-brown.

This medusa was found by the Albatross in the Paumotos Islands, South Pacific, in
September, 1899. Later, 29 specimens were found by the Szboga in the Malay Archipelago.
Maas, 1903, describes these specimens and fortunately corrects certain serious mistakes in the
description written by me in the account published under the names of Agassiz and Mayer.
I am inclined to regard this medusa as merely a large, highly-colored variety of N ausithoé
punctata.

Genus LINUCHE Eschscholtz, 1829.

Linuche, Escuscuoutz, 1829, Syst. der Acal., p. 91.
Linerges+ Liniscus+ Linuche, Harcxer, 1880, Syst. der Medusen, pp. 495, 496, 498, 642.
The type species is Linuche unguiculata Eschscholtz, 1829, of the West Indian region,
tropical Atlantic.
GENERIC CHARACTERS.

Ephyropside with 8 (4 perradial, 4 interradial) rhopalia. 8 adradial tentacles. 16
lappets. 8 gonads grouped in 4 pairs close to the 4 perradii. With zones of hernia-like sacs
upon the floor of the subumbrella. The 16 peripheral stomach-pouches break up into
branches in the lappets. A marginal ring-sinus may or may not be present.

558 MEDUS OF THE WORLD.

There are, I believe, but two species, L. ungurculata of the tropical Atlantic and L. aquila
of the Pacific and Indian Oceans. These are closely related, but in the Atlantic form
there is no marginal ring-canal, whereas this is present in the Pacific species; moreover, the
subumbrella warts of the Pacific form are arranged in 2 rows and in the Atlantic medusa in
3 rows.

Linuche unguiculata Eschscholtz.
Plate 59, figs. 1 to 10.

Medusa unguiculata, Scawartz, 1788, Neue Abhandl. Schwed. Acad. Deutsche Uebers., 1789, p. 195, taf. 6, fig. 1.—Linn&é
(Gren), 1788, Syst. Nature, tomus 1, pas. 6, p. 3159.

Pelagia unguiculata, PERon et Lessuer, 1809, Annal du Muséum Hist. Nat., Paris, tome 14, p. 349.

Linuche unguiculata, Escuscuotz, 1829, Syst. der Acal., p. 91. —Brainvitte, 1834-1836, Manuel d’ Actinologie, p. 289, planche

Crees pease pegasus+Liniscus ornithopterus (?) +L. sandalopterus (?) +L. cyamopterus+Linuche unguiculatat
L. vesiculata, HArcKet, E., 1880, Syst. der Medusen, pp. 495, 497, 498, 499, taf. 29, fign. 4-6.

Linerges mercurius, FewxKes, 1882, Bull. Mus. Comp. Zool. at Harvard College, vol. 9, p. 259, plate 2, figs. 3-5; plate 3, figs. 4-8,
11, 13; plate 4, figs. 3-22; 1886, Report Commiss. Fish and Fisheries for 1884, p. 950.—Acassiz, A., 1888, Bull. Mus.
Comp. Zool. at Harvard College, vol. 14, p. 186, fig. 93.—Mayer, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol.
37, p- 68.—Conk in, 1906, Year Book of Carnegie Institution of Washington, No. 4, p. 115; 1908, Papers from Tortugas
Laboratory Carnegie Institution of Washington, vol. 2, p. 153, 8 plates (development).

Bell about 13 mm. high and 16 mm. wide. Lenticular apex flat and horizontal, separated
from the vertical sides by a distinct but shallow annular furrow. Occasionally a few irregu-
larly arranged, radiating clefts are found in the margin of the lenticular apex of the bell,
but this is usually plain. Sides of bell composed of 16 vertical pedalia, similar each to each,
and separated one from another by 16 clefts in the radii of the mid-axial lines of the lappets.
Thus the pedalia are in the radii of the tentacles and rhopalia and alternate with the lappets
(plate 59, fig. 2).

The 16 lappets are bluntly oval with rounded edges and are all inclined inward at an
angle such that when one observes the medusa by looking down upon the aboral end of the
bell the animal rotates with the hands of the watch as it swims through the water, upon each
contraction of its margin. The lappets being inclined as are the vanes of a wind-mill cause
this peculiar spinning on its axis as the medusa progresses rapidly along. This was discovered
by Prof. E. G. Conklin in 1905. 8 small, simple, marginal sense-organs, perradial and inter-
radial in position, arise from clefts between the lappets and are not protected by covering
scales. The entoderm of each rhopalium contains a spherical mass of concretions. No
ocelli. The 8 adradial tentacles are small, neither very flexible nor contractile, and only
about 1.5 times as long as lappets.

The 8 (4 double) gonads form 4 cleft crescents on both sides of the perradial lines of
the subumbrella, the cleft being in the perradius itself and the horns of the crescents extending
outward toward the margin of the bell. These gonads begin to develop as 8 separate sacs
diverging outwardly on either side of the 4 perradii when the ephyra i is about 5 mm. in diameter.
The subumbrella sacs are not male gonads as was conjectured by Haeckel, and the medusa
is not hermaphroditic, the sexes being separate.

The proboscis is urn-shaped, 4-sided, and with 4 slightly recurved lips with their perradial
angles truncated so as to present a nearly octangular appearance when viewed upon looking
into the bell-cayity. The mouth does not extend to level of bell-margin, but is usually at
about two-thirds the distance down from the inner apex of the bell-cavity. There are 4 cres-
centic interradial rows of simple unbranched gastric cirri, about 15 to 20 in each row at the
interradial septal nodes. Beyond these, and connected with the central stomach by 4 perradial
ostia, is the broad bell-sinus, which in turn gives rise to 16 radiating pouches in the radii of the
sense-organs and tentacles. The edges of these pouches break up into numerous, ragged-edged
branches in the lappets, but I am unable to find any marginal ring-canal, for although I have
often injected the lappet-pouches with air, carmine, or other stains, each pouch is evidently
completely separated from the two adjacent to it. This appears the more remarkable from
the fact that Maas has found a marginal ring-canal in the Pacific species; a fact which I am
enabled to confirm in specimens from the Philippine Islands, and Claus, Vanhéffen and Bige-
low have demonstrated that such a structure exists in other Ephyropside.

Projecting from the floor of the subumbrella into the bell-cavity are 48 hollow sac-like
or wart-shaped protuberances which arise from the radial stomach-pouches and are arranged

PLATE 59.

Figures 1 to 10 are of Linuche unguiculata; figure 11 is of Linuche aquila.

Fig.

Fig.

Fig.

nN

g. 6.

= 9;

. 10. Side view of an ephyra 3 mm. in diameter, showing the beginning of

“BI

. Side view of mature medusa. Key West, Florida, June 1, 1897.
. Side view of bell of mature medusa showing sculpturing of the exum-

brella. Tortugas, Florida, May 12, 1906.

. Oral view of an ephyra 2 mm.in diameter. Nassau Harbor, Bahamas,

March to, 1893.

. Marginal sense-organ of the ephyra shown in figure 3.
. Oral view of an ephyra 7 mm. in diameter, showing the 4 pairs of

gonads beginning to develop on both sides of each perradius, and
masses of brown cells. Ragged Islands, Bahamas, April 5, 1907.
The mouth-parts of the ephyra shown in figure 5. Showing gonads

and gastric cirri.

. Oral view of mature medusa, showing the subumbrella saccules.

Tortugas, Florida, May 8, 1907.

. Oral view of a mature medusa, showing the 16 radial pouches of the

central stomach. There is no marginal ring-canal.
Marginal sense-organ of a mature medusa.

the tentacles. Nassau Harbor, Bahamas, March 14, 1893.

. Linuche aquila, Wailangilala Atoll, Fiji Islands, South Pacific,

November 20, 1897.

Drawn from life, by the author.

PLATE 59
MAYER

AHcens Ca.

CORONATA—LINUCHE. 559

in 3 zones (text-fig. 3564). The 2 inner rows are each composed of 8 large perradial and
interradial saccules which lie between the gonads, the 8 innermost saccules being partially
cleft and bean-shaped (plate 59, fig. 7); a third row of 32 smaller saccules, 2 for each lappet-
pouch, lies at the level of the outer ends of the horns of the crescentic gonads. In the sub-
umbrella we find a broad, unitary, marginal area of ring-muscles, and centripetal to this are
more or less isolated strands of radial muscle-fibers.

The entoderm of the gonads, of the wart-like saccules, and of numerous, separated,
more or less polygonal areas of digestive cells in the gastric pouches is brown. 8 rows of
dark-brown spots extend longitudinally down the inner surface of the lips.

Vast numbers of the ephyre of this medusa appear among the Bahama Islands and
West Indian region in February and March, and become mature from April until early June,
usually disappearing at or about the middle of May. I have seen hundreds of such swarms,
all composed of but one species. I have never seen any of the so-called “species” described
by Haeckel from the West Indian region, and it appears to me that he has constituted species
out of preserved material displaying various well-known characteristics of abnormal con-
traction and in various stages of growth. In fact I have seen Haeckel’s “species” only in
medusz which I have myself preserved. They appear‘not to exist except in alcohol.

In the Bahama-Florida region in spring these meduse form swarms, miles in extent,
filling the water with myriads of brown thimbles, all actively spinning clockwise as they
progress through the water.

Conklin, 1908, has studied the habits and early development of Linuche unguiculata.
When mature the medusz rise in vast swarms to the surface and the eggs are discharged and
fertilized at 8 p.m. The female gonads are slaty or blue-gray in color while those of the male
are brown. When the gonads have been emptied the medusz sink down to the bottom and
die. Each egg 1s closely invested by a very thin transparent membrane which persists to the
gastrula stage. The eggs are laid near the surface but gradually sink downward. The newly
laid egg is 0.24 mm. in diameter. It consists of a peripheral layer of clear protoplasm, an
intermediate layer of densely packed yolk spherules, and a central sphere composed appar-
ently of dissolved yolk. The peripheral layer becomes the peripheral layer of the blastula and
gastrula, and gives rise to the cilia of the ectoderm. The intermediate “shell” of closely
crowded yolk spherules constitutes the principal part of all of the cells of the gastrula and
blastula, while the central mass of dissolved yolk is poured into the cleavage cavity and probably
serves as a source of nourishment for the surrounding cells.

Two polar bodies are formed. The first two cleavages are meridional and cut downward
from the animal (polar body) pole to the vegetative pole, and the third cleavage is equatorial.
Up to the 64-cell state the divisions are wholly mitotic. Cleavage is total and practically equal.
When the embryo consists of about 1,000 cells protoplasmic processes appear over the entire
periphery, and these push off the egg membrane and form the vibratile cilia. Gastrulation
usually takes place by invagination of the small, rounded cells of the vegetative pole of the
embryo; but sometimes there is a unipolar ingression of cells at the vegetative pole and no
invagination. The blastopore closes so that the entoderm becomes entirely incased within
the ectoderm, and the larva elongates and becomes a free-swimming planula.

Isolated blastomeres, at least as late as the 4-cell stage, may give rise to apparently normal
free-swimming larve.

The center of the egg is more nearly fluid than the peripheral layers, and this fact favors
the cutting inward of the cleavage furrow from the animal pole to the vegetative during the
first two divisions, and the unilateral constriction in the third (equatorial) division.

When the ephyra is 1.5 mm. in diameter, it has 16 lappets and 8 sense-organs, but neither
tentacles nor gonads. 4 pastric cirri, one im each interradial side of stomach. Disk very flat,
brown in color (fig. 3, plate 59). When 3 mm. wide the tentacles begin to develop, and the
gonads appear when the medusa is about 5 mm. wide. The polygonal areas of pigmented
digestive cells then develop in an inner ring of 16 large areas centrifugal to the gonads, and
still farther out an outer annulus of 32 areas. There are also irregularly shaped and arranged
areas of brown cells in the lappets (plate 59, fig. 5). The subumbrella saccules do not develop
until later.

560 MEDUSZ OF THE WORLD.

Linuche aquila.
Plate 59, fig. 11-

Linerges aquila, Hance, 1880, Syst. der Medusen, p. 496.—Goerre, 1886, Sitzungsber., Akad. Wissen., Berlin, Jahrg., p. 833.—
~ Acassiz, and Mayer, 1899, Bull. Museum Comp. Zool. at Harvard College, vol. 32, p. 170, plate 10, figs. 33, 34; 1902,
Mem. Jbid., vol. 26, p. 156.
(?) Linerges draco, HarckEL, loc. cit., p. 496.
Linerges draco (young medusa), Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 24, taf. 1, fign. 1, 2.

This form is widely distributed over the tropical Pacific and is closely related to the
tropical Atlantic L. ungurculata, with which it is identical in form and dimensions, being
about 13 mm. high and 16 mm. wide. It has 48 wart-like protuberances upon the subumbrella
arranged in 2 rows instead of in 3 as in the Atlantic medusa. 8 of the subumbrella sacs in
L. aquila alternate with the gonads and 8 arise from the sides of the gonads themselves.
Thus in the Pacific medusa we have two zones of protuberances, an inner zone of 16 large
sacs, and an outer of 32 small subumbrella saccules. The 16 large sacs lie in the mid-regions
of the gonads, while the 32 small saccules lie at the zone of the outer ends of the gonads.
The areas of brown cells are developed only centrifugal to the zone of gonads in the Atlantic,
while they occur between the gonads as well as beyond them in the Pacific medusa. A marginal
ring-canal is present.

O

o@,e°

Fic. 356.—A, arrangement of subumbrella warts in Linuche unguiculata of the
tropical Atlantic. B, arrangement of subumbrella warts of Linuche aquila
of the tropical Pacific. C, enlarged view of central part of subumbrella of
Linuche aquila, showing lips, gonads, and saccules.

Vast swarms of these meduszw are found among the atolls of the Fiji and Paumotos
Islands, and they extend westward to the coast of Africa. They abound in the spring months,
in Fiji in December and at Singapore in April. I have studied a large collection of these
medusz taken in the Philippine Islands at Mactau, near Sibu, on April 6, 1908, by the U.S.
Bureau of Fisheries steamer Albatross. All were mature.

There appear to be no valid distinctions in Haeckel’s descriptions between L. aguila
and his “1. draco,” the differences being such as one would expect to find in two contracted
preserved specimens. Haeckel’s Linantha lunulata (Syst. der Medusen, p. 494, taf. 29, fign.
I to 3) is possibly the young of L. aguila. It is said to have 4 interradial horse- -shoe-shaped
gonads, but in all known species of Linuche the gonads are more nearly perradial than inter-
radial. It is evidently an immature form, being only 10 mm. wide, and has no subumbrella
saccules; indeed, the figure itself shows its ephyra- -like condition. It comes from the Gala-
pagos Islands, off the Pacific coast of South America.

CORONATZ—ATOLLA. 561

Family COLLASPIDZ Haeckel, 1880.

Collaspide, HaEcKet, 1880, Syst. der Medusen, p. 488.—VANHGFFEN, 1906, Nordisches Plankton, Nr. 11, p. 44.
Atollide, Bicetow, H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, pp- 21, 37-

FAMILY CHARACTERS.

Coronate with numerous (more than 8) marginal sense-organs which alternate with an
equal number of tentacles. Marginal lappets twice as numerous as the tentacles.

Atolla is the only known genus.

Genus ATOLLA Haeckel, 1880, sensu Fewkes.

Atolla+ Collaspis, Harcke, 1880, Syst. der Medusen, p. 488; 1881, Deep-Sea Meduse, Challenger Report, Zool., vol. 4, p. 111.

Atolla, Fewxes, 1886, Report Commiss. Fish and Fisheries U. S. A. for 1884, p. 934; (?) Ephyroides, 1885; Ibid. for 1883,
p- 597; and 1886, p. 948; 1889, p. 532-—VANHOFFEN, 1892, Ergeb. der Plankton Expedition, Bd. 2, K. d., p. 16; 1902,
Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p- 5; 1906, Nordisches Plankton, Nr. 11,
p- 44.—Maas, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, No. 1, pp. 78, 79; 1899, Bull. Soc. Zool. de
France, p. 165; 1904, Résult Camp. Sci. Prince de Monaco, fasc. 28, p. 48; 1903, Scyphomedusen der Siboga Expedition,
Monog. 11, p- 14; 1907, Ergeb. und Fortschritte der Zool., Bd. 1, p. 195.—Browne, 1908, Trans. Roy. Soc. Edinburgh,
vol. 46, p. 240.—BiceLow, H. B., 1909, Mem. Mus. Comp. Zool. at Harvard College, vol. 37, p- 37-

GENERIC CHARACTERS.

Coronate with numerous (g or more) tentacles and equally numerous marginal sense-
organs. Twice as many marginal lappets as sense-organs. 8 adradial gonads and 4 inter-
radial subgenital ostia. 4 lips. The tentacles and marginal sense-organs alternate regularly,
but the insertions of the tentacles and their pedalia are higher up on the exumbrella than are
the insertions of the pedalia of the sense-organs.

The type species is wyville: Haeckel.

Haeckel distinguished “A tolla” with 8 gonads grouped in 4 pairs and “Collaspis” with
8 separate, equally spaced gonads. As was first shown by Fewkes, 1886, both of these con-
ditions may exist upon one and the same medusa, some of the gonads being paired, others
separated by equal spaces. The name “Collaspis” must therefore be dropped, being equiva-
lent to Atolla.

The medusz of this genus are deep-sea forms and are inhabitants of all oceans, and
large numbers of them have been taken in open nets dragged at 350 to 2,500 fathoms. A few
specimens have also been found upon the surface, but this is unusual. They are often brought
up from depths of about 100 fathoms where the ocean is more than 2,000 fathoms deep.

In the sculpturing of the exumbrella, the structure of the tentacles, the general features
of the anatomy of the gastrovascular system, and in the arrangement of the muscular system,
these medusz are strikingly similar to Per:phylla but differ in the absence of funnel cavities
in the floor of the subumbrella and in the flat, discoidal shape of the bell. The central stomach-
cavity is thus less complex, but not essentially different from that of Periphylla.

The details of the structure of gastrovascular system, sense-organs, gonads, etc., are
given in the description of 4. bairdit.

Ephyrotdes rotaformis Fewkes, 1886 (Report U. S. Fish Commission for 1884, p. 949),
may be closely related to Atolla. The 16 to 32, or more, pedalia are separated by wide intervals
instead of by narrow clefts, as in Atolla. Several specimens are mentioned by Fewkes as
having been dredged from depths of 389 to 1,555 fathoms in the Gulf Stream off the coast
of the United States, but were not sufficiently well preserved to admit of description.

Vanhéffen, 1902, discovered in Atolla, a system of excretory openings which consists of
8 pores, 2 in each principal radius near the perradial angles of the stomach and centripetal
to the zone of the gonads, with which this system has no connection. The position of these
openings is marked by 8 oval spots upon the floor of the subumbrella. The closely allied
E phyropside have numerous openings on the subumbrella at the place of connection between
the tentacular canals and the ring-canal, but these have not been observed in Afolla.

Vanhiffen (1902), Maas (1904,) and Bigelow (1909), have given the best detailed descrip-
tion of the genus 4tolla. The development is unknown.

562 MEDUS#® OF THE WORLD.

Bigelow concludes that the sculpturing of the central lens of the exumbrella, the presence
or absence of warts, etc., constitute the best criteria for the separation of species. The furrows
of the central lens are, when present, always one less in number than the tentacles. Bigelow
doubts the existence of 4. alexandr: and finds slight furrows on the central disk of the specimens
which I described from Hawaii as 4. alexandri. He therefore merges “A. alexandri” with
A. wyuviller.

A study of the large number of specimens of 4tolla in the National Museum at Washington
convinces me that 4. bairdii and A. wyviller are very closely related if not mere individual
variations of one and the same species. In some specimens the annular ridge in the outer side of
the ring-furrow is so narrow and indistinctly separated from the marginal zone of pedalia that
one hesitates to decide whether to consider it to be present or absent. Moreover the margin
of the central lens in all Atollas is apt to be more or less notched with faint sinuosities in its
outline, and thus 4. barrdiz and A. wyville: may be only varieties of one and the same species.

8 so-called species of Atolla have been described, but I believe only three can be distin-
guished upon non-intergrading characters. These are:

A. bairdi: Fewkes, with an annular ridge within the ring-furrow between the central lens and
the marginal zone of pedalia. Exumbrella surface of lappets smooth. 4. valdivie and
A. gigantea are probably identical with 4. barrdi.

A. chunt Vanhoffen has the exumbrella surface of lappets besprinkled with papilla, very wide
central lens, and an annular ridge in ring-furrow.

A. wyviller Haeckel, with margin of central lens notched with radial furrows; no annular ridge
in the ring-furrow. This is probably identical with 4. achillis and A. verrillit.

A synopsis of the distinctive characters of the forms is presented in the following table:

Synopsis of the So-called Species of Atolla.

| A. bairdii*

lens.

Annular ridge on
the outer side of
the ring furrow.

Radical furrows
of central lens.

Number of ten-
tacles.

Exumbrella sur-
face of lappet
zone.

Where found.

more than half
as wide as me-
dusa.

Smooth with
even margin.

None; but the
margin is in-
dented with 17
to 23 notches.

18 to 24

Smooth.

| North Adantic. |

medusa.

As in A. bairdii.

As in A. bairdii.

20 to 29

Smooth.

Indian Ocean.

Notched in outer
margin. Con-
tracted ?

As in A. bairdii.

Smooth.

Gulf of Panama.
Panama.

Hidden within

the ring-furrow.

Notched with 23
shallow radial
furrows.

24+

Covered with
papille.

Cape of Good
Hope, Africa.

Hidden within
the ring-furrow.

Notched with 15
to 31 radial
clefts or grooves,

16 to 32

Smooth.

Antarctic.
Tropical Pacific.

A. valdivie* | A. gigantea* | A. chuni. A. wyvillei= A. verrillii-f
| A. achillis->
Width of bell in | 12 to 72 132 | 150 27 to 50 58 to 66 95
mm.
Width of central) 5 to 40, usually | Half as wide as | 70 22.5 to 38 About 29 to 33.| About so.

Hidden within
ring-furrow.

Notched with
13 to 27
shallow radial
clefts.

14 to 28

Smooth.

Atlantic, Indian,
and Pacific
Oceans.

#These are probably all A. He

fThese are probably 4. wyvillei.

To a great extent the conditions exhibited by these so-called species of Atolla intergrade.

For example we can draw no sharp line of distinction throughout the series between a smooth

b

central lens with faint notches in its margin and as is seen in 4. bairdi1, and a lens deeply
furrowed by radiating valleys as in 4. wyviller. Similarly the wide, annular ridge, which pro-
jects prominently on the outer side of the ring-furrow in A. bairdi grades insensibly to the
condition seen in 4. wyville: wherein the ridge is so narrow that it is quite hidden by the ring-
furrow which over-arches it. With the exception of 4. chuni, which stands apart, all of the
others may be mere local races or varieties of one variable species, 4. wyvillet.

CORONATA—ATOLLA. 563

Atolla bairdii Fewkes.

Atolla bairdi:, Fewxes, J. W., 1886, Report Commiss. Fish and Fisheries U. S. A. for 1884, p. 936, plates 1-3, 4 figs.; 1888,
American Journ. Sci., ser. 3, vol. 35, p. 172; Ann. Mag. Nat. Hist., ser. 6, vol. 1, p. 253; 1889, Report Commiss. Fish
and Fisheries U. S. A. for 1886, p. 530.—Acassiz, A., 1888, Bull. Mus. Comp. Zool. at Harvard College, vol. 15, p. 132,
fig. 427.—VANHGFFEN, 1892, Ergeb. der Plankton Expedition, Bd. 2, K. d., p. 16, taf. 4, fign. 3-9; 1902, Wissen. Ergeb.
deutsch. Tiefsee Exped., Dampfer Valdivia, Bd. 3, Lief. 1, p. 9; 1906, Nordisches Plankton, Nr. 11, p. 44, fign. 4, 5.—
Maas, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. 79; 1904, Résult. Camp. Sci. Prince de Monaco,
fasc. 28, p.49, planche 4, figs. 29-34; planche 5, figs. 38-43.—Browne, 1906, Trans. Linnean Soc. London, ser. 2, Zool., vol.
10, p. 179.

The medusa is 30 to 144 mm. in diameter, disk quite flat. Central lenticular part of
exumbrella somewhat more than half as wide as entire medusa. It is flat aborally and sur-
rounded by a recurved rim, the periphery of which displays about 20 to 22 slight notches,
which are one less numerous than the tentacular pedalia in whose radii they tend to lie. Central
lens separated from peripheral part of exumbrella by a deep, broad, annular groove, slightly
over-arched by the recurved rim of the central lens itself. The peripheral zone of the exum-
brella includes all that part of bell which lies centrifugal from the annular groove. It may
in turn be divided into 4 zones: (1) The innermost zone is a simple, smooth-surfaced, annular
ridge bounded on the inner side by the deep annular groove and on the outer by a very slight,
reddish-colored groove which separates it from (2), the zone of the tentacular pedalia. There

Fic. 357.—Atolla bairdii, after Maas in Résult. Camp. Sci. Albert 1° Prince de Monaco.
A, and B, side views of the medusa; C, marginal canal-system and ring-muscle (cut across).

are about 18 to 24, usually 22, pedalia, one in the radius of each tentacle. These pedalia are
thick ridges separated one from another by shallow radiating furrows, which do not cut very
deeply into the gelatinous substance, so that the tentacular pedalia form a thick, continuous,
gelatinous zone. Each pedalium bears a short, solid tentacle, usually carried recurved upward
and provided with strong, longitudinal muscles upon its subumbrella side.

The third zone is that of the pedalia of the sense-organs and lies immediately centrifugal
to the zone of the tentacular pedalia. These pedalia of the sense-organs alternate in position
with the equally numerous, tentacular pedalia. They are lower than the tentacular pedalia
and are separated from one another by deep, wide, radiating furrows which occupy the radii
of the tentacles. Sense-organs very small with 2 swollen regions upon the ventral (subum-
brella) side of the bulb where one finds thickened, sensory ectodermal epithelium. There
is no ocellus and there are no pigment spots either in the ectoderm or the entoderm, but
there is a terminal sac-like swelling which contains a crystalline mass of entodermal origin
and which is protected by a wide, scale-like expansion on the exumbrella side of the sense-
club. (See Vanhoffen, 1902; Maas, 1904.)

The fourth or outermost zone of the exumbrella consists of long, slender, marginal lappets
supported by the pedalia of the sense-organs, and which are twice as numerous as the latter.

564 MEDUS OF THE WORLD.

The center of the subumbrella is occupied by the shallow proboscis, the 4 lips of which
are simple and cruciform. Surrounding the base of the proboscis is a zone in which are
situated the 8 adradial gonads. The gonads vary considerably in size and shape, but normally
appear to be composed of bean-shaped sectors, each genital ridge being constricted radially
by one or more deep furrows (see Vanhoffen, 1892, taf. 4, fig. 4, g). 8 groups of radial-muscles
lie in the floor of the subumbrella between the 8 gonads. Immediately centrifugal to the
zone of the gonads lies a zone of circular muscle-fibers which is divided into separate parts
by furrows which lie in the radii of the sense-organs. Beyond this zone lies the outermost
ring of circular muscles developed so remarkably that it projects outward from the floor of
the subumbrella as a thick annular mass, which is entire, not divided, as are the inner, circular
muscles. The cavity of the central stomach communicates with a broad ring-sinus in the
subumbrella by means of 4 perradially situated ostia. This ring-sinus corresponds in position
to the zone of the gonads upon the subumbrella. Centrifugally it gives rise to a radiating vessel
in each tentacular radius and also to an equal number of vessels to the sense-clubs. The ten-
tacular vessels each give off a pair of side-branches which lead into the rhopalar yessels near
the bases of the sense-clubs. At the point of origin of each tentacular vessel there is a region
of fusion of the subumbrella and exumbrella walls of the gastrovascular cavity thus forming
a small cathammal plate in the middle of each tentacular vessel (see fig. 359 C).

Fic. 358.—Avolla valdivie, after Vanhoffen, in Valdivia Expedition. View of subumbrella.

The excretory system of the exumbrella has been described under the genus Atolla.

In the corners of the central stomach are 4 interradial rows of gastric cirri forming a
cross with perradial axes. Altogether the gastrovascular system, the pedalia of the exumbrella,
and the structure of the muscular system, with the exception of the remarkable unitary ring-
muscle of Atolla, are quite alike in both Atolla and Periphylla and indicate a genetic rela-
tionship between the two forms.

The gelatinous substance of the bell of Atolla bairdit is translucent and milky-blue in
color. The central disk of the exumbrella is sprinkled over with rust-red colored patches
which become more numerous in the region of the annular furrow, which is of a decided red-
brown color. The powerful centrifugal circular-muscle band of the subumbrella is dark-red
and a radial red streak extends inward on the subumbrella side from the base of each sense-
organ to the band of circular muscles. Gonads and entoderm of central stomach red. The 8
adradial excretory pores near the 4 angles of the central stomach are dark-red of the hue
so commonly seen in deep-sea medusz of all ports.

CORONATH—ATOLLA. 565

This species is widely distributed over the North Atlantic. It has often been obtained
in open nets dragged at depths from goo to 2,000 fathoms, but has also been taken occasionally
upon the surface, especially upon the Gulf Stream off the Carolina coast. It is a creature of
the deep ocean, not found in bays or estuaries, but is evidently a deep-sea medusa that may
only occasionally and under exceptional conditions come to the surface.

When young the 8 gonads are grouped in 4 interradial pairs, but in later life they swing
into adradial positions. The number of antimeres does not alter with age; but in some
specimens one finds some small tentacular pedalia inserted between larger rhopalar ones.
Vanhoffen records Ir specimens ranging from 5.5 to 114 mm. in diameter and with 18 to 24
tentacles. These were found by the Valdivia in the Gulf of Guinea, west coast of Africa.

Atolla bairdii forma valdivie Vanhdéffen.

Atolla valdivie, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lief. 1, p. 13, taf. 1,
fig. 3; taf. 5, fig. 21; taf.6, fign. 41-46; taf. 7—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 17,
taf. 1, fign. 3, 4; taf. 3, fig. 23; taf. 12, fig. 108.

Medusa 132 mm. in diameter. Central disk only half as wide as bell, smooth, without
radial furrows. 20 to 2g tentacles. The 4 interradial septal nodes of the subumbrella are
wider than in the typical 4. bairdi1. g specimens were found by the Valdivra in the Indian
Ocean. Maas records g specimens obtained by the S:boga in the Malay Archipelago. These
ranged from 12 to 82 mm. in diameter, the central disks ranging from 6 to 42 mm. in width;
and the antimeres range from 20 to 29 without reference to the size of the medusa. These
medusz were obtained from depths ranging from about 280 to 1,000 fathoms (see fig. 358).

This form is very closely related to 4. bairdi: but the central disk is only half as wide
as the medusa, and the 4 septal nodes are wider than in 4. bairdit.

The Albatross obtained 3 specimens of this medusa in the Philippine Islands on April ro,
1908, at dredging station No. 5202. The characters and dimensions, in mm., of these specimens
are as follows:

Specimen | Specimen | Specimen |
A. B. c.
Exumbrella: | |
Dream car @ Ween oocceeccbeccspocno connote taoocepe>oaderodeoeae 45-5 38 37
Diameter of central lens of exumbrella............-.-2.-00-eeeeeee 21.5 19 18.5
Width of annulus between coronal furrow and tentacular pedalia...... 1.5 1.25 I
Wadthyorstentacnlars pedaliajorc cists = clcleicifeleeicie siete oir ss niels ie) e.einle + ace 4 Res: 3-25
Length of tentacular pedalia (in radial direction)................. 3 2.75 3
Width of ocular pedaliass. 2... c. ence ce cence cence senses Bos 3 Da |
Length of ocular pedalia (in radial direction).......-........-...--5 TS) 5 5-75
Subumbrella: |
Diameter to outer side of ring-muscle..............-.---- se se0 555 || 43 35 34
\Viiatin Gimn panied Es 60 enoaoqesede pp peas Oaseancobocuaocedorisce: 2.25 3-5 2.5
Diameter, to) outer, edge of gonads...- 2.2.0 000. sews etme cman ecsere 31 20.5 23
DY ame berg obs art anne su rele fe asaya esate (ate late lele)o) ele wl ctse= eie=ieie/a elie’ se violet» 15 13 14
Length (circumferential) of gonad.........2...2+:222.0-cseeeeeeeeneees 7 Zink fol
Width (radially) obs pomad tees risers Sie eivie i= = 21 sialslaielateiel> aisle melee oie 6 3-5 3
General characters:
Numberjofitentaclessncje2s-c2.4 22+ ols Paths se aea aes cio ae ss 24 22 24

Atolla gigantea Maas =A. bairdii (?)

Atolla gigantea, Maas, 1897, Mem. Mus. Comp. Zool. at Harvard College, vol. 23, p. 80, taf. 12, fign. 2-4; taf. 13, fign. 7-9;

taf. 14, fig. 6.

This form resembles 4. valdivia, but the outer edge of the annular ridge in the ring-furrow
is notched, not simple and entire as in 4. bairdi1 and A. valdivie. Thus the central lens is half
as wide as the bell, as in 4. valdivie. Ring-furrow wide, and peripheral to it there is the
notched, annular ridge, and beyond this is fhe: zone of pedalia and lappets. Tentacular pedalia
somewhat shorter and wider than the rhopalar pedalia. Medusa is 150 mm. wide. Num-
ber of tentacles (?) Gulf of Panama, Pacific coast of Central America (see fig. 359).

Maas is uncertain as to whether or not the margin of the central lens is plain or notched.
Radial furrows of the ridge in the ring-furrow may be due to contraction in preservative fluids.

566 MEDUSA OF THE WORLD.

Atolla chuni Vanhéffen.
Atolla chuni, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lief. 1, p. 12, taf. 1,

figs. 1, 2; taf. 5, fig. 26.—Browne, 1908, Trans. Roy. Soc. Edinburgh, vol. 46, p. 240.

Bell 27 to 50 mm. wide, 9 to 15 mm. high. Central lenticular disk 14 to 27.5 mm. wide
with 23 faint radial furrows at the margin. Annular furrow 0.5 to 1.75 mm. wide. Zone
of pedalia 2.25 to 3.5 mm. wide. 24 tentacles. Species distinguished by 7 to g small, pearl-
colored, papilla-like protuberances over the exumbrella surfaces of each marginal lappet;
commonly with one papilla in the center and the others in two lateral rows. 2 specimens found
by the Valdivia off Cape of Good Hope, Africa, November 18, 1898; and 1 by the Scot-
tish Antarctic Expedition, in a trawl at 1,332 fathoms, in the same region.

\B' >:
Le L
ae aeaaee&

fi

Fic. 359.—Atolla gigantea, after Maas, in Mem. Mus. of Comp. Zool. at Harvard College.
Fic. 360.—Atolla chuni, after Vanhoffen in Valdivia Expedition.
Fic. 361.—Atolla wyvillei, 0.75 natural size, drawn by the author, from a specimen in the National Museum, Washington.

Atolla wyvillei Haeckel.

Atolla wyvillei, Haecxer, 1880, Syst. der Medusen, p. 488; 1881, Report, Challenger Expedition, Zool., vol. 4, p. 113, plate 29,

figs. 1-9.—Rou te, 1896, Résult. Scientifique Camp. Caudan, Tome 1, Lyon, p. 302 (Bay of Biscay, France, depth of 350

to 850 fathoms).—Maas, 1897, Mem. Museum Comp. Zool. at Harvard College, vol. 23, p. 79 —VANHGFFEN, 1902,

Wissen. Ergeb. deutsch. Tiefsee Expedition Valdivia, Bd. 3, Lfg. 1, p. 13, taf. 5, fig. 22; 1908, deutsch. Siidpolar Expe-

dition, Bd. 10, Zool. 2, p.37—Brownge, 1908, Trans. Roy. Soc. Edinburgh, vol. 46, p. 241.—Bicetow, H. B., 1909, Mem.

Museum Comp. Zool. at Harvard College, vol. 37, p. 39, plates 8-10.

(2) Collaspis achillis, Harcxet, 1880, loc. cit., p. 489.
Atolla alexandri, Maas, 1897, Mem. Museum Comp. Zool. at Harvard College, vol. 23, p. 81, taf. 11, fig. 2; taf. 14, figs. 4, 5—

Acassiz, A., and Mayer, 1902, Mem. Museum Comp. Zool. at Harvard College, vol. 26, p. 156.—Mayer, 1906, Bull.

U. S. Fish Commission, vol. 23, p. 1138, plate 2, fig. 7; plate 3, figs. 10, 11.

This species is characterized by the numerous, wide, radial notches or furrows in the
margin of the central lens of the exumbrella. These are much wider and deeper than in 4.
verrillii. Exumbrella surface of lappets smooth, not beset with papilla as in 4. chuni. This
medusa is probably identical with Collaspis achillis Haeckel, but in the latter the furrows of
the central lens are represented as deep, narrow clefts, whereas in 4. wyville: they are shallow
notches which vary greatly in prominence in individual meduse. Moreover, in 4. wyviller
the pedalia are short and broad, while in 4. achillrs they are long and narrow. The central
lens and the pedalia are separated only by a ring-furrow and there is no prominent ridge periph-
eral to the ring-furrow such as is seen in 4. barrdit. The medusa becomes 73 mm. wide and
there are usually about 22 to 28 tentacles. The bell is flatter than a hemisphere. Found in the
Antarctic and Southern Atlantic and Pacific. The Albatross obtained it in the Philippine

Islands, tropical Pacific.

CORONATAS—ATOLLA. 567

Tam inclined to believe that 4. wyvillet, A. verrillit, A. alexandri, and A. achillis are only
varieties of one and the same species. “A. alexandri” isa form of A. wyviller with indistinct
notches in the margin of its central lens.

A specimen was dredged by the Albatross on April 10, 1908, dredging station No. 5201,
off the south end of Leyi Island, Philippine Islands, from a depth of 554 fathoms. A side
view 1s shown in fig. 361. “There were 23 pedalia and tentacles, and 22 radial furrows in the
margin of the general disk. “The dimensions in millimeters are as follows:

Diameter of bell.............. 55 Width of tentacular pedalia..... 6 Width of ring-muscle.......... 3

Thickness of belli .5:..<(<.015 << -s 26 Length of pedalia of lappets..... 6.5 Diameter across zone of gonads.. 28.5

Diameter of central Jens....... 40 Width of pedalia of lappets...... 5 Diameter of manubrium....... 21

Depth of coronal furrow....... 3 Length of tentacles............ 13 Length of manubrium......... 22

Length of tentacular pedalia.... 6.5 Diameter to outer edge of ring- Gonads 7 long in the circumferential direc-
DU BCLG a tatals aioszistaieystetsloratclo 48 tion, 5 wide in the radial direction.

Atolla wyvillei forma verrillii Verrill.

Atolla verrilliz, Verritt, 1885, Report Commiss. Fish and Fisheries U.S. A. for 1883, p. 594-—Fewkes, 1885, Report Commiss.
Fish and Fisheries U. S. A. for 1883, p. 596; Ibid., 1886, Report for 1884, p. 939, plates 4, 5, 4 figs.; Ibid., 1889, Report
for 1886, p. 530; 1888, American Journ. Sci., ser. 3, vol. 35, p- 172; 1888, Ann. Mag. Nat. Hist., ser. 6, vol. 1, p. 253.—
Maas, 1897, Mem. Museum Comp. Zool. at Harvard College, vol. 23, p. 79.—VANHOFFEN, 1902, Wissen. Ergeb. deutsch.
Tiefsee Exped., Valdivia, Bd. 3, Lief. 1, p. 10, taf. 5, fig. 23; taf. 6, fig. 39; taf. 7; 1908, deutsche Siidpolar Expedi-
tion, Bd. 10, Zool. 2, p. 36.

This form is distinguished from the typical 4tolla hairdi and resembles 4. wyville: by the
absence of a projecting annular ridge on the outer side of the ring-furrow, the numerous,
fine, radial furrows of its central dome, the long, narrow pedalia of the sense-organs, and the
smallness of its marginal lappets. The medusa becomes 95 mm. wide and has between 14 to 28,
usually 22, tentacles and marginal sense-organs and 28 to 56 small lappets. Central lens of
exumbrella wider than in 4. bairdit. It is found in deep water in the Atlantic, Indian, and
Pacific Oceans. Most of the specimens have been obtained in open nets dragged from depths
of 373 to 2 ,369 fathoms; but several have been found upon the surface. It is probably only
a variety of A. wyvillet.

Family ATORELLIDZ Vanhoffen, 1902.

Atorellide, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition Valdivia, Bd. 3, Lfg. 1, p. §t.—Bicrtow, H.B., 1909,
Mem. Museum Comp. Zool. at Harvard College, vol. 37, pp. 21, 30.

FAMILY CHARACTERS.

Coronatze with 6 rhopalia.

There is but a single known genus, this being Atorella.

The relationship between the Atorellida and other Coronate is unknown; for in the
Periphyllida, Paraphyllinidz, and Ephyopside there are 4 or 8 thopalia, and in the Collaspide
a large (more than 8) but indefinite number of these organs; and it is impossible at present to
determine how the number 6 may have been derived in the Atorellida. It is probable, how-
ever, that its affinities are much closer to 4folla than to the other Coronate.

Genus ATORELLA Vanhioffen, 1902.

Atorella, VANHGFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 33-—Maas, 1903,
Scyphomedusen der Siboga Expedition, Monog. 11, p. 10.—B1GEtow, H.B., 1909, Mem. Museum Comp. Zool. at Harvard
College, vol. 37, p. 30.

The type species is Atorella subglobosa Vanhoffen, of Dar es Salaam, Africa, and from
the Malay Archipelago.

GENERIC CHARACTERS.

Coronatz with 6 marginal sense-organs, 6 tentacles, 12 marginal lappets, and 12 pedalia.
A coronal furrow is present, and the 12 pedalia alternate with the lappets. There is a poorly
developed ring-muscle in the subumbrella. 4 lips, 4 interradial gonads.

568 MEDUS OF THE WORLD.

Atorella subglobosa Vanhdffen.

Atorella subglobosa, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Valdivia, Bd. 3, Lfg. 1, p. 33, taf. 3, fig. 11.—
s AAS, 1903, Scyphomedusen der Ssboga Exped., Monog. 11, p. 10, taf. 3, fign. 16-18.—Bigelow, H. B., 1909, Mem. Mus.
Comp. Zool. at Harvard College, vol. 37, p. 30.

Bell globular, 15 to 17 mm. in diameter. 6 mainly solid, tapering tentacles, about as long
as the bell-radius. 6 marginal sense-clubs, each with a ‘terminal mass of concretionary
crystals, arise from yery shallow niches in bell-margin. There are 12 wide, shallow, slighty
cleft marginal lappets. Ring-furrow on exumbrella not very deep. Genel disk of exum-
brella more than twice as wide as zone of pedalia. The 12 pedalia alternate with the lappets
and are separated one from another by shallow furrows. Throat tube 4-sided, mouth cruci-
form. ‘There are 4 clusters of gastric filaments, each cluster consisting of about 20 filaments.
‘The ring-sinus gives rise to 12 pouches in the radii of the tentacles and sense-organs and there
is a marginal ring-canal as in Atolla or Pertphylla. 4 interradial, sac-like, swollen gonads
arise from floor of subumbrella beyond the zone of the
gastric filaments. Muscular system of subumbrella quite
similar to that of Nausithoé, but the ring-muscle is very
poorly developed.

Stomach and gastric filaments brown, gonads yel-
lowish-brown. Subumbrella muscles white, all other
parts translucent.

Vanhoffen describes a specimen from Dar es
Salaam, east coast of Africa, and Maas describes
another from the Malay Archipelago. Our description
is derived mainly from that of Mass, his specimen being
the more perfect. Vanhoffen’s figure shows 6 gonads.

Atorella vanhoffeni Bigelow.

Atorella vanhoffeni, Bicetow, H. B., 1909, Mem. Museum Comp. Zool. at

Fic. 362.—Atorella subglobosa, after Maas in Harvard College, vol. 37, p. 30, plates 1, 11, and 12.

Siege Eapelicen: Bigelow had three specimens. In two of these the
bell was 5 mm. high and 6 mm. wide, and one was 3 mm. high and 7 mm. in diameter. The
ring furrow is a deep cleft. The entire exumbrella surface is besprinkled with wart-like, nema-
tocyst-bearing prominences, thus being very different from the smooth surface of the bell of
Atorella subglobosa. The 6 tentacles are each about as long as the bell-diameter. They taper
outwardly but each terminates in a knob-like tip, instead of having simple, pointed ends as
in A. subglobosa. The 6 rhopalia closely resemble those of Atolla, but the exumbrella surface
of its covering scale is covered with thickened ectoderm, not with a thin layer as in Atolla.

There is a large lithocyst and ventral bulb, but no ocellus. There are 12 long, oval, marginal

Fic. 363.—Atorella vanhoffent, after H. B. Bigelow, in Mem. Mus. Comp. Zool. at Harvard College, 1909.

CORONATAS—ATORELLA., 569

lappets. The stomach is flat and shallow and the 4 lips are short and thickened. The 4
interradial septal nodes are narrow and the perradial ostia wide. The gastric cirri are
arranged in 4 interradial groups, each group arising from a stout gelatinous stalk, and consist-
ing of 80 to 100 filaments. The canal-system of the bell resembles that of Periphylla. The 4
perradial ostia of the central stomach lead into a wide ring-sinus, which gives rise on its outer
side to 12 broad radial-canals in the radii of the tentacles and sense-organs. These radial-
canals branch at their ends and unite to form a marginal festoon canal. The ring-muscle of
the subumbrella is very weak, but the subumbrella plates at the bases of the tentacles are
very prominent.

There are 4 gonads, each being a leaf-shaped body folded so as to leave a deep groove
along its middle line on the inner surface, and this groove causes the gonad to appear as if
double, although this is not truly the case for each gonad is attached along a single line. The
gonads are orange-yellow, all other parts colorless. This is a surface species. Bigelow records
it from off the Pacific coast of Panama.

Order SEMAEOSTOMEZ L. Agassiz, 1862.

Semaeostomea, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp- 9, 159-

Semostoma, HAEcKEL, 1880, Syst. der Medusen, p. 499.

Semaostomata, VANHGFFEN, 1888, Bibliotheca Zoologica, Heft. 3, pp. 6,21; 1906, Nordisches Plankton, Nr. 11, Acraspede, p. 45-
Semaeostoma, Maas, 1907, Ergeb. und Fortschritte der Zool., Bd. 1, p. 200; 1906, Fauna Arctica, Bd. 4, Lfg. 3, p. 504.
Discomedus@ (in part), Harcket, 1880, Syst. der Medusen, p. 450.

CHARACTERS OF THE SEMAEOSTOME#,

Scyphomedusze without a coronal furrow and without pedalia. With a simple, central
mouth-opening, the 4 perradial angles of which are developed into large curtain-like or gelat-
inous lips. With hollow tentacles and marginal rhopalia. The gonads are in sac-like folds of
the entodermal wall of the subumbrella. Without interradial septal nodes in the stomach.

The families of the Semaeostomez are as follows:

(1) Pelagide Grecennaur, 1856. The central stomach gives rise to completely separated, unbranched radiating pouches.
No ring-canal. Tentacles arise from the bell-margin between the clefts of the lappets.
(2) Cyaneidz Acassiz, L., 1862. The central stomach gives rise to branched but completely separated radial-canals.
No ring-canal. Tentacles arise from the floor of the subumbrella.
(3) Ulmaride Harcxet, 1880, sens ampl. The central stomach gives rise to simple or branched radial-canals which
are put into connection one with another by a marginal ring-canal.
A. Subfamily Umbrosidii. The tentacles arise singly from the clefts between the marginal lappets at the bell-
margin. Protrusive, sac-like gonads, without subgenital pits.
B. Subfamily Sthenonidii. The tentacles arise in linear clusters from the floor of the subumbrella.
C. Subfamily Aurelidii. The tentacles and lappets arise from the sides of the exumbrella above the margin.
Invaginated gonads with subgenital pits.

Family PELAGIDA Gegenbaur, 1856.

Pelagide, GeGENBAUR, 1856, Zeit. fiir wissen. Zool., Bd. 8, pp. 210, 267.—Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, pp
121, 163.—Acassiz, A., 1865, North Amer. Acal., p. 47-—HaeckEt, 1880, Syst. der Medusen, p. 499.—von LENDENFELD,
1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 265.—C1aus, 1886, Arbeit. Zool. Inst. Univ. Wein., Bd. 7, p. 110.—
Acassiz anv Mayer, 1898, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 1-—Craus, 1883, Organisation und
Entwick. Medusen, p. 24.—VANHOFFEN, 1906, Nordisches Plankton, Nr. 11, p. 45-

FAMILY CHARACTERS.

Scyphomeduse with 8 or 16 marginal sense-organs, 4 perradial and 4 interradial, and,
when present, 8 adradial. 8 or more tentacles which arise singly from the clefts between the
marginal lappets. 16 to 64 marginal lappets. The mouth is simple and cruciform, and is
situated at the extremity of an cesophagal tube, the 4 perradial corners of which are produced
to form 4 long mouth-arms, the free edges of which are complexly crenulated. The simple,
lenticular, central stomach gives rise to completely separated, radiating pouches the centrifugal
ends of which give rise to simple, unbranched lappet-pouches. There is no ring-canal. The
gonads occupy 4 interradial folds in the wall of the subumbrella. In some cases they project,
but they are usually sunken, forming 4 pits in the floor of the subumbrella.

570 MEDUS® OF THE WORLD.

The medusa of this family are readily distinguished from the Coronate by the absence
of a coronal furrow and by the remarkable development of the 4 perradial corners of the
mouth, which extend outward as 4 long palps or mouth-arms, carrying the free edge of the
lips along with them in double curtain-like fringes. Also the exumbrella of the Pelagidz is
smooth and displays none of the complex sculpturing seen among the Coronate; finally, the
gastrovascular system of the medusa of the Pelagide is simpler than in the Coronate, for
the central stomach is without interradial fusions of its upper and lower walls, and it con-
sists merely in a wide, lenticular, central space which gives off completely separated, radiating
pouches in the radii of the tentacles and sense-organs.

The Pelagidz are also closely related to the Cyaneidz. In the Cyaneide, however, the
tentacles arise from the floor of the subumbrella at some distance in from the bell- -margin,
whereas in the Pelagidz they arise from notches between the marginal lappets. Also the
tentacles of the Cyaneidz are usually grouped in clusters, while in the Pelagide they arise
singly. A still further distinction lies in the fact that the radiating pouches of the stomach
are simple in the Pelagidz, while in the Cyaneida they give forth numerous blindly ending,
non-anastomosing canals, which enter the marginal lappets.

Medusz of the genus Pelagia develop directly from the planula without going through a
sessile scyphostoma stage. The planule of Chysaora and Dactylometra, however, attach
themselves and develop into Scyphostomz, which in the case of Chysaora is known to strobi-
late and produce a number of ephyre.

The Pelagide are of world-wide distribution, but are most abundant in the tropical
regions. Many of them congregate in great swarms in bays and estuaries, and none are known
to ie permanently at great depths.

The development of Pelagia has been studied by L. Agassiz (Cont. Nat. Hist. U. S.) and
by A. O. Kowalevsky, 1874 (Memoirs of the Imperial Society of the Friends of Natural
History, Anthropology and Ethnography of Moscow, vol. 10, p. 7, plate 3 [Russian text]).
A synopsis of the genera of the Pelagidz follows:

Pelagia PEron anv Lesueur, 1809. With 8 marginal sense-organs. 8 tentacles alternating with 8 marginal sense-
organs, 16 marginal lappets.

Chrysaora Péxon anv Lesueur, 1809. With 8 marginal sense-organs. (3 X8) 24 tentacles, 3 between each successive
pair of marginal sense-organs. 32 marginal lappets.

Dactylometra LL. Acassiz, 1862. With 8 marginal sense-organs. (5X8) 4o tentacles, 5 between each successive pair
of sense-organs. 48 marginal lappets.

Kuragea Kisutnouye, 1902. (7X8) 56 tentacles, (8 X8) 64 lappets.
Sanderia Gortre, 1886. 16 marginal sense-organs, 16 tentacles, 32 cleft Jappets.

Genus PELAGIA Péron and Lesueur, 1809.

Pelagia, PERON ET ae rR, 1809, Ann. Mus. Hist. Nat. Paris, tome 14, p. 349.—Escuscuoxtz, 1829, Syst. der Acalephen,
p- 72.—Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 163.—Haecxet, 1880, Syst. der Medusen, p. 504.—VaAn-
HOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 6,21; 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Valdivia,
Bd. 3, Lfg. 1, p. 34; 1906, Nordisches Plankton, Nr. 11, 14, 5—Maas, 1904, Résult. Camp. Sci. Prince de Monaco,
fasc. 28, p. 56; 1906, Revue Suisse de Zool., tome 14, p. 100; 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 29.

GENERIC CHARACTERS.

Pelagide with 8 adradial tentacles, alternating with 8 rhopalia. With 16 marginal
lappets. 16 radiating stomach-pouches in the rhopalar and tentacular radii, each of which
ends in 2 side branches in the marginal lappets. No ring-canal.

The type species is Pelagia noctiluca of the Mediterranean, first described by Forskal,
1775, aS Medusa noctiluca. At least 14 so-called “species” of Pelagia are known, 1 from the
Mediterranean, 1 from the Mediterranean and Atlantic, 6 from the Atlantic, 5 from the
Pacific, and 1 from the Indian Ocean. They are more abundant in warm or torrid regions,
but one species is found in Behring Sea and another near the Cape of Good Hope. All of
the Atlantic species are closely related one to another, and future researches may demonstrate
that they are only geographical races. In fact the distinctions between “ species” have been
largely determined upon preserved material, and some of them may be separated upon un-
natural conditions of contraction due to the effects of preservation; thus Vanhdffen, 1888,
distinguishes a number of “species” upon the folding and wrinkling observed in the exum-
brella warts of preserved medusa. At present the “species” are in almost hopeless confusion,
as will appear from the following table based largely upon Vanhoffen’s work. Indeed it

PLATE 60.

Fig. 1. Pelagia noctiluca, mature medusa, 1.25 times natural size. Naples
Zoological Station, November 29, 1907.

Fig. 2. Pelagia noctiluca. Aboral view of a marginal sense-organ.

Fig. 3. Pelagia noctiluca. Side view of one of the nettling-warts near the
mid-radius of the exumbrella.

Fig. 4. Nausithoé punctata. Oral view of mature female seven times the
natural size. Tortugas, Florida, April 17, 1906.

Fig. 5. Nausithoé punctata. Side view of mature medusa showing sculptur-
ing of exumbrella. Tortugas, Florida, June, 1906.

See page 553 for figures 4 and 5.

Drawn from life, by the author.

PLATE 60

MAYER

AHoen& Co

SEMAEKOSTOMEAS—PELAGIA,.

571

seems probable that the foldings observed by Vanhéffen in the nettle-warts are largely due
to shrinkage in alcohol.

The medusz of Pelagia, being pelagic in all stages, are creatures of the high seas; and
one would expect the species to be of world-wide distribution and at the same time to have
developed many local varieties which are not very clearly differentiated from their parent
stocks. I believe that all of the forms may be grouped into 4 cohorts as follows:

(1) P. noctiluca, neglecta, and crassa, of the Mediterranean and Atlantic, with large, elongate nettle-warts over the

exumbrella.

(2) Allied to (1) in the Pacific, we find P. flaveola, denticulata, tahitiana, and papillata with large oval, erect nettle-warts.

(3) P. cyanella, perla, discoidea, phosphora and minuta of the Atlantic with small, rounded nettle-warts.

(4) P. panopyra and placenta of the Pacific with small, flatly dome-like nettle-warts.

Cohorts (1) and (2) are closely related; and (3) and (4) form another group.
As in Cyanea and Aurellia so in Pelagia we find that the Linnean system 1s inadequate
to express the relationship of the numerous, closely related forms.

Synopsts of the Forms of Pelagta.

P. cyanella

P. denticulata

Color.

Variable. Yellow-
brown to reddish-
brown. Gonads
and tentacles us-
ually dark-red.
Mouth-arms yel-
lowish. Nettle-
warts reddish-
brown.

coast of Brazil and
Florida to Cape
Cod, in Gulf
Stream.

Variable. Usually
blue-violet to pale
blue. Nettle-warts
reddish-brown.
Tentacles reddish-
purple.

Variable. Varying
from light rose-red
to violet-red.
Mouth-arms more
violet. Gonads
more purple.
Nettle-warts
violet.

Paumotos
Islands, tropi-
cal Pacific.

Light-yellowish.
Tentacles citron-
yellow. Gonads
brownish-yellow.
Nettle-warts
tipped with
orange.

P. noctiluca P. panopyra P. flaveola P. discoidea
Peron et Lesueur.*| Péron et Lesueur.t} Péron et Lesueur. | Eschscholtz= Eschscholtzf. Brandt. A

P. tahitiana variety of P.
Agassiz and flaveola ( ? )
Mayer.

Width of disk in} 60 50 50 16 to 30 70 to 80 60

mm.

Height of disk | 32 40 30 8 to 15 15 to 20 50

in mm.

Character of Large, elongate, | Small, roundish, | Small, round, Very large, No warts. Ex- | Large, elon-

nettle-warts on | elliptical, with thick-set. Found | elliptical, with thick-set, and umbrella gate over entire’

exumbrella. cross-foldings. only in middle longitudinal fur- | egg-shaped. smooth ? exumbrella.

zone of disk. row and cross- Especially
foldings. thick at apex.
Shape of mar- | Square-cornered, | Twice as wide as | Quadratic. Rounded, double,| Flat and cleft so| Quadratic.
ginal lappets. | quadratic. high. Outer edge twice as wide as} as to be double.
convex. long.

Length of ror r— r ar r — O.5 7

mouth-tube in

terms of disk

radius.

Length of art ar ar r 3r 4r

mouth-arms in

terms of disk-

radius (r).

Where found. | Mediterranean. Pernambuco, Tropical Pacific. | Coast of Japan,| Cape of Good | Behring’s Sea,

Hope, Africa.

Light-reddish.
Palps rose-red.
Tentacles pur-
ple-red. Gonads
whitish.

Aleutian
Islands.

Light violet-
red. Tentacles
red. Nettle-
warts brown.

*Development without alternation of generations.
{Development through a pelagic larva without alternation of generations.
{Too imperfectly known to be retained.

The remarkable development, which is direct, without a sessile larval stage, is described
in detail under P. noctiluca and P. cyanella. The young medusa passes through a 4-tentacled
stage, before acquiring its 8 adradial tentacles.

“Zonephyra corona,” Agassiz and Mayer, 1902 (Mem. Museum Comp. Zool. at Harvard
College, vol. 26, p. 157, pl. 4, figs. 19, 20), is probably a young Pelagia.

or
ca |
bo

MEDUS OF THE WORLD.

Synopsts of the Forms of Pelagia—Continued.

P. perla Haeckel. | P. phosphora P. placenta P. neglecta P. crassa P. minuta
A variety of P.| Haeckel* Haeckel. A Vanhoffen=a | Vanhdffen. Vanhoffen.
cyanella (?) variety of P. variety of P. =P. phos-

panopyra. noctiluca. phora (? )

Width of disk | 50 to 60 40 to 50 40 53 to 60 35 12 to 25
in mm.

Height of disk | 40 to 50 25 to 30 12 23 to 28 13 3 to 6
in mm.

Character of Numerous, Small, rounded, | Numerous, thick-| Large, rounded | Large, flat, ellip-/ Rounded,
nettle-warts on} rounded, small, | withlongitudinal| set but flat and | to elliptical. tical with a small, thick-
exumbrella. and flat. furrow and cross-| small. Surface | Without longi-| longitudinal set, without

foldings. None | nodular. tudinal furrow,| furrow and in-| longitudinal
on upper third | but with cross-| distinct cross- | furrows, but
of exumbrella. foldings. foldings. None| with very
at bell-margin.| thick cross-
furrows.
Shape of mar- | Quadratic, with | Rounded, almost | Twice as wide as} Quadratic. Twice aswide as| Wider than
| ginal lappets. | concave outer semicircular. high. Flatly high. high.
edge. rounded.

Length of 0.337 0.57 0.57 — 0.5r 0.66r
mouth-tube in
terms of disk

| radius.
Length of 3r ar 2r 2.597 2r ar
mouth-arms in
terms of disk-
radius (r).
Where found. | Atlantic coast of | Tropical parts of | Tropical Pacific, | Mediterranean | In middle of Coast of
Europe. the Atlantic. West} South America, | and coast of | tropical Atlan-| Brazil, Per-
coast of Africa. | Philippines. Africa. tic. nambuco.

Color. Variable. Orange | Variable. Rose- | ? ? ? ?

| to rose-red or | red to violet-red
flesh-colored, or | or purple. Arms,
with rusty-yel- | ribs, tentacles,
low flecks. and gonads
Nettle-warts usually darker
orange. Tenta-| red than the bell.
cles and gonads
red.

*Development through a pelagic larva without strobilization or alternation of generation, Haeckel, 1867.

Griffiths and Platt, 1895 (Nature, vol. 52, p. 564), find that the violet pigment of Pelagia
has the composition C,, H,, NO,. It is soluble in alcohol, ether, and acetic acid, and especially
soluble in CS,. Insoluble in water. It gives no characteristic absorption bands. It is thus
quite distinct from the blue coloring matter of Hydromedusz as determined by Colasanti, 1888
(Centralblatt fiir Physiol., Bd. 2, p. 10).

Pelagia noctiluca Péron and Lesueur.
Plate 60, figs. 1 to 3.

Medusa noctiluca, ForsxKAt, 1775, Descript anim. itin. orient., p. 109.

Pelagia noctiluca, PEnon ev Lesueur, 1809, Annal. du Mus. Hist. Nat., tome 14, p. 350; P. purpura, Aurellia phosphorica,
loc. cit., pp. 350, 358.—Kroun, 1855, Miiller’s Archiv. Anat. Physiol., p. 491, taf. 20 (development).—Harcket, 1880,
Syst. der Medusen. p. 505 (list of authors and names).—Kowatevsky, 1873, Mem. Imp. Soc. Lovers of Nat. Hist.,
Moscow, vol. 10, part. 2, p.7, plate 3 (development).—Hamann, 1883, Zeit. fiir wissen. Zool., Bd. 38, p. 422, taf. 32 (develop-
ment and structure of gonads).—Mertscunikorr, 1886, Embryol. Studien an Medusen, Wien., p. 24 (egg); 67 (segmenta-
tion); 100 (larva); taf. 10, fign. 23-28.—Monaco, Prince of, 1887, Comp. Rend. Paris, tome 104, p. 452 (swarming habits
of the medusa).—Vanunrren, 1888, Bibliotheca Zoologica, Heft. 3, p- 8, taf. 1, fign. 5, 6; taf. 6, fign. 1-5; 1908, Deutsch.
Stidpolar Expedition, 1901-1903, Bd. 10, Zool. 2, p. 38.—Gorrtre, 1893, Zeit. fiir wissen. Zool., Bd. 55, p. 659, 11 fign.,
taf. 30-31; 1893, Sitzungsber. Akad. Wissen. Berlin, p. 853 (development).—Scuaxet, 1910, Zool. Anzeiger, Bd. 35, p.
407 (histology of odgenesis).

The following is a description of a typical, adult specimen from the Bay of Naples:
Disk somewhat higher than a hemisphere when contracted, but flatter than a hemisphere

SEMA EOSTOMEA—PELAGIA, (8)

when expanded. In ordinary contraction it is about 49 to 55 mm. in diameter and 31 mm,
high. Sides of bell relatively straight and sloping, the apex flat. Numerous nettle-warts
over the exumbrella, arranged in more or less irregular lines radiating from aboral apex of
exumbrella. These warts are rich orange-red in color and are elongate and linear, some-
times with, but more often without, cross-foldings. Near the bell-margin, however, they
lose their linear shape and become small, simple, and more or less oval.

The 8 marginal sense-organs are set in deep niches in the perradii and interradii. The
sense-club has no ocellus, but contains only a terminal mass of deeply pigmented orange-colored
crystalline concretions of entodermal origin. There is no sensory pit in the ecunibrella above
the sense-club. The 8 hollow, tapering tentacles are each about twice (115 mm.) as long as
the bell-diameter.

There are 16 subrectangular marginal lappets, with shallow median notches and rounded
angles. The septum between the ultimate branches of the radiating stomach-pouches in the
marginal lappets is twice as wide as the ultimate pouches themselves. The 4-sided throat-
tube is as long as the bell-radius. The 4 lanceolate lips or palps, with their complexly folded
margins are each about 1.33 as long as the bell-diameter. Thus in an adult medusa with a
disk 49 mm. wide the palps were 68 mm. long.

The bell has a rich rose-purple tinge; the gonads, the entodermal cores and the ten-
tacles being especially deep in this color. The warts upon the exumbrella and along the
outer edges of the palps are orange-brownish red.

This medusa is abundant at times in the Mediterranean especially in summer, although
large specimens are rarely seen in winter. It may be locally abundant during several suc-
cessive seasons and then vanish for years. For many years it was all but unknown in the
Bay of Naples but since 1900 it has been one of the commonest Scyphomeduse in this
region. It ranges widely over the warm regions of the Atlantic.

The development has been studied by Kaohot Kowaleysky, Hamann, Goette, Hyde, and
Metschnikoff. Hamann, 1883, has made a detailed study of the development of the gonads,
and their structure has been described by the brothers Hertwig, 1878. ‘They appear as 4 .
interradial, elongate ridges in the entoderm of the subumbrella. anne entoderm forms a series
of follicles in which the sex-cells develop and then migrate into a gelatinous lamella between
the layers of entoderm.

According to Metschnikoff, the egg is violet-brown and is laid between 12 and 2 in the
afternoon, in December, in the Mediterranean. Segmentation is total and nearly equal,
and a very large central segmentation-cavity is formed. The gastrula results from invagination
at the hinder end of the body. The blastopore does not binees but forms the Pot of the
larva. Thus, according to Goette, 1893, the mouth is ectodermal and forms by invagination
at the hinder end of the larva, but the invaginated sac by no means fills the segmentation
cavity. The first pair of stomach-pouches arise from the entoderm and are 180° apart, then
follows an ectodermal pair go° away from the first. The latter then develop 2 lateral pouches
each, and at a later period the entodermal pair each gives rise to 2 lateral pouches, thus giving
a larva with 6 ectodermal and 6 entodermal stomach-pouches; finally the ectodermal pouches
give rise to 4 new adradial pouches and the larva has 16 stomach-pouches—1o ectodermal
and 6 entodermal. There is thus a striking analogy between its development and that of
the scyphostoma of 4urellia, according to Goette.

The external features of the transformation of the free-swimming larva into the medusa
have been studied by Krohn (1855), Kowalevsky (1873), etc. The mouth-end of the larva
becomes expanded and crater-like, with the mouth at summit of central cone of crater. The
depressed region around the cone becomes the subumbrella. The lappets, into which the
gastrovascular cavity is continued, grow out at intervals around the margin. The covering
of cilia is lost from the body of the fee and it begins to swim by means fae rhythmical con-
tractions of its oral disk. Thus the free- -swimming scyphostoma is converted into a medusa
without strobilization (see Goette, 1893.)

Reasoning by analogy from the excellent work of Hyde, 1894 (Zeit. fiir wissen. Zool., Bd.
58, p- 531), upon Aurellia, it is probable that only the subumbrella floor of the second pair
of evaginated gastric pouches is formed from ectoderm, their exumbrella sides being of
entoderm. (See also Hadzi’s work upon Chrysaora.)

574 MEDUS.® OF THE WORLD.

Pelagia noctiluca var. “neglecta.”
Pelagia neglecta, VANHOFFEN, 1888, Bibliotheca Zoologica, Heft. 3, p. 9, taf. 6, fign. 6-12.

This variety is distinguished by the large, elliptical nematocyst-warts upon its exumbrella.
These warts are usually about twice as long as they are wide and display cross-furrows. Speci-
mens in which the bell is 53 to 60 mm. wide have a bell- -height of 23 to 28 mm. Mouth-tube
15 to 25 mm. long and mouth-arms 68 to 85 mm. Color (?) This species is found at Naples
and at the Canary Islands. Were it not for the very large, elliptical nettle-warts of the exum-
brella, it would be identical with the ty pical Pelagia noctiluca Péron and Lesueur. It is so
closely related to P. noctiluca that I believe in view of the ordinary variability of individuals
of the same species in Scyphomedusz, it had best be omitted from further consideration and
merged with P. noctiluca.

Pelagia cyanella Péron and Lesueur.
Plate 61, fig. 1.

Medusa pelagica, Linn#, 1758, Systema Natura, Ed. 10, p. 660.

Medusa pelagica, Linn, 1766, Systema Natura, Ed. 12, p. 1098.—1788 (Gmelin), tomus 1, pars 6, p. 3154.

Pelagia cyanella, PERon et Lesurur, 1809, Ann. Mus. Hist. Nat. Paris, tom. 14, p. 349, No. 66.

Dianaea cyanella, LaMARCK, 1816, Hist. Anim. sans vert., tome 2, p. 507.

Pelagia cyanella, Escuscuoutz, 1829, Syst. der Acalephen, p. 75, taf. 6, fig. 1.—Bosc, 1830, Hist. Nat. des Vers., Ed. 2, tome 2,
p- 140, plate 17, fig. 3.—Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 128, 164; Ibid., 1860, vol. 3, plate 12,
figs. 1-16.—Acassiz, A., 1865, North Amer. Acal., p. 47, fig. 63 —Harcket, 1880, Syst. der Medusen, p. 507.—VAN-
HOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 22.—Bicetow, 1890, Johns Hopkins Univ. Circ., vol. 9, No. 80,
p- 66.—Harcirr, 1904, Bull. U. S. Bureau of Fisheries, vol. 24, p. 70, plate 7, fig. 1.

This American medusa is very closely related to the European P. noctiluca, of which it
is apparently only a local variety.

Bell about 40 mm. high and 50 mm. broad; somewhat fuller than a hemisphere, being a
little less broad at margin than a short distance above. Numerous small wart-like nemato-
cyst capsules are sprinkled thickly over the exumbrella and are especially thick in a zone at
about mid-height of bell; these protuberances are reddish in color and tend to be arranged in
radiating lines. 8 very long, highly contractile, hollow tentacles alternate with 8 marginal
sense-organs. Each sense-club is set within a niche between two adjacent lappets and is
protected on the outer side by a partial web between the lappets. The club is hollow and has
no ocellus, but contains a terminal, entodermal mass of crystalline concretions which are deeply
pigmented. 16 marginal lappets, hemispherical in shape. There is a long, narrow, 4-sided
proboscis, the radial corners of which extend downward as 4 long, flexible mouth-arms, the
free edges of which are complexly crenulated. The proboscis, together with the mouth-arms,
or palps, is about 3 times as long as bell-height. There are 4 complexly folded horse-shoe-
shaped gonads in interradial positions upon the floor of the subumbrella, and immediately
centripetal to them are 4 subgenital pits or cavities extending inward from the outer surface
of the subumbrella. The quadrangular cesophagus leads into a circular, disk-shaped, central
stomach which gives rise to 16 radial pouches extending outward in the radii of the sense-
organs and tentacles. Each of these pouches gives off a pair of unbranched, curved canals
which enter the lappets, but do not form a ring-sinus. There are 16 well-developed strands
of radiating muscle fibers in the wall of the exumbrella adjacent to the gastrovascular cavity.
These extend outward in the radii of the tentacles and sense-organs, and fork as they approach
the bell- -margin.

The color is quite variable, sometimes bluish, sometimes slightly yellowish. Exumbrella
and mouth-arms sprinkled over with brownish-red nettling-warts, tentacles reddish- -purple.

This species is found among the West Indies and Florida Reefs, and in summer it may
drift northward in the Gulf Stream so as to appear off the southern coast of New England
from July to September.

L. Agassiz, 1860 and 1862, found that the planule of this species, as in P. noctiluca,
develop directly into medusz without going through a sessile scyphostoma stage and without
alternations of generations. The planule are set free into the water where each develops into
a single medusa. The minute details of the development have been worked out upon Pelagia
noctiluca by Metschnikoff, 1886 (Emb. Stud. an Medusen, Wien.), and by Goette, 1893
(Zeit. fiir wissen. Zool., Bd. 55, pp. 659-692). The gastrula is formed by invagination. The
first pair of radial stomach-pouches appear, according to Goette, as outpocketings from the

: oe cal mais pai ash S Wive
Bi did oz. A sy ewer. |,

| :
Ane Par i He bhai i mh

tia

> isx
.

Pate 61.

Pelagia cyanella, from the borders of the Gulf Stream off Woods Hole,
Massachusetts.

Drawn from life, by the late Prof. William K. Brooks and kindly pre-
sented for publication in this work.

PLATE 61

BROOKS

Adoent Co,

SEMAEOSTOMEA—PELAGIA, 575
entoderm and these are quickly followed by another pair from the ectoderm of the throat-
tube, the two latter being go° away from the former. ‘The ectodermal pouches then give rise
each to two side branches and soon thereafter the entodermal do the same. Thus the cen-
tral stomach comes to have 12 radial pouches. 4 more radial pouches are soon formed
from the ectodermal pouches, so that the young medusa finally possesses 16 radial pouches.
It follows in adult medusa that the center of the exumbrella side of the central stomach is derived
from entoderm. 2 diametrically opposed, perradial pouches are ectodermal in origin and the
other 2 are entodermal. he 4 interradial pouches are ectodermal, and of the 8 adradial
pouches, 4 are ectodermal and 4 entodermal. The wall of the cesophagus is of ectodermal
origin. The young medusa soon develops 8 lobes which bifurcate, giving 16 marginal lap-
pets. The 8 marginal sense-organs develop before the tentacles. The mouth is at first a sim-
ple, round opening at the center of the crater-like ectodermal depression. It soon acquires
4 lips, but the mouth-arms do not develop until a later stage. It is probable that the ecto-
derm does not take so large a share in the formation of the stomach-pouches as Goette sup-
poses (see Chrysaora and Aurellia).

Pelagia panopyra Péron and Lesueur.

Medusa panopyra, Péron et Lesurur, 1807, Voyage aux Terres Australes, planche 31, fig. 2.

Pelagia panopyra, Péron et Lesurur, 1809, Annal. du Mus. Hist. Nat., tome 14, p. 349.—Escuscuortz, 1892, Syst. der Acal.,
P- 73, taf. 6, fig. 2—Branpr, 1838, Mém. Acad. St. Pétersbourg, tome 4, sér. 6, Sci. Nat., p. 382, taf. 14, fig. 1; taf. 14
A, fign. 1-5.—Harcke, 1880, Syst. der Medusen, p. 509 (literature).—Vanuorren, 1888, Bibliotheca Zoologica, Heft. 3,
p- 14.—FewkeEs, 1889, American Naturalist, vol. 23, p. 592, fig. 1; 1889, Bull. Essex Inst. Salem, vol. 21, No. 7, p. 122,
plate 5, fig. —Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 29.—Mayer, 1906, Bull. U. S. Fish Com-
mission, vol. 23, part 3, p. 1139, plate 2, figs. 3, 4. —Bicetow, H.B., 1909, Mem. Museum Comp. Zool. at Harvard College,
vol. 37, p- 43-—Maas, 1909, Abhandl. Akad. Wissen. Miinchen, Suppl. Bd., 1 Abhandl. 8, p. 43—Kisuinouye, 1910,
Journal College Sci. Tokyo, vol. 27, art. 9, p. 9.

The characters of this widely distributed Pacific form are described in the table under
the genus Pelagia. It is distinguished from the closely allied P. flaveola by its pink colora-
tion, whereas P. flaveola is yellowish. Moreover the nettle-warts of P. panopyra are low
and domelike, while in P. flaveola they are erect and bluntly pointed.

Fic. 364.—Pelagia flaveola, from Tahiti, after Agassiz and Mayer, in Mem.
Mus. Comp. Zool. at Harvard College.

P. panopyra is common off the coast of California and extends across the Pacific to the
Malay Archipelago, and northward to Japan.

Vanhoffen gives the dimensions of a specimen as follows: Bell 27 mm. wide, 6 mm. high;
mouth-tube 15 mm. long; lips 27 mm. long.

When young the medusa has only 4 tentacles. When the bell is 15 mm. wide the gonads
begin to develop.

Pelagia panopyra var. placenta.

Pelagia placenta, Harcxet, 1880, Syst. der Medusen, p. 510.—VaNnuOrreN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 12,

taf. 6, fig. 20.

This appears to be a very close variety of P. panopyra. Both medusx are widely dis-
tributed over the tropical Pacific. See synoptic table of forms of Pelagia. I believe that this

576 MEDUS OF THE WORLD.

form can not be distinguished from P. placenta for I have found specimens which intergrade
in one character or another. Many specimens of this medusa were found by the Albatross in
the Philippine Islands in March and April, 1908. The dimensions in mm. of a mature speci-
men are as follows: Diameter of bell, 35; height of bell, 15; length of cesophagus, 14; length
of lips, 30; length of tentacles, 50.

The largest exumbrella warts are near the center of the bell and they decrease in size
toward the margin, where they are very small. The surface of these warts is nodular and
lacks the cross-foldings seen in the typical P. panopyra, but this character intergrades.

Pelagia flaveola Eschscholtz.

Pelagia flaveola, Escuscuortz, 1829, Syst. der Acal., p. 76, taf. 6, fig. 3.—Gorrre, 1886, Sitzungsber. Akad. Wissen. Berlin,
Jahrg. 1886, p. 883.

(2) Pelagia denticulata, Branot, J. F., 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 383, taf. 14, fig. 2.

Pelagia tahitiana, AGassiz, A.. anv Mayer, 1902, Mem. Museum Comp. Zool. at Harvard College, vol. 26, p. 158, plate 8, figs.

34) 35:
Pelagia papillata, Harcket, 1880, loc. cit., p. 509-

For description see synoptic table of forms of Pelagia, and figure 364.

This form is distinguished by its yellow color and long, pointed nettle-warts which cluster
thickly at the aboral apex of the bell. Tropical Pacific from South America to East Africa.

Pelagia perla Haeckel.

Medusa perla, StanpeEr, 1781, Physikal. Belust., p. 58, taf. 13, fign. 1, 2.

Pelagia cyanella, Fores, 1847, Annals and Mag. Nat. Hist., vol. 19, p. 390, plate 9, fig. 5.

Pelagia perla, Harcke1, 1880, Syst. der Medusen, p. 506.—Maas, 1904, Résult Camp. Sci. Prince de Monaco, fasc. 28, p. §7.—
Detar, M.S., 1906, Fisheries of Ireland, Sci. Invert., 1905, No. 7, p. 22, 1 plate-—VANHOFFEN, 1906, Nordisches Plankton,
Nr. 11, p. 45, fign. 6, 7—Browng, 1908, Trans. Royal Soc. Edinburgh, vol. 46, p. 242.

Pelagia discoidea, Escuscuoxtz, 1829, Syst. der Acal., p. 76, taf. 7, fig. 1.
This appears to be a North Atlantic variety of P. cyanella. The nettle-warts of the

exumbrella are more numerous and the outer margin of the lappets are concave instead of

conyex as in P. cyanella. For description see the synoptic table of the forms of Pelagra.

P. discoidea from the Cape of Good Hope is probably identical with P. perla.

Pelagia phosphora Haeckel.

Pelagia phosphora, HArcket, 1880, Syst. der Medusen, p. 506.—VanuGrreEn, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 11,
22, taf. 6, fign. 18, 19; 1892, Ergeb. der Plankton Expedition, Bd. 2, K. d., p. 19; 1902, Wissen. Ergeb. deutsch. Tiefsee
Expedition, Valdivia, Bd. 3, Lfg. 1, p. 36.

This form is closely related to P. cyanella, but may possibly be distinguished from it by
the larger nettling-warts upon the exumbrella, which are round and 0.5 to 1 mm. in diame-
ter and each gives rise to a longitudinal ae like crest. Marginal lappets wider than long.
(Esophagus about one-fourth as long as bell-diameter, moat about equal to bell-diam-
eter in length. Bell hemispherical, 40 to 50 mm. in diameter. Color quite variable, either
purple, violet, or reddish; gonads, tentacles, and median ribs of the mouth-arms of darker
color than other parts of the animal.

Found in the eastern Atlantic from 58° N. to 42° S. lat.; also in the Indian Ocean,
being especially abundant in the tropics.

* Vanhéffen, 1902, gives the following dimensions for this species: Diameter of bell, 76
mm.; height of bell, 25 mm.; tentacles, 76 mm. long; nettling-warts on exumbrella, 2.5
mm. high.

Pelagia “‘minuta” Vanhoffen, 1888 (Bibliotheca Zoologica, Heft. 3, p. 12, taf. 6, fign.
16, 17), is a variety of, or possibly the young of, P. phosphora. The nettling-warts on the
exumbrella, in specimens preserved in alcohol, are elongate with numerous transverse furrows
(due to contraction?). No mature specimens were described by Vanhoffen. Found at
Pernambuco, Brazil, early in July.

Pelagia crassa Vanhoffen.
Pelagia crassa, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 10, 22, taf. 1, fign. 1, 2; taf. 6, fign. 13, 14.
Pelagia crassa yar. sublaevis, Ibid., p. 11, taf. 6, fig. 15.
This appears to be a small variety of P. noctiluca.
Bell about 13 mm. high and about 35 mm. in diameter. Gelatinous substance of exum-
brella thicker than in any other Pelagia. Marginal lappets about twice as wide as long.

SEMAEOSTOMEA)—PELAGIA, CHRYSAORA. 577

(sophagus shorter than in P. cyanella, only about one-quarter as long as bell-diameter.
Mouth-arms also much shorter, only about as long as bell-diameter. Nettling-warts upon
exumbrella larger and differently arranged than in either P. cyanella or P. phosphora; they
are concentrated near apex of bell and not found near margin, elliptical in shape, sometimes
quite flat, or with a well-developed longitudinal comb-like ridge crossed by corrugations.
The largest warts are found at the apex of the exumbrella and they become smaller toward
the periphery. In the variety sublevis the bell is arched and more hemispherical than in
P. crassa. The gelatinous substance is thicker and the nettling-warts are larger and flatter
than in P. crassa. ‘These forms are found in the middle of the tropical Atlantic between
Africa and South America.

Genus CHRYSAORA Péron and Lesueur, 1809.

Chrysaora, Peron et Lesurur, 1809, Ann. Mus. Hist. Nat. Paris, tome 14, p. 364.—Escuscuoxtz, 1829, Syst. der Acalephen,
p- 78.—Branpt, 1838, Mém. Acad. des Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 384.—Harcket, 1880, Syst. der
Medusen, p. 510.—Craus, 1883, Untersuch. Organisation und Entwick. Medusen, pp. 1-22.—von LENDENFELD, 1884,
Proc. Lin. Soc. New South Wales, vol. 9, p. 268.—Kisuinoure, 1899, Zoolog. Anzeiger, Bd. 22, p. 44 —VANHOFFEN, 1888,
Bibliotheca Zoologica, Heft. 3, pp. 14, 22; 1906, Nordisches Plankton, Nr. 11, p. 47——Hapi1, 1907, Arbeit. Zool. Inst.
Univ. Wien, Bd. 17, p. 17, fign. 1-15, taf. 3, 4.—Heric, Ibid., p. 95, taf. 9. ;

Chrysaora+ Melanaster, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 162.

The type species is C. mediterranea Péron and Lesueur, of the Mediterranean.

GENERIC CHARACTERS,

Pelagide with 8 marginal sense-organs, 24 tentacles (3 in each octant), and with 32
marginal lappets.

The meduse of Dactylometra pass through a “Chrysaora stage” in their development,’
and there can be but little doubt that some of the so-called Chrysaora meduse are only im-
mature Dactylometra. The difficulty in distinguishing Chrysaora is still further complicated
by the very variable, individual coloration of these medusz, and by the fact that some species
of Dactylometra, such as D. quinquecirrha itself, become sexually mature in the Chrysaora
stage when living in brackish water. The Chrysaora and Dactylometra meduse are widely
distributed over the tropical and temperate seas and there are many local races. I believe
that future study will reduce the species to two or three, with numerous local varieties, as in
Pelagia or Cyanea.

Chrysaora hysoscella of the Mediterranean and Atlantic, and C. melanaster and C. helvola
of the North Pacific appear to be distinct species, while the other so-called species may be
varieties of the three above named or merely immature specimens of Dactylometra. C. helvola
and C. melanaster finally develop small, lateral lappets upon the sides of their 16 ocular lappets
and thus approach the Dactylometra condition in having 48 marginal lappets. The tentacles,
however, remain 24 (3 in each octant) as in Chrysaora. L. Agassiz proposes the generic
name Melanaster for Pelagide with 48 lappets and 24 tentacles, but as this appears to be
but a transient stage in the process of growth of Dactylometra, or a late stage in the devel-
opment of Chrysaora, we prefer not to adopt it.

Chrysaora hysoscella, which is probably identical with C. mediterranea, is often hermaph-
roditic; young individuals being male, middle-aged ones hermaphroditic, and old ones
female. In other cases, however, the medusz are throughout life of one sex; while in others
male saccules develop among the old female gonads, and in other parts of the entodermal
layer of the subumbrella.

The development of Chrysaora is through a sessile scyphostoma-stage, and the ephyra
is produced by polydiscus strobilization. The ephyra passes through an 8-tentacled stage
in which it recalls the condition seen in the adult of Pelagia.

The mode of origin of the 4 primary stomach-pouches of the scyphostoma, and their
derivatives, has been the subject of a prolonged discussion between Claus and Goette, and
others. The view of Goette that the cesophagus of the scyphostoma is derived from invaginated
ectoderm received support from Hyde, and was generally accepted until 1907 when Hadzi and
also Heric (Arbeit. Zool. Inst. Wien, Bd. 17, Heft. 1) made further studies of the development
of Chrysaora. It should also be remembered that R. P. Bigelow, 1g00, in his study of the
development of Casstopea is in accord with the views which were later put forth by HadzZi.

578 MEDUS2 OF THE WORLD.

Hadii finds that some of the free-swimming planulz of Chrysaora are 4 or 5 times as large
as others. They swim with the broad end forward and soon settle upon ulva, etc., attaching
by means of their forward ends. The entoderm, which was previously a solid mass, then
hollows out and the larva becomes two-layered, and the uppermost (the former posterior) end
becomes the widest. The mouth then breaks through, the oral pole flattens laterally, and 4
tentacles develop, 2 in the short and 2 at the ends of the long diameter. The stomach-pouches
do not begin to form until after the mouth and 4 tentacles have developed.

The view of Claus has received strong support from Hadzi who casts serious doubt upon
Geotte’s interpretation that the cesophagus of the scyphostoma is always composed of invag-
inated ectoderm.

Hadzi, whose research upon Chrysaora appears to have been carefully studied, finds that
the cesophagus of the scyphostoma is entodermal and that the mouth breaks through from the
inside, the entoderm thus taking the active share in its formation, and no invagination of ecto-
derm occurring. Indeed Hadii finds that the cells lining the throat of the scyphostoma resem-
ble ectodermal cells in having nematocyst capsules and glands, but they are nevertheless solely
of entodermal origin. From this it follows that the 4 primary stomach-pouches are also ento-
dermal, not 2 ectodermal and 2 entodermal as claimed by Goette. Hadzi finds also that the 4
intertentacular taniola are formed from 4 simple, longitudinal infoldings of the entoderm of
the stomach wall, the ectoderm taking no part in their formation. The primary stomach-
pouches are thus the passive result of the infoldings which form the tzniola, not of an active
outgrowth of pouches as Goette believes.

Hadzi’s view appears to be the more reasonable, for if Goette were correct one half of the
gonads would be ectodermal and one half entodermal, whereas according to Hadzi they are all
entodermal; moreover, according to Goette, the mouth of the first ephyra set free in strobili-
zation has its cesophagus lined with ectoderm, while those ephyra which follow it have their
throats lined with entoderm, an anomalous condition. According to Hadzi and Heric, how-
ever, all of the ephyra have their throats lined with entoderm.

Heric finds in the strobilization of the scyphostoma of
Chrysaora that with the exception of the terminal ephyra all of
the mouth-tubes of the chain of ephyre are formed from the
connecting tube which joins all of the ephyra together. The
external wall of this connecting tube is ectodermal and its inner
wall entodermal. 4 perradial clefts develop in the side wall of
each tube near the upper end where it joins with the exumbrella of
the overlying ephyra. The lower edges of these clefts grow out-
ward and form the 4 lips of the ephyra, while the 4 connections
are interradial and are in the radii of the taniola which consti-
tute their inner sides.

The 4 subgenital cavities of the ephyra are new formations
and not derived from the 4 funnel-cavities of the scyphostoma.
The 4 interradial septa of the stomach-cavity of the ephyra are,
however, derived from the taniolz of the scyphostoma. These
soon disappear, and the central stomach of the medusa is a
simple lenticular space.

The forms of Chrysaora are so imperfectly separated one from
another that were it not for the fact that many minute distinc-
tions have been pointed out between them, I would greatly
prefer to consider them all to be one variable species, C. hysoscella.

Fic. 365.—Diagrammatic section of wever. we may. possibly distinpuish more less sabaeine
a strobilla of Chrysaora after Howe T, we y P v2 suing or f) y:

Heric, in Arbeit. Zool. Inst. C. hysoscella=C_ mediterranea with its varieties blossevillei and plocamia (?)
Wien. bigs of the Atlantic, Mediterranean, and South Pacific.

als, perradial cleft. am, beginning of C. helvola, with its varieties calliparea, and chinensis of the Pacific and
the lip. p, /, 1, stages in the Indian Oceans.
growth of the throat-tube. C. melanaster with its variety gilberti of the North Pacific.
sm, septal muscle. Ectoderm
cross-hatched, entoderm plain, I believe that a study of the following synoptic table will

intermediate lamella dotted.
NLCAMEEBENMEL SSS convince one that we have here only one species, the varieties of

which defy classification in terms of the Linnean system.

SEMAEOSTOMEA:—CHRYSAORA,

Synopsis of the “Species” of Chrysaora.

579

C. mediterranea Péron
et Lesueur=C. hyso-
scella.*

C. hysoscella Esch-
scholtz.f Identical
with C. mediterranea.

C. blossevillei Lesson, a
variety of C. hyso-
scella.t

C. fulgida Haeckel=

| Rhizostoma fulgidum
Reynaud; a variety of
C. hysoscella.

Shape of disk.
Width of disk in mm.
Height of disk in mm.

Shape of marginal lap-
pets.

Shape of the 8 ocular
stomach-pouches.

Shape of curtain-like lips.

Length of mouth cur-
tains (lips) in terms of
disk-radius r.

Length of longest ten-
tacles in terms of disk-
radius r.

Color.

Where found.

Flatly rounded.
100 to 300
40 to 80

Flatly rounded. The
16 ocular lappets only
half as wide as the 16
tentacular lappets.

At periphery of central
stomach as wide, in the
middle three-fourths, at!
margin half as wide as
tentacular pouches.

Tapering from base to
pointed ends. Edges
are curtain-like and

very complexly folded.
3 to 4r
2r
Variable. Disk is

whitish to yellow. Us-
ually there is a reddish
ring around apex, and
radiating from this are
16 radial streaks.

Mediterranean.

Flatly rounded.
100 to 200
40 to 60

All semicircular and of
equal width. The 16
ocular lappets project
more than the 16 ten-
tacular lappets.

In middle, the 8 ocular
pouches are same
width as 8 interocular.
At bell-margin, ocular
pouches are only one-
third as wide as ten-
tacular pouches.

Tapering from base to
pointed ends. Edges
very much folded.

2r

2r

Variable, and similar
to C. mediterranea, but
usually more intense.

Atlantic coasts of
Europe.

Nearly hemispherical.

100

40

All half-egg-shaped
(oval). Ocular and
interocular clefts about
twice as deep as clefts

which alternate with
them.

In middle, ocular
pouches are equal to
tentacular pouches in
width. At margin,

ocular pouches are only

half as wide as the ten-
tacular.

Lancet-shaped. Taper-

ing with folded margins,

2r

Bell amber to rusty in
color. Mouth-arms
rusty-yellow.

Coast of Brazil, Island
of Santa Catharina to
Pernambuco.

Hemispherical.
300 to 400
T00 to 200

All alike. Semicircular.

Lancet-shaped, widest
in middle where they
are as wide as 0.5r.

4 to 6r

r. Reynaud figures only
16 tentacles.

Bell yellowish-brown.
Radial streaks and
marginal lappets red-
dish-brown. Mouth-
arms reddish; gonads
carmine.

Cape of Good Hope,
False Bay, Algoa Bay,

Africa.

*Development through alternation of generations with strobilization (Claus).

{Development as in C. mediterranea.

as in Dactylometra.

{One of Vanhéffen’s specimens had 4 tentacles in one octant.

127, 166.

Urtica marina, etc., Borrase, 1758, Nat. Hist. Cornwall, p. 256, plate 25,

Chrysaora hysoscella Eschscholtz.

Medusa hysoscella, Linné, 1766, Systema Nature, Ed. 12, p. 1097.
Chrysaora cyclonata, asptlonota, spilogona, spilhemigona, pleurophora, lesueurii, macrogona, heptanema, mediterranea, PERON ET
Lesueur, 1809, Annal. du Mus. Hist. Nat., tome 14, pp. 365, 366.
Chrysaora hysoscella, Escuscuoxtz, 1829, Syst. der Acal., p. 79, taf. 7, fig. 2—Craus, 1877, Denksch. Wien. Acad., p. 33, taf.
6, 7-—KRruKENBERG, 1880, Zool. Anzeiger, Jahrg. 3, p. 306 (the medusa contains 95.75 to 96.3 per cent of water).—GRaEFFE,
1884, Arbeit. Zool. Inst. Wien., Bd. 5, p. 342.—VANHOFFEN, 1908, Deutsche Siidpolar Expedition, Bd. 10, Zool. 2, p. 39.
Chrysaora mediterranea+ C. isosceles, Tee 1880, Syst. der Medusen, pp. 511, 513, taf. 31, fign. 1-3 (list of literature).
Chrysaora mediterranea, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft 3, p- 14.

Chrysaora hysoscella=

figs. vii and vir.

Haeckel found a specimen 160 mm. wide, with 40 tentacles, and 48 marginal lappets

See also L. Agassiz, 1862, Cont. Nat. Hist. U.S., vol. 4, pp.

C. mediterranea, VANHOFFEN, 1906, Nordisches Plankton, Nr. 11, p- 47, fign. 9-10 a, b.

Chrysaora isosceles, Detar, M. J., 1901, Irish Naturalist, vol. 10, p. 27 (rearing the larva in an aquarium).

For a synopsis of the characters of C. hysoscella, see table of characters of Chrysaora.

There appear to be no definite distinctions between Chrysaora “

mediterranea”

of the

Mediterranean and C. hysoscella of the Atlantic coasts of France, England, Germany, and
Holland, and in the entrance to the Baltic Sea. It is probable that the Chrysaora of the
Chesapeake Bay, in America, is identical with that of Europe, but I believe the Chesapeake
medusa to be only a brackish-water variety of Dactylometra quinquecirrha, which becomes
mature in the 24 tentacle stage. Judging from Haeckel’s figures the Mediterranean medusa
is more highly colored than that of the northern shores of Europe. In the Chesapeake the

580

MEDUS OF

THE WORLD.

Synopsis of the “Spectes” of Chrysaora—Continued.

C. plocamia
Haeckel, a variety
of C. blossevillei.

C. helvola Brandt.

C. calliparea
Haeckel, a variety
of C. helvola.

C. chinensis Van-
hoffen, a variety of
C. helvola.

C. melanaster.*
See C. gilberti
Kishinouye.

Shape of disk.

i
Width of disk in
| mm.
Height of disk in
mm.

Shape of marginal
lappets.

| Shape of the 8
ocular stomach-

pouches.
|

Shape of curtain-
like lips.

Length of mouth-
curtains (lips) in
terms of disk-
radius r.

Length of longest
tentacles in terms
of disk-radius r.

Color.

Where found.

Hemispherical.

85 to 100
4o to So

All alike. Semicir-
cular.

All 16 radial-
pouches of equal
width in middle.
At margin ocular
are only half as
wide as tentacular
pouches.

Curtain-like, folded.
In middle as wide
as radius of um-
brella.

Hemispherical, or
flatter. Exum-
brella smooth.

100 to 300

5°

Oval. The adradial)
clefts between
tentacular lobes are}
deeper than others,
causing general
outline of each
octant to be convex.

Ocular pouches are
in middle twice as
wide, at the mar-
gin half as wide,
as tentacular
pouches.

Lancet-shaped, in
middle two-thirds
as wide as disk- |

3r

2r

| Bell rusty-yellow.
Mouth-arms color-
less, with yellow

red.

West Coast of South
America. Cape
Horn to Peru.

margins. Tentacles)

radius. Margin |
complexly folded.
4r

Flat, ribbon-like. 4r
long.

Bell and lips light
yellowish-brown,
with marginal lap- |
pets and mouth-
arms speckled with
rusty-red. Tenta-
cles dark rusty
color.

North Pacific Ocean,
Asia to California.

Flatly rounded to
hemispherical.

160 to 200
50 to 100

Kidney-shaped,
wider outward than
at bases. The 16
ocular lappets
wider and longer
than the 16 tentac-
ular lappets.

Ocular stomach-
pouches spindle-
shaped. In middle
as wide as tentacu-
lar pouches.

Curtain-like, very
wide and long.

6 to 8r

Ribbon-like at their
bases 2r—-

Ground color of um-
brella bright red-
dish-yellow, with a
32-rayed chestnut-
brown star on ex-
umbrella. Lappets
brown. Mouth-
arms yellow,
spotted with brown.|
Gonads yellow.

Indian Ocean to
east coast of Africa.

Flatly rounded to
hemispherical.

7o
3°

Lappets longer than
wide, triangular
with blunt points.

Twice as wide in
middle; at margin,
half as wide as

tentacular pouches.

Curtain-like. In
middle as wide as
radius of umbrella.

7

Southern parts of
the China Sea.

Hemispherical or
flatter.

200 to 300
100 to 150

Tongue-shaped,
narrower at base
than beyond this
point. All of equa’
size. When old,
the 16 ocular lap-
pets develop side
lappets as in
Dactylometra.

Ocular and tentacu-
lar pouches of ae
lar form and size.

Tapering from a
wide base to
pointed ends. In
middle one-third r
wide. Margins
much folded.

2r

Ribbon-like r-

Bell, mouth-arms,
and tentacles light-
bluish. 32 star-
like brown rays on
exumbrella. 16
dark-brownish to
black radial streak:
on subumbrella in
radii of 16 lateral
tentacles. Gonads
reddish-brown.

North Pacific. Asia
to California.

*This species shows a tendency to approach the Dactylometra stage in having 48 marginal lappets, but the tentacles remain
24 in number as in other species of Chrysaora. L. Agassiz, 1862, calls this Melanaster mertensit.

medusa is highly variable in color, sometimes pale, nearly uniform, milky ocher-yellow, and
at other times with radiating streaks of rich red-brown.

The medusa is locally common but rare in many other places, such as Naples, where
it is hardly ever seen. ‘

According to Graeffe, C. hysoscella is found at Trieste, Adriatic Sea, from November to
April, and is mature in winter. It is hermaphroditic, and the eggs segment in the ovaries and
are set free as pear-shaped planula in March and April. At the end of 8 to 14 days they become
attached by the broad anterior end and finally acquire 16 tentacles in the scyphostoma-stage.
These strobilate in September and October and each gives rise to 8 to 12 ephyre.

SEMAEOSTOMEA—CHRYSAORA, 581

Miss M. J. Delap finds that the scyphostoma of Chrysaora does not eat copepods or small
fish, but feeds voraciously upon hydromeduse, siphonophore, ctenophora, and pelagic worms.

Claus, 1877, has studied the growth of the ephyra. At first the 8 ephyra lobes are very
long and slender with deep, aaa clefts between them and with sharp-pointed, marginal
lappets. There are 8 sense-organs but no tentacles. Later 4 and then 8 tentacles develop
and the medusa is in the Pelagia-stage. The first 4 tentacles are in the 4 adradial clefts which
precede the 4 perradii in the direction of the hands of the watch (see text-fAgure 372). The
adradial stomach-pouches develop later than the perradial and interradial.

The hermaphroditism of Chrysaora was first observed by Derbés (1850, Annal. des Sci.
Nat., tome 13, p. 377) and later by Wright (1861), Claus (1877), and Haeckel (1880). Young
sadivdluals are apt to be male, middle- “aged ones Reacepiearas older ones female. Some
are, however, of one sex only throughout life. In other cases when the production of eggs
in the entoderm of the gonads begins to decline, a number of sac-like cavities, of irregular size,
develop not only in the gonads but in other parts of the subumbrella wall, and these give rise
to spermatozoa.

Chrysaora hysoscella var. blossevillei.
Chrysaora blossevillei, Lesson, 1829, Voyage de la Coquille Zool., p. 115, planche 13, fig.2; 1843, Hist. Zooph. Acaléphes, p. 401.—

HaceckEL, 1880, Syst. der Medusen, p. 514.—LENDENFELD, R. von, 1884, Proc. Linnean Soc. New South Wales, vol. 9,

p- 268 —Vanuorren, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 15, taf. 1, fig. 3.

Lobocrocis blossevillei, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. es p. 166.

This is closely allied to C. hysoscella, of which it is probably a local variety.

Disk 28 to 37 mm. wide, or larger, and about 7 to g mm. thick. Exumbrella covered
with small, round, nettling warts. According to Vanhoffen there are 4 marginal lappets in
each octant between successive sense-organs. The § interocular clefts are twice as deep as are
the 16 clefts adjacent to them. The longest tentacles arise from the interocular clefts. These
are about as long as bell-radius. The 2 ote tentacles in each octant are only half as long as
the interocular iors There are thus 24 tentacles, 32 lappets, and 8 ‘marginal sense-
organs. 16 radial pouches extend out from the central stomach and these are twice as wide
at margin in the interocular as in the ocular radii. The palps are shorter than bell-diameter.
Bell and palps yellowish, speckled with rusty-red. Coast of Brazil. Best description by
Vanhoffen.

Chrysaora hysoscella var. fulgida.

Rhizostoma fulgidum, ReyNaup, 1830, Lesson’s Centurie Zoologique, p. 79, plate 25.

Chrysaora fulgida, Harcxet, 1880, Syst. der Medusen, p. 514.—VANHGFFEN, E., 1902, Wissen. Ergeb. deutsch. Tiefsee Expedi-
tion, Valdivia, Bd. 3, Lfg. 1, p. 38.
See synoptic table of forms of Chrysaora.
This form is very imperfectly known and may be identical with C. hysoscella. It is found

off the Cape of Good Hope, Africa.

Chrysaora blossevillei var. plocamia.

Cyanea plocamia, Lesson, 1829, Voyage de la Coquille Zool., p. 116, plate 12, figs. 1, 2.
Chysaora plocamia, Harcxet, E., 1880, Syst. der Medusen, p. 516. Sy core 1888, Bibliotheca Zool. Bd. 1, Heft. 3, p. 16.

See synoptic table of forms of Chrysaora.

This form is found on the South Pacific coast of South America from Peru southward,
and is allied to C. blosseville: of the Atlantic side. Its marginal lappets are said to be semi-
circular instead of oval as in C. blosseville’, and the contour of the bell-margin is circular, not
octangular with concave inter-rhopalar sides. It is only misleading to attempt to separate
species upon distinctions such as these.

Chrysaora helyola Brandt.

Chrysaora (Polybostrycha) helvola, Branvt, 1838, Mém. Acad. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p.384, taf. 15, fign. 1-4.
Chrysaora helvola, HarcKet, 1880, Syst. der Medusen, Pp. 515.—VANHOFFEN, 1906, Nerdieclice Plankton, Nr. 11, p. 48, fig. 11.
Chrysaora convoluta, Kisttnoure, 1910, Journal College of Sci. University Tokyo, vol. 27, art. 9, p- 11, plate 2, figs. 11-14.

See synoptic table of the species of Chrysaora.

Distinguished by the considerable length of the 16 ocular lappets, which project beyond
the zone of the velar ones, thus giving an octangular appearance to marginal outline of bell.
The medusa is found in the North Pacific from Asia northward to Alaska and eastward to
the coast of California. Kishinouye finds that young cod are often found under the bell of

582 MEDUS OF THE WORLD.
this medusa. He gives good figures of it showing the considerable depth of the adradial clefts
between the lappets.
Chrysaora helyola var. calliparea.
(?) Cyanea calliparea, ReyNauD, 1830, Lesson’s Centurie Zoologique, p. 67, planche 20.
Chrysaora calliparea, HarckeL, 1880, Syst. der Medusen, p. 516.
See synoptic table of forms of Chrysaora.
This medusa is reported from Pondichery and Zanzibar, and our knowledge of it is still
imperfect.
Chrysaora helvola var. chinensis.

Chrysaora chinensis, VANHOFFEN, 1888, Bibliotheca Zoologica, Heft. 3, p. 16.

In the single specimen described by Vanhéffen the disk is 70 mm. wide and 30 mm. high.
Exumbrella thickly covered with numerous, very small warts arranged in a scale-like manner
from summit to margin. Ocular and tentacular marginal lappets of equal width and length,
longer than wide, and each one ends in a blunt point; lappets overlap each other on their
sides and this causes the ocular lappets to appear smaller and sharper than the tentacular
lappets. The 16 radiating stomach-pouches are all of equal width at periphery of central
stomach. The septa bordering the 8 ocular pouches trend so as to become wider apart for
the proximal three-fourths of their lengths, so that the ocular pouches are about twice as wide
as the tentacular ones at this zone. The septa then trend toward the ocular radii so that at
the bell-margin the 8 ocular stomach-
pouches are only half as wide as the 8
tentacular ones. The 4 lips, or palps,
are each 7 times as long as the radius
of the disk. Near the mouth-opening
they are almost as wide as bell-radius
and are curtain-like. Midrib of each
mouth-arm well developed, the folded
margins thin. Outer parts of palps
about one-third as wide as bell-radius,
not tapering outwards, but ending
bluntly. Color (?) Gonads (?) Tenta-
cles (7) Found near Hongkong, China,
in October.

: This form is distinguished by its
Fic. 366.—Chrysaora helvola, according to Brandt, after Vanhoffen, peculiar exumbrella warts and its very
ene long mouth-arms. It is separated from

Chrysaora calliparea by its long, pointed, marginal lappets.

Chrysaora melanaster Brandt.

Chrysaora melanaster, Branvt, 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 385, taf. 16, 17.—HarckeL,
1880, Syst. der Medusen, p. 515.—Ferwxes, 1889, Bull. Essex. Inst. Salem, vol. 21, No. 7, p. 121.—VANHOFFEN, E., 1906,
Nordisches Plankton, Nr. 11, p. 49, fig. 12.

Melanaster mertensiiy AGAssiz, L., 1862, Cont. Nat.' Hist. U. S., vol. 4, pp. 126, 166.

Melanaster, sp., Acassiz, A., 1865, North Amer. Acal., p. 50.

See synoptic table of species of Chrysaora.
This abundant North Pacific form is found from Kamtschatka to California. Bell

with reddish-brown radial lines extending from center to margin. Marginal sense-organs
bright yellow and brown. ‘Tips of tentacles are red. When old it develops 48 lappets, but

the tentacles remain 32.
It is closely allied to Dactylometra quinquecirrha of the North Atlantic, and C. gilbertt

is one of its varieties.
Chrysaora melanaster var. “‘gilberti’’? Kishinouye.
Chrysaora gilberti, Kisuinovye, 1899, Zool. Anzeiger, Bd. 22, p- 44, 1 fig.

Distinguished by its semicircular lappets all similar each to each and its relatively long
tentacles. It is probably only a variety of C. melanaster.

SEMAEOSTOMEA—CHRYSAORA, DACTYLOMETRA. 583

Umbrella 70 to 100 mm. wide, 25 to 30 mm. high. Exumbrella thickly besprinkled with
nematocyst-warts. 32 marginal lappets, all similar each to each and all nearly semicircular.
24 laterally compressed tentacles, all somewhat longer than bell-diameter. The 4 subgenital
ostia are round to oval. The septa between the 16 radial stomach-pouches are nearly straight,
but the ocular pouches are a little broader than the tentacular pouches at the annulus of the
marginal sense-organs. The 4 oral curtains are as broad as the bell-radius near the mouth
and are very long with finely frilled margins. In contraction they are coiled, screw-like;
and in this condition are about as long as bell-radius. They taper to blunt points and their
thick proximal parts are covered with nettle-warts.

Umbrella light brown, tentacles and midrib of mouth-arms darker brown. Common
in Monterey Bay, California, in summer.

Genus DACTYLOMETRA L. Agassiz, 1862.

Dactylometra, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 166.—Acassiz, A., 1865, North Amer. Acal., p. 48.—HarckeEL,
1880, Syst. der Medusen, p. 517.—Kisuinovuye, 1892, Zoological Mag., Tokyo, vol. 4, p. 261.—Acassiz anp Mayer,
1898, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 1—VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee
Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 39; 1906, Nordisches Plankton, Nr. 11, p. 50.

The type species is D. lactea L. Agassiz, of the West Indies and South American coast.
GENERIC CHARACTERS.

Pelagide with 40 tentacles, 5 be-
tween each successive pair of sense-
organs, and with 48 marginal lappets.

Fic. 367.—Chrysaora melanaster, according to Brandt, after Vanhoffen, in Nordisches Plankton.
Fic. 368.—Chrysaora gilberti, after Kishinouye, in Zool. Anzeiger.

There is reason to believe that certain meduse may become mature with 24 tentacles,
while others, possibly better fed individuals of the same species, may develop 40 tentacles
before becoming mature. Similarly Dactylometra longicirra of Japan may develop 40 or
even 56 tentacles before becoming mature. The conditions characteristic of the genera
Chrysaora, Dactylometra, and Kuragea may therefore be transitional stages in the growth
of one and the same medusa.

Dactylometra lactea L. Agassiz.

Chrysaora lactea, Escuscuoutz, 1829, Syst. der Acal., p. 81, taf. 7, fig. 3- ;

Dactylometra lactea, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 125, 126, 166.—HarckeL, 1880, Syst. der Medusen,
p- 517.—Lenpenrexp, R,, von, 1884, Proc. Linn. Soc. New South Wales, vol. 9, p. 271.—Acassiz, A., AnD Mayer, 1898,
Bull. Mus, Comp. Zool. at Harvard College, vol. 32, p. 7, plates 12, 13 and fig. 10, plate 7.

584 MEDUS4 OF THE WORLD.

Mature medusa.—Bell hemispherical, smaller than in D. quinquecirrha being about
7o mm. in diameter. Clefts in lappets adjacent to sense-organs fully as deep as those between
the remaining lappets instead of being mere shallow notches, as in D. quinquecirrha. The
tertiary tentacles arise from clefts between the lappets, not from the floor of the subumbrella,
asin D. quinquecirrha. The 8 primary tentacles are about 3 times as long as the bell-diameter.
The 16 secondary tentacles, however, are only about half, and the 16 tertiary, one-quarter
as long as the primary ones.

General color dull milky-white, exumbrella sprinkled over with ocher-yellow-colored
spots, thickly clustered at aboral pole. Genital organs slightly yellowish, a delicate irides-
cence oyer the long, flexible mouth-arms.

Fic 369.—Dactylometra lactea, aboral view, three-fourths natural size. From nature,
by the author. Showing variable character of the lobes and tentacles.

This species is found at Rio Janeiro, Brazil, and at Jamaica and Cuba. In Havana
Harbor it is abundant and mature in February.
A mature specimen found by me off Port Royal, Kingston

SO ale Harbor, Jamaica, on March 20, 1909, was of the following
Nowun|itectaclert | elobes dimensions in millimeters: Bell 66 wide, somewhat flatter
— == — than a hemisphere, palps 50, longest tentacles 60 long. Exum-
1 | 3 4 brella regularly and thickly besprinkled with very small, low-

| 2 4 4 lying, milky-yellow colored nettling-warts. 16 spoke-like stripes
: : : of dull ocher color and numerous russet-brown nematocyst-warts
5 4 5 at the margin of the exumbrella. Gelatinous substance and ten-
|e 3 tacles milky. Gonads dull milky-pink. This specimen was very

; | 3 : irregular in the development of its tentacles and lobes, the 8

octants being as shown in the table (fig. 369).

PLATE 62.

Fig. 1. Dactylometra quinquecirrha, natural size. In the “‘Chrysaora” stage.
From the brackish water of the St. Mary’s River, Chesapeake
Bay, Maryland, November 13, 1905.

Fig. 2. Dactylometra quinquecirrha, natural size. In the “Chrysaora” stage.

Agassiz Laboratory, Newport, Rhode Island, August 2, 1892.

Drawn from life, by the author.

MAYER PLATE 62

Lc ee

Peiniees ac irvine
el of &

—

rin
re ee

al
vehi

PLATE 63.

Dactylometra quinquecirrha, natural size. Mature in the “Chrysaora” stage,
in the brackish water of Norfolk Harbor, Virginia, November 4, 1904.

Drawn from life, by the author.

PLATE 63
_ MAYER

PLaTE 64.

Mature Dactylometra quinquecirrha, half natural size. Highly-colored, mature
specimen from Tiverton, Narragansett Bay, Rhode Island.
September, 1896.

Drawn from life, by the author.

MAYER

PLATE 64

ie
r
e
a
ell ee He! og eo
: TTS) ; Aes
iL m
7
f rs
1g .
:
“=

Sor

PLATE 64A.

Dactylometra quinquecirrha, drawn by the late Prof. William K. Brooks from
a specimen obtained at Beaufort, North Carolina.

PLATE 644

BROOKS

AMeon tle.

SEMAEOSTOMEA—DACTYLOMETRA. 585

Dactylometra quinquecirrha L. Agassiz.
Plates 62 to 64A.

Pelagia quinquecirrha, Desor, E., 1848, Proc. Boston Soc. Nat. Hist., vol. 3, p. 76.

Dactylometra quinquecirrha, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 125, 166.—AGassiz, A., 1865, North Amer.
Acal., p. 48, fig. 69.—Harcket, 1880, Syst. der Medusen, p. 518.—Fewxes, 1882, Bull. Mus. Comp. Zool. at Harvard
College, vol. 9, No. 8, p. 293, plate 1, figs. 25-28, 38, 39.—BicrLow, 1890, Johns Hopkins Univ. Circulars, vol. 9, No. 80,
p- 65.—Acassiz, A., anp Mayer, 1898, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 1, plates 1-11, 33 figs.—
Harcirt, 1904, Bull. U.S. Bureau of Fisheries, vol. 24, p. 69, plate 7, fig. 2—Haraitr, 1905, Journal Exper. Zool., vol. 2,
P- 575 (variations)—VANHOFFEN, 1906, Nordisches Plankton, Nr. 11, p. 50, fign. 13-14.

Bathyluca solaris (damaged and regenerating specimen?), Mayer, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37,
p- 2, plate 1.

Chrysaora, BicEtow, R. P., 1880, Johns Hopkins Univ. Circulars, vol. 9, No. 8, p. 66 (brackish-water variety from Chesapeake
Bay).

Adult medusa.—Bell nearly hemispherical, 170 to 190 mm. in diameter. Numerous
small, wart-like clusters of nematocysts thickly scattered over the exumbrella, especially

abundant at aboral apex where they appear as little hemispherical projections above the
general surface; near the margin they are elongate in shape, while at the margin itself they

S
ISS

Cs ¥

Fics. 370.—Dactylometra lactea from Havana, Cuba; after Agassiz and Mayer in Bull. Mus. of Comp. Zool.
A, side view; B, oral view.

are again hemispherical as at the apex. 8 marginal sense-organs, 40 tentacles, and 48 marginal
lappets. The marginal sense-organs are set within niches between the lappets, 4 being per-
radial in position and 4 interradial; these niches are protected above by a small web between
the lappets. A ciliated, pit-like depression extends downward from the surface of the ex-
umbrella immediately above each sense-organ. The sensory-club projects slightly down-
ward and contains a distal, entodermal mass of crystalline concretions but no ocellus. The
entodermal core of the sense-club is hollow and its lumen is connected with the general
gastrovascular space of the medusa.

There are 5 tentacles between each successive pair of sense-organs. 3 of these tentacles,
the primary and secondary, arise from the clefts between the lappets, but the other 2 (tertiary)
are generally found to spring from the under or subumbrella side of the ocular lappets; for
even in very large medusz the ocular lappets exhibit but a slight notch adjacent to the ter-

586 MEDUSZ OF THE WORLD.

tiary tentacles; in fact, the tertiary tentacles do not usually make their appearance until
the medusa is about 130 mm. in diameter and the lappets remain undivided until the medusa
is mature, although Hargitt shows that this is subject to great individual variability. Thus
in immature medusz of large size there are usually but 24 tentacles and 32 marginal lappets,
and the animal is in the “‘Chrysaora stage.” I believe, also, that they often mature in this
stage and never reach the Dactylometra condition.

The primary and secondary tentacles are very long and flexible while the tertiary ten-
tacles are only a few millimeters in length. In like manner the lappet-clefts of the primary
and secondary tentacles are deep and the lappets almost as long as they are broad; while
the lappet clefts of the tertiary tentacles are mere shallow notches in the contour of the lappets
adjacent to the sense-organs. Mouth-opening cruciform, in center of subumbrella, at extremity
of a 4-cornered cesophagus and surrounded by 4 mouth-arms or palps, which when fully
extended are about 3 or 4 times as long as the bell-diameter. The 8 free edges of the mouth-
arms are complexly crinkled and highly flexible. The central stomach occupies a wide lentic-
ular space in the midst of the bell and gives rise to 16 simple, radiating pockets, 8 in the
tentacular and 8 in the rhopalar radii. These pockets are completely separated one from
another by 16 radiating septa which join the upper and lower walls of the umbrella cavity
together. The tentacles are hollow
throughout the greater part of their
length and their entoderm is ciliated
as is that of the stomach itself.

The gonads are contained in 4
interradially situated, entodermal in-
foldings of the wall of the subum-
brella, and their position is marked
by 4 deeply sunken, subgenital pits.
The genital organs are provided
with numerous, simple, unbranched
gastric cirri which project inward
into the stomach-cavity. There are
two sets of radial muscle-fibers; the
principal set is found in the 16 septa
between the gastric pouches, and
alternating with these in position are
16 strands in the exumbrella, 8 of
which lead outward to the sense-

; organs and 8 to the primary tentacles.

Fic. 371.—Dactylometra quinquecsrrha, after Agassiz and Mayer, in Color quite variable. In some

Bull. Mus. of Comparative Zoology at Harvard College. ogee eaesn Fhs 5

individuals the disk is pink, in others

yellow with a bluish opalescence. The exumbrella is thickly sprinkled with yellow-ocher
colored nettling-warts and there are 16 radiating stripes of reddish color upon the exumbrella in
the radii of the septa of the peripheral stomach. These reddish stripes extend about half-way
from the bell-margin toward the center of the exumbrella and are due to highly refractive,
rosin-colored pigment granules within the epithelial cells of the disk. The male gonads are
generally pink, while the ovaries are yellowish or ashy-gray. The radial muscle-strands of
the subumbrella are of a glistening white and the entodermal cores of the tentacles are pink.
The mouth-arms are pink or yellow and always sprinkled over with red-colored pigment spots.
The marginal sense-organs contain each a mass of glistening white concretions, but no ocelli.

This species extends from the southern coast of New England to the tropics. In August
it is abundant in Tampa Bay, Florida. It has been taken by Bickmore at the Bermudas,
and by Drayton between the Bermudas and the Azores. “A well-marked southern variety”
was found by Brooks at Beaufort, North Carolina, and is figured in plate 64A. It makes its
appearance upon the surface along the coast of New England in August when large meduse
are found. The young rarely come to view, but remain in deep water.

Varieties and development—The egg develops into a free-swimming planula which
soon attaches itself to the bottom and develops into a scyphostoma having normally 4 ten-

SEMAEOSTOMEAS—DACTYLOMETRA. 587

tacles. R. P. Bigelow, 1880, states that the so-called “‘Chrysaora” of the Chesapeake, which
is only a brackish-water, abortive variety of Dactylometra, develops from an ephyra through a
Pelagia stage, wherein it has only 8 tentacles and 16 lappets, and Brooks has figured the ephyra
in the text figures here shown. ;

The present writer found considerable numbers of Chrysaora-like meduse in Hampton

Roads and Norfolk Harbor, Virginia, and in St. Mary’s River, Maryland, early in Novem-

i

S.

es

—s

.

\\

Zi

Fic. 372.—Young ephyre of Dactylometra quinquecirrha. Figures drawn by the late Prof. William K. Brooks at the
Chesapeake Bay Laboratory of the Johns Hopkins University. Presented by the Department of Biol-
ogy of the Johns Hopkins University for publication in this work.
ber, 1904 and 1905. These were generally pale milky-yellow in color and lacked the rich
brown pigment and the 16 pigmented, radial areas seen upon the exumbrella of Dactylometra
quinquecirrha. Others had a red-brown spot at the apex of the exumbrella, and surrounding
this was a star-like zone of red-brown streaks with pointed ends directed outward. The

588 MEDUS OF THE WORLD.

axial ribs of the mouth-arms (palps) were red-brown. Although all were in the Chrysaora
condition and had only 3 tentacles and 4 lappets in each octant, some appeared to be fully
mature or with gonads nearly ripe. The exumbrella surface and the palps were covered
with dull milky-yellow clusters of nematocysts. There were 8 marginal sense-organs as in
Dactylometra, but only 24 tentacles and 32 marginal lappets. None of the medusz were,
however, as large as is commonly seen in full-grown Dactylometra quinquecirrha, the largest
Chrysaora-like medusa seen in Norfolk harbor being only 105 mm. in diameter. It should
be borne in mind that D. quinquecirrha does not usually attain 48 marginal lappets and 40
tentacles until the medusa is 120 mm. in diameter, and it seems therefore that the so-called
Chrysaora of the Chesapeake is only a stunted Dactylometra which becomes mature in the
Chrysaora stage, and its pale coloration may be a local peculiarity due to unfavorable
conditions of confinement in brackish water. In the purer ocean water at the mouth of
Chesapeake Bay the medusz develop into the Dactylometra condition with 40 tentacles.
These conditions are also found in Narragansett Bay, Rhode Island, where in relatively pure
clean water the medusa have 40 tentacles, but in brackish estuaries they often become
mature with only 24 tentacles and are pale in color.

Fic. 373-—Dactylometra africana, after Vanhoffen, in Valdivia Expedition.

Dactylometra africana Vanhdffen.

Dactylometra africana, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p- 40,
taf. 4, fig. 20.

Disk 100 to 130 mm. wide. 6 well-developed marginal lappets and 5 long tentacles in
each octant. Lappets and tentacles red. Red radial streaks over exumbrella. C&sophagus,
palps, and gonads not highly colored. Colors of large specimens duller than those of small
meduse and not unlike the coloration of D. quinquecirrha. Distinguished by its red lappets.
Coast of German Southwest Africa. Common in the Great Fish Bay on October 10, 1898.

Dactylometra ferruginaster Kishinouye.

Dactylometra ferruginaster, Kisntnouye, 1892, Zoological Magazine, Tokyo, vol. 4, p. 264, taf. 3.
Dactylometra pacifica var. ferruginaster, Maas, 1909, Abhandl. Akad. Wissen. Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 44, taf. 2,

fig. 14.
Bell flatly rounded, 3 to 4 times as wide as high, 100 mm. in diameter. 48 oval, marginal
lappets. The 8 primary tentacles longer than the others. The lappets next to the ocular

SEMAEOSTOMEA3—DACTYLOMETRA, KURAGEA, 589

lappets are the smallest, and only about half as wide as the others. The 16 radial stomach-
pouches of nearly similar form and size. Mouth-arms longer than bell-radius and tapering
to pointed tips. Bell white with reddish-brown star on exumbrella. Inner sides of mouth-
arms and tentacles brown. Found on the coast of Japan in autumn. This form is imper-
fectly separated from D. quinquecirrha of which it appears to be a variety, if, indeed, it be not
identical with the American medusa.

“Dactylometra longicirra’”’ Kishinouye.

Dactylometra quinquectrrha var. pacifica, Gorter, 1886, Sitzungsber. Akad. Wissen. Berlin, Jahrgang 1886, p. 834.
Dactylometra longicirra, Kisninouye, 1892, Zoological Magazine, Tokyo, vol. 4, p. 261, taf. 2.

Bell flatly rounded, 3 times as wide as high and 75 mm. in diameter. 48 tongue-shaped
marginal lappets, all similar each to each, somewhat narrower near their bases than near
their outer ends. Accessory lappets sometimes seen on sides of ocular lappets. 40 tentacles
of nearly equal length each to each, and 10 times as long as bell-diameter; these tentacles
are wide, flat, and ribbon-like near their bases; sometimes one finds small accessory tentacles
arising from between the ocular lobes and their accessory lateral lappets or from the sub-
umbrella surface of the ocular lappets. “The medusa then has 56 tentacles, 7 in each octant
as in Kuragea depressa of Japan, and this illustrates the close genetic relationship which
exists between the two forms. In the middle the ocular stomach-pouches are twice as wide
and in their distal parts half as wide as the tentacular pouches. The mouth-curtains are
very wide and much folded; they are about 5 times as long as bell-diameter.

Bell white with 32 reddish-yellow radiating streaks. Mouth-arms yellow. Gonads and
tentacles reddish.

Found on the Pacific coast of Japan, in Owari Bay. Its common Japanese name is
Aschinaga Kurage, or Akakurage. I am inclined to believe that this is only a growth-stage of
Kuragea depressa, and that it is identical with D. ferruginaster.

Genus KURAGEA Kishinouye, 1902.
Kuragea, Kisuinouye, 1902, Journ. College Sci. Tokyo, vol. 17, art. 7, p. 9, plate 1, fig. 7.

The type species is K. depressa of Japan.

GENERIC CHARACTERS.

Pelagide with 8 marginal sense-organs. 87 (56) tentacles. 8X8 (64) marginal lobes.
4 interradial gonads.

This genus bears the same relation to Dactylometra that the latter does to Chrysaora,
being a stage wherein the lappets and tentacles have progressively increased by 16 in number.

Thus Chrysaora has 24 tentacles and 32 lappets, Dactylometra has 40 tentacles and 48
lappets, Kuragea has 56 tentacles and 64 lappets. I am inclined to look upon this form as an
hypertrophic Dactylometra rather than as a separate genus.

Kuragea depressa Kishinouye.

Kuragea depressa, Kisninovuye, 1902, Journ. College Sci. Tokyo, vol. 17, art. 7, p. 9, plate 1, fig. 7.

Umbrella 85 mm. wide and 30 mm. high. 8 marginal sense-organs. 87 (56) tentacles.
8X8 (64) marginal lobes. 4 gonads, each folded
in the form of the Greek letter ». Gastric
filaments long and numerous. Color (?) Misaki,
Japan. A single specimen. .

The 16 ocular lobes and the lobes by the
sides of the adradial tentacles are larger than
the others, while those adjacent to the ocular
lobes are the smallest. The lips are broad and
complexly folded. The exumbrella exhibits a
16-rayed, star-shaped marking in the inter-radi,
adradii, and perradii. The central stomach gives

Fic. 374.—Kuragea depressa, after Kishinouye, in Journal RISE KD 16 peripheral pouches ae uae Dactylometra
College of Sci., Tokyo University. longicirra. Inthe middle of their lengths the 8

590 MEDUS2 OF THE WORLD.

rhopalar pouches are 1.5 times as wide as the 8 tentacular pouches, but at the bell-margin
only half as wide.

This medusa is clearly derived from Dactylometra longicirra of Japan. D. lactea of
America tends to attain to the Kuragea condition, but all of the octants do not usually develop
equally (see text-fig. 369).

Genus SANDERIA Gitte, 1886.
Sanderia, Gorrte, 1886, Sitzungsber. Akad. Wissen. Berlin, Jahrg. 1886, p. 835.—VANHGFFEN, 1902, Wissen. Ergeb. deutsch.

Tiefsee Expedition, Dampfer, Valdivia, Bd. 3, Lfg. 1, p. 37-

Neopelagia (an aberration), Kisurnovye, 1910, Journal College Sci. Tokyo, vol. 27, art. 9, p. 14.

The type species is Sanderia malayensis Goette, of the tropical Indian Ocean and Malay
Archipelago.

GENERIC CHARACTERS.

Pelagide with 16 marginal sense-organs, 16 tentacles, and 32 cleft marginal lappets.
4 lips, 4 interradial gonads, and 32 peripheral stomach-pouches in the radii of the tentacles
and sense-organs. No marginal ring-canal.

Sanderia malayensis Goette.

Sanderia malayensis, Goretre, 1886, Sitzungsber. Akad. Wissen. Berlin, Jahrg. 1886, p. 835.—VANHOFFEN, 1902, Wissen. Ergeb.
deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 38, taf. 3, fig. 12; taf. 8, fign. 69-74.
Neopelagia eximia, Kisu1novye, 1910, Journal College of Sci. Tokyo, vol. 27, art. 9, p. 14, plate 3, fig. 15, text-fig. 1.
Bell flat, go mm. wide, large, rounded nettling-warts at the center of the exumbrella but
diminishing in size outwardly so as to be absent at the bell-margin. 16 marginal sense-organs

Fic. 375.—Sanderia malayensis, after Vanhoffen, in Valdivia Expedition.

alternating with 16 long, ribbon-like tentacles, 32 cleft lappets. A long, tubular cesophagus is
bounded on the 4 perradial corners by long, complexly folded lips. The 4 interradial, heart-
shaped, genital ostia are each bordered externally by about 24 to 30 finger-shaped papilla. The
central stomach gives rise to 32 peripheral pouches in the radii of the sense-organs and tentacles.
These are completely separated one from another by straight, radiating septa which converge
slightly near the bell-margin in the rhopolar radii. There is no marginal ring-canal. The gen-
eral color is yellow with the aboral center of the bell dusted over with reddish flecks which
extend outward, spoke-like, in the radii of the sense-organs and tentacles. The mouth-parts
are also covered with reddish spots.

SEMAEOSTOMEAS—SANDERIA, DESMONEMA. 591

Found in the Indian Ocean, Gulf of Aden, at Singapore, and off the east coast of Africa.
Some specimens of this medusa were found by the U. S. Fisheries Bureau Steamer Albatross
in the Philippine Islands in March and April, 1g08, and Kishinouye found it at Misaki,
Japan.

A perfect specimen found by the d/batross on March 8, 1908, at station D 5175 in the Sulu
Sea, southeast of Cagayanes Islands, Philippine Islands, hada bell 75 mm. wide, alte 46 long,
eel stomach 35 wide, contracted tentacles 65 long, and with 25 to 30 finger-shaped pro-
jections bordering each mantel ostium.

Kishinouye, 1910, found an abnormal specimen with 13 tentacles, 13 rhopalia, 26 lappets
and irregularly developed gonads. He found that a fish of the genus Psenes accompanied
the medusa.

Family CYANEIDZ L. Agassiz, 1862.

Cyanetde, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 114, 161.—Acassiz, A., 1865, North Amer. Acal., p. 44
HarckEL, 1880, Syst. der Medusen, p. 518; 1881, Challenger Report, Zool., Bd. 4, part. 2, p. 1244.—Craus, 1883, Organ.
und Entwick. Medusen, p- 24.—von LenpeENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p.271. ne LAUS,
1886, Arbeit. Zool. Inst. Univ. Wien., Bd. 7, p. 110.—VANHOFFEN, 1906, Nordisches Plankton, Nr. I1, p. 51.—Maas,
1906, Fauna Arctica, Bd. 4, Lfg. 3, p. 505.

FAMILY CHARACTERS.

Semaeostomez with a single, 4-sided, central mouth surrounded by 4 perradially situated,
curtain-like lips. The tentacles arise from the floor of the subumbrella, at some distance
inward from the margin, and are usually in clusters. The gonads are situated in 4 complexly
folded, interradial outpocketings of the wall of the subumbrella. The central stomach gives
rise to radiating, peripheral pouches, which in turn give rise to numerous branching, non-
anastomosing, blind canals in the lappets. There is no ring-canal. The tentacles are hollow.

The medusz of this family are apparently descended from some such forms as the Pelag-
ide. They resemble the Pelagidz in the structure of the oral appendages, the general plan
of the gastrovascular system, and in the lobulation of the bell-margin. They differ mainly
in the complex branching of the peripheral edges of the radiating stomach- -pouches, and
above all in that the tentacles arise from the fear of the pati brelia: not from the notches
between the lappets as in the Pelagide. The young medusz are strikingly similar to the
Pelagidz in all respects, for their radiating stomach- -pouches are simple and the tentacles
first appear in the notches between the lappets. The margin grows beyond the bases of the
tentacles as development proceeds, however, and thus they come secondarily to arise from
the floor of the subumbrella. Indeed the tentacles of all Scyphomedusz are structures of the
subumbrella.

In Cyanea the development is known to be through a sessile scyphostoma which strobi-
lates, giving off a number of ephyra which develop into mature meduse. The Cyaneide are
of universal distribution, but the great majority of the species are found in the temperate
regions and in the colder waters. Unlike the Pelagide the Cyaneide are creatures of the
shallower waters along shores, not animals of the high seas.

A synopsis of the genera of the Cyaneidz follows:

Desmonema L. Acassiz, 1862. 8 rhopalia, 8 adradial clusters of tentacles. No radial-muscle strands in the subum-
brella.

Cyanea Péron Et Lesueur, 1809=Procyanea+ Medora+ Stenoptycha+ Desmonema (in part)+ Cyanea, Harcke1,1880.
8 rhopalia, 8 adradial clusters of tentacles; each cluster contains several rows of tentacles. Both radial and circular
muscles in the subumbrella.

Drymonema Harcxet, 1880. 8 rhopalia. Tentacles not grouped in isolated clusters, but arising from a wide zone in
the subumbrella.

(?) Patera Lesson, 1843 (doubtful). 16 rhopalia. 16 clusters of tentacles alternating with the radii of the rhopalia.

Genus DESMONEMA Agassiz, 1862.

Couthouyia, used for Mollusca by Adams, 1860, Annal. and Mag. Nat. Hist., vol. 5, p. 410.

Couthouyia, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 118, 163.

Couthouya, Maas, 1906, Fauna Arctica, Bd. 4, Lfg. 3, pp. 487, 505; 1908, Expédition Antarctique Frangaise, Meduses, p. 3.

Desmonema, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 166.— HarckeL, (in part), 1880,Syst.der Medusen, p. 526.
—LeEnpDEnFELD R., von., 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 273-—VANHOFFEN, 1888, Bibliotheca Zoo-
logica, Bd. 1, Heft. 3, p- 17.—Browne, 1908, Trans. Royal Soc. Edinburgh, vol. 46, p. 242.—VANHOFFEN, 1908, Deutsche
Siidpolar Exped., Bd. 10, Zool. 2, p. 40.

Medora (young stage), AGAssiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 118.

Medora, preoccupied for Mollusca by Adams, 1858, Genera of Mollusca, vol. 2, p. 183

592 MEDUS OF THE WORLD.

GENERIC CHARACTERS.

Cyaneide with 8 marginal sense-organs and with 8 adradial clusters of marginal tentacles.
The tentacles of each cluster may arise in several rows from the subumbrella surface. There
are 8 chief lobes and 16 to 32 secondary lappets. Without radial-muscles in the lobes.

The names Couthouyia and Medora are preoccupied, but Agassiz’s Desmonema may be used.

The oldest species is the little-known “Chrysaora” gaudichaudu erroneously described
by Lesson, 1829 (Voyage de la Coquille, Zooph., p. 114), from the region of Cape Horn, South
America, and recently revealed through the studies of Maas, 1908.

Some of the medusa which Haeckel included in his genus ‘‘Desmonema’’ represent only
immature Cyaneas in which the tentacles of each cluster arise in a single row. Vanhoffen, 1888,

Fic. 376.—Desmonema gaudichaudii, after Maas, in Meduses Expédition Antarcticque Frangaise.

shows, however, that there are considerable anatomical differences between Cyanea in any
stage and Desmonema. In Desmonema, for example, the 8 clusters of tentacles arise in linear
arcs concentric with the general contour of the bell-margin, whereas in Cyanea they arise in
crescents, the horns of which are directed outward toward the margin. Moreover, there are
no radial-muscle strands in the 8 velar lobes of Desmonema, but these are found in Cyanea.

Desmonema annasethe Haeckel is clearly a young Cyanea, as was pointed out by Van-
hoffen, 1888.

The rediscovery of Lesson’s, Cape Horn, medusa by Maas makes it practically certain
that ‘ is specifically referred to by Agassiz, 1862, p. 118, under the designation Couthouyia
pendu a,

SEMAEOSTOME A2—DESMONEMA, 593

Demonema gaudichaudii Agassiz.

Chrysaora gaudichaudii, Lesson, 1829, Voyage de la Coquille, Zooph., p. 114, Zoophytes planche 13, fig. 1.

Couthouyia pendula and Medora capensis, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 118, 163.
Desmonema gaudichaudt, Acassiz, L., 1862, Cont. N at. Hist. sie S., vol. 4, p- 166.—HarckEL, 1880, Syst. der Medusen, p. 527.
Couthouya gaudichaudi, Maas, 1908, Expé ee Antarctique Frangaise, Meduses, p. 5, planche 1, fig. 1.

Bell at least 500 or 600 mm. wide, only 5 to 7 tentacles in a single line, in each inter-
rhopalar cluster. Thus when the medusa is of a greater size than D. chierchiana, it has fewer
tentacles. Maas states that the color of the gastric cavity is brownish- -purple, accentuated
in the gonads, while the muscle-system is lighter and the gelatinous substance is bluish and
transparent. The medusa is found in the Antarctic region in April. Future studies will
probably demonstrate that D. chierchiana is only a variety of this medusa for they are alike
in all respects excepting that in Desmonema chierchiana the tentacles arise in several rows,
whereas in D. gaudichaudii they remain as a single row even when the medusa is much larger
than D. chierchiana. ‘This difference may be due to environmental causes, or may be of
the nature of a variation such as one observes in the development of tentacles in Dactylometra
and Chrysaora, but until further studies have been carried out it will be safer to keep the
two forms specifically distinct one from the other. It is probable, however, that D. gaudi-
chaudu is only a variety in which the tentacles remain in a single row, as in the young of D.
chierchiana (see fig. 376).

Further details of the structure of D. gaudichaudi may be obtained from the description
of D. chierchiana which follows.

Agassiz’s Medora capensis is apparently a young stage of this medusa.

Desmonema chierchiana Vanhdffen.

Desmonema chierchiana, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft 3, p. 18, taf. 1, fig. 4; 1908, Deutsche Siidpolar
Expedition, 1901-1903, Bd. 10, Zool. 2, p. 41, taf. 2, fign. 2, 3; Abbild. 5-9.—Browne, 1908, Trans. Royal Soc. Edinburgh,
vol. 46, p. 244, plate 2, fig. 2.

(??) Chrysaora gaudichaudii, Lesson, 1829, Voyage de la Coquille, Zooph., p. 114, planche 13, fig. 1.

Umbrella 310 mm. or more wide and too mm. high. Exumbrella smooth. The 8 pairs
of ocular appers are only about one-third as wide as ate 8 tentacular lappets, which are flatter
than a semicircle in outline. The 16 small, ocular lappets are bluntly rounded and sharply
set off from the 8 tentacular lappets. The subumbrella is divided into 16 equal sectors cor-
responding to the 16 stomach-pouches. ‘These sectors are areas in which the circular muscles
are well developed and separated one from another by narrow septa. 8 of the 16 muscular
sectors are in the radii of the 8 marginal sense-organs and 8 are intermediate. There are no
radial-muscle strands.

There are 8 groups of tentacles, each of which, in old medusz, arises in several rows
from the subumbrella at some distance inward from the bell-margin. These tentacles are
developed along the outer edges of the 8 interocular, circular muscle-sectors. There are
about 60 tentacles in each cluster, the oldest and longest being along the innermost, and the
shortest and youngest in the outermost rows. The rows are not U-shaped, as in Cyanea, but
are nearly straight. The tentacles are hollow, their entoderm being thin and the ectoderm
thick-walled. When young the tentacles are somewhat flattened and the ectodermal longi-
tudinal muscle strands are set within infolded, gutter-like lines down the length of the ten-
tacle. As growth proceeds these gutters sink deeper and deeper into the ectoderm and finally
become inclosed tubes, sunken beneath the surface. Thus the longitudinal muscles become
tubular strands of fibers. A full description of the tentacles is given by Vanhoffen, 1908.
When expanded the tentacles of a large medusa may be at least 20 meters long.

The 4 mouth-curtains are well- developed and resemble those of Chrysaora. They are
narrow near the mouth- -opening but expand outwards and then taper gradually to a point.
Each mouth-curtain is 1.5 times as long as the bell-radius and its margins are much folded.

The 4 sac-like, protrusive gonads are folded and resemble those of Cyanea, but are smaller.

There are 16 sectors of circular muscles in the subumbrella, 8 in the ocular and 8 in the
interocular radii. These circular muscles do not extend beyond the zone of the clusters of
tentacles and sense-organs and are not found in the subumbrella of the lappets. There are
no strands of radial-muscles in the subumbrella, or in the lappets, such as are seen in Cyanea.

594 MEDUS.2 OF THE WORLD.

The 16 stomach-pouches break up into numerous forked, branched, radiating vessels
in the marginal lappets, but their ultimate branches rarely anastomose. There are about
12 to 18 main branches from the outer edge of the stomach-pouch in each lappet and these
branch dendritically so that about 100 ramuli reach the bell-margin. There is no marginal

ring-canal.

“3 FS xe

Fic. 377-—Desmonema chierchiana, after Vanhoffen, in deutsch. Siidpolar Expedition.

The medusa is salmon-red or brownish-red, the canal-system being darker and the tenta-
cles light in color. When young the bell and tentacles are bluish and the curtain-like lips
salmon or brownish-red in color.

SEMALEOSTOME

DESMONEMA, CYANDA, 595

The medusa appears to be common from December to June in the Antarctic region, and
is reported from both sides of the Straits of Magellan, Kerguelen, and the Falkland Islands;
and from Kaiser Wilhelm Coast, South Victoria and Alexander I Land along the ice-edge
of the Antarctic continent.

Ephyre 3 to 10 mm. wide are found in January and February, and Vanhdffen records a
young medusa in the Medora stage from Gauss Station, Kaiser Wilhelm Land on April 14.
This medusa was 38 mm. in diameter, the mouth-arms 16 mm. long. There were 8 principal
tentacles about two-thirds as long as bell-diameter, and 4 of these were bordered on one side
by a small tentacle of recent development, figure 379. The lips and gastric cirri were
brownish-red, other parts being translucent milky-blue. A later stage is described by Browne
(see figure 378).

Genus CYANEA Péron and Lesueur, 1809.

Cyanea, Péron anv Lesueur, 1809, Ann. Mus. Hist. Nat. Paris, tome 14, p. 363.—Escuscnoxtz, 1829, Syst. der Acalephen,
p- 67.—Lesson, 1843, Hist. Zooph. Acal., p. 379.—Branpt, 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome
4, P- 77-—Fornes, 1848, British Naked-Eyed Meduse, p. 77.—Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 115,
161.—Acassiz, A., 1865, North Amer. Acal., p. 44.—von LenDENFELD, 1882, Zeit. fiir wissen. Zool., Bd. 37, p. 465; 1884,
Proc. Linnean Soc. New South Wales, vol. 9, p. 275.—Bicrtow, R. P., 1900, Mem. Boston Soc. Nat. Hist., vol. 5, p. 211.—
Haraitt, C. W., 1902, American Naturalist, vol. 36, p. 555 —Maas, 1904, Résult. Camp. Sci. Prince de Monaco, fasc. 28,
P- 55; 1906, Fauna Arctica, Bd. 4, Lfg. 3, pp. 486, 505; 1903, Scyphomedusen der Stboga Expedition, Monog. 11, p. 28.—
VANHOFFEN, 1906, Nordisches Plankton, Nr. 11, p. 51-

Procyanea+ Medora+ Stenoptycha+ Desmonema (in part)+ Cyanea, Harcket, 1880, Syst. der Medusen, pp. 524-528.

The type species is C. capillata of the North Atlantic, Pacific, and Arctic Oceans. It is
the largest of all known medusz.
GENERIC CHARACTERS.

Cyaneide with 8 marginal sense-
organs and with 8 adradial crescentic
groups of tentacles. Each group consists
of several rows of tentacles. With radial
muscle strands in the subumbrella.

ASS =

When young only 8 simple tentacles
arise in the adradial clefts between the
ephyra lobes, but later the margin grows
beyond them, leaving them to project from
the floor of the subumbrella. In the mean-
time the tentacles increase in number,
becoming a row in each adradius, but
finally they come to lie in two or more
rows. Haeckel has constituted a special
genus for each of these growth-stages.
He calls the 8-tentacled stage“* Procyanea.”
The stage with 24 tentacles, 3 in each
adradius, he names ‘‘ Medora,’ and when
there are 5 tentacles in each rowthe medusa
becomes a “‘Stenoptycha”; then as long as
the medusa remains with the tentacles of
each cluster in a single row it is a “‘Des-
monema,’ and finally when older and the
tentacles begin to develop in two or more
rows in each cluster the medusa is called a
Cyanea. Itis possible that some meduse
may become mature in, and never advance

Desmonema chierchiana. ) =
Ba) b oe
Fic. 378.—After Browne, in Trans. Royal. Soc. Edinburgh. beyond, Haeckel’s Desmonema stage,

Fic. 379.—Young medusa, after Vanhoffen in deutsch. Siidpolar }yt it is certain that others pass through this
Expedition. :

condition and become mature as Cyanea.

Medusz of Cyanea are abundant in the Arctic and Antarctic, but are not found in the
tropics. Being dependent upon a fixed scyphostoma-stage for development, they are confined
to the proximity of coasts where the water is relatively shallow.

596 MBEDUS# OF THE WORLD.

The early development of the planula takes place among the folds of the copious mouth-
curtains of the adult medusa. Segmentation is total and regular but unequal, the cells of one
pole being smallest. The gastrula results from simultaneous delamination and invagination
at the small-cell pole. The blastopore closes. The planula attaches itself by its forward end
and becomes a scyphostoma which acquires 15 to 20 tentacles and strobilates producing a
number of ephyre. The details of this development are given under C, capillata.

It appears that the numerous so-called species of Cyanea intergrade to such a degree
that we can not maintain them, and I believe there are only two species: C. capzillata of the
north temperate and Arctic regions and C. annaskala of the south temperate and Antarctic.
In common with Pelagia, Chrysaora, Dactylometra, Aurellia and other world-wide forms of
meduse, growth-stages, color varieties and local races have frequently been described as
separate species, but as our knowledge increases many intergrading forms come to light thus
reducing the so-called species to a few dominant types with numerous, closely related offshoots.
It is unfortunate that the aim of the old systematic zoology was mainly toward the empha-
sizing of distinctions rather than the indication of affinities and the discovery of relationships.

Cyanea is readily distinguished from Desmonema by its radiating muscle strands in the
subumbrella, and its horse-shoe shaped clusters of tentacles.

Cyanea capillata Eschscholtz.
Plate 65, figs. 3 and 4.

Medusa capillata, Linné, 1746, Fauna Suecica, Ed. 1, p. 368, Nr. 1286; Systema Natura, Ed. 10, 1758, tome 1, p. 660; 1788,
tomus 1, pars 6, p. 3154.—Fasricius, 1780, Fauna Groenlandica, p. 364.

Cyanea capillata, Eschscholtz, 1829, Syst. der Acalephen, p. 68.—Van Benepen, 1886, Fauna. littor, Belg., p. 77, taf. 1, 2.—

Ec age Haecker, 1880, Syst. der Medusen, p. 529.—Hamann, 1890, Internal. Monatsschrift Anat. Physiol., Bd. 7, p. 259,

®, =, taf. 12.—BrowneE, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p- 775-—Maas, 1906, Fauna Arctica, Bd. 4, Lfg. 3, pp. 486,

* §11.—Hott, 1902, Report Fisheries of Ireland, part 2.

Cyanea artica, Péron Ev Lesueur, 1809, Ann. Mus. Hist. Nat. Paris, tome 14, p. 363.—Morcu, 1857, Beskriv. af Groénland,
Pp- 95-—Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, pp. 87, 162; Ibid. 1860, vol. 3, plates 3-5a; plate 10, figs. 1-17,
19-21, 23-30, 33-35) 37) 38; plate 10a, figs. 1-4a, 5-124, 14, 15, 17-40.—PAcKARD, 1863, Canadian Naturalist and_Geol.,
vol. 8.—Acassiz, A., 1865, North Amer. Acal.,
P- 44, fig. 67.—Harcker, 1880, Syst. der
Medusen, p. 530.—Frwkes, 1881, Bull. Mus.
Comp. Zool. at Harvard College, vol. 8, No. 8,
p. 166, plate 7, figs. 1, 4, 5, 8-14.—Wacner,
1885, Wirbellosen des Weissen Meeres, Bd. 1,
p- 83, taf. 5, 6-—MacMurricn, 1891, Amer.
Naturalist, vol. 25, p. 287.—Haraitr, 1902,
Science, ser. 2, vol. 15, p. 571.—MAcALLUM,
1903, Journal of Physiology, Cambridge, Eng-
land, vol. 29, pp. 213-241.—Hypr, 1894, Zeit.
fiir wissen. Zool., Bd. 58, p. 531, taf. 34, fign.
54-62; taf. 35, fign. 63-79; taf. 36, fign. 94-96.
—Harairtr, 1904, Bull. U.S. Bureau of Fisher-
ies, vol. 24, p. 68.—Maas, 1904, Résult. Camp.
Sci. Prince de Monaco, fasc. 28, p. 56.—Van-
HOFFEN, 1906, Nordisches Plankton, Nr. 11, p.
53, fign. 16-19; C. capillata, p. 52, fig. 15; C.
lamarcki, pp. 53, 64, fign. 35-37-—Harairt,
1905, Journal Experimental Zool., vol. 2, p.
574 (variations).

Cyanea lamarchti, Péron er Lesurur, 1809, Ann.
Musée Hist. Nat., Paris, tome 14, p. 363.
Cyanea postelsii, Goutv, A. A., 1841, (non Brandt)
Report Invert. Massachusetts, p. 347.—S1iMp-
son, 1853, Marine Invert. Grand Manan, p. 11.

Cyanea — » M’Kenprick, J. G., 1881, Journal
Anat. and Physiol., vol. 15, p. 261 (coloring
matter).

Cyanea lamarcki, Detar, 1905, Annual Report Fish-

Fic. 380.—Cyanea capzsllata, after Vanhéffen, in Nordisches Plankton. eries of Ireland 1902-03, part 2, Appendix 1

(reared from the egg in an aquarium),

Cyanea lamarckit, HarcKer, 1880, Syst. der Medusen, p. 530 (literature).— Jounin, 1906, Bull. Musée Oceanograph., Monaco,
No. 66, p. 27, fig. 28 (after Delap).—Vanudrren, 1908, Deutsche Stidpolar Expedition, Zool. 2, Bd. 10, p. 39-

Cyanea lamarckii=C. capillata, M. Inrosu, 1885, Annals and Mag. Nat. Hist., ser. 5, vol. 15, p. 148.

For literature of C. fulva and C. versicola see the detailed account of these varieties.

Cyanea ferruginea, Escuscuortz, 1829, Syst. der Acalephen, p. 70, taf. 5, fig. 1.

Cyaneopsis behringiana, BRANDT, 1838, Mém. Acad. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 379, taf. 11, fig. 1.

Cyanea ferruginea, Harcker, 1880, Syst. der Medusen, p. 531.—Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11,
p- 28.—Vanuirren, 1906, Nordisches Plankton, Nr. 11, p. 55, fig. 20.

PLATE 65.

Fig. 1. Cyanea capillata var. versicolor, mature medusa, natural size. In
the ocean off St. Catherines Sound, Georgia, December 29, 1904.
Two of the oral palps have been removed in order to reveal the
structures of the subumbrella.

Fig. 2. Cyanea capillata var. versicolor, young ephyra. 2.5 mm. in diameter.
Surface tow off Cape Fear, North Carolina, December 1, 1904.

Fig. 2’. Sense-organ of the ephyra shown in figure 1.

Fig. 3. Cyanea capillata, mature medusa, half natural size. Oral view of
quadrant of exumbrella, with mouth-parts and gonads removed.
Biological Laboratory, South Harpswell, Casco Bay, Maine,
August 28, 1908.

Fig. 4. Cyanea capillata, young ephyra, 3.5 mm. in diameter. gt. internal
gastric cirri. Agassiz Laboratory, Newport, Rhode Island, June
29, 1893.

Drawn from life, by the author.

MAYER
PLATE 65

AHoenk lo

~I

SEMA EOSTOMEAS—CYANEA. 59

(22) Melusina formosa, Haecxer, 1881, Report Deep-Sea Meduse Challenger Exped., Zool., vol. 4, p. 1-

Cyanea postelsis, BRanpt, 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 375, taf. 12, 13, 13a.—HarcxeL,
1880, Syst. der Medusen, p. 532.—VANHOFFEN, 1906, Nordisches Plankton, Nr. 11, p. 55, fig. 21.—Maas, 1906, Fauna
Arctica, Bd. 4, Lfg. 3, p. 506.

Cyanea tmporcata, Norman, 1865, Nat. Hist. Trans., Northumberland and Durham, vol. 1, p. 11, 1 taf. (a young Cyanea from
the British coast).

Cyanea citrea, KisHINOUYE, 1910, Journal College of Sci. University Tokyo, vol. 27, art. 9, p. 16, plate 4, figs. 16, 17 (this is C.
ferruginea).

It is practically impossible to draw any fixed distinctions between the various forms of
the great Cyanea of the North Atlantic. Intergrading forms are commonly met with and
many of the races are separated only geographically or upon color distinctions which are
neither wholly characteristic nor stable. Medusa capillata Linné, 1746, is the oldest name
applied to any of these meduse. The following are probably all varieties of one and the same
species, C. capillata:

Cyanea capitllata. Bell about 500 to 1,200 mm. wide. Ocular and interocular clefts of bell-margin not sharp and narrow,
but with evenly rounded curves. Bell, palps, and tentacles reddish or yellowish-brown, with rose-colored or red gonads.
Vanhdffen, 1906, states that when the bell is 13 mm. wide there are 7 tentacles in each adradial cluster, the middle
one of each group being the longest. When 20 mm. wide there are 15 tentacles in each cluster with 3 long ones
in the middle. When 41 mm. wide there are 35; and when 86 mm. there are 63 tentacles. This variety is not un-
common in the English channel, North Sea, and coast of Norway in summer and autumn.

Var. /amarckii. Distinguished by the decided blue color of the bell and palps, the color being lighter at the margin
than at the center of the bell. Gonads and tentacles nearly white. In other respects this form resembles C. capillata.
Vanhoffen states that when the medusa is 43 mm. wide there are only 20, and when 85 mm. wide only 31 tentacles
in each group, thus being only about half as many as in the typical C. caprl/ata of the same diameter. On the other
hand the gonads in a medusa of C. /amarcki 43 mm. wide are about as long as the palps, thus being larger than in
C.capillata. This form is found in the English channel along the coasts of France and Great Britain, at Helgoland,
and in the entrances to the Baltic, but it does not extend into the Baltic. It becomes 300 mm. wide.

Var. arctica. Supposed to be distinguished from the first two forms by the indistinctness of its rhopalar tappets which
do not project as far beyond the general contour of the bell-margin as in C. capillata. It is very large, though speci-
mens over 800 mm. in diameter are rarely met with. The bell is very variable in color but is usually rich brown and
yellow, with deeply colored gonads and rich rosin-yellow muscles and tentacles. It is found off the American coast
north of Cape Cod, where it is abundant during the summer, becoming mature and disappearing in early autumn.

LBD

SID
Ss

ES fe

ge OF

i
1
TA
ately

leetrrs pre) >
i ey
sy steranieee
2,

Fic. 381.—Cyanea ferruginea, according to Eschscholtz, after Vanhoffen, in Nordisches Plankton.

Var. fulva. A small yellowish-colored variety of C. arctica which ranges from Cape Cod southward to the Carolina
coast. It is rarely over 200 mm. wide. At Newport, Rhode Island, it becomes mature early in June and is not seen
in July, but on the coast of New Jersey it is found in August.

Var. nozakit. Similar to C. fulva, but milk-white in color. Found in the Inland Sea of Japan.

Var. versicolor. A pink-colored southern variety of Cyanea fulva, which is even smaller, rarely more than 110 mm.
wide. It forms large swarms during the winter and spring off the coast of the United States between Cape Hatteras,
North Carolina, and southern Florida.

Var. ferruginea. Of the North Pacific coasts of America and Asia. It is a variety of C. cap#llata and is apparently identical
with the variety C.“‘arctica.”” It becomes about 450 mm. wide and is light-yellow or orange with the stomach and radial
pouches light-brown. The gonads are yellow and the tentacles reddish, although these colors are probably somewhat
variable as in other forms of Cyanea.

Var. postelsii appears to be only a local variety of C. capsljata. It is found in the North Pacific from the Aleutian
Islands to Oregon and is distinguished by the 16 well-developed clefts which flank the rhopalar lappets and are about
half as deep as the adradial clefts; moreover, according to Mertens, the contours of the lappets are evenly rounded
and even the clefts are not narrow, but widen inwardly with rounded contour. A modern study of this medusa is
to be desired, for it is possible that Mertens over-emphasizes these peculiarities. The medusa is found between
Sitka and the Aleutian Islands, Alaska.

The following is a detailed description of Cyanea “arctica” of the coast of North America:
Adult medusa.—The disk is quite flat and lenticular and attains a diameter of 2,300 mm.
Medusz of this size are very rare, however, and the majority are not over 800 mm. wide. The

598 MEDUS.Z OF THE WORLD.

umbrella margin is divided by 8 deep, adradial clefts into 8 principal lobes, which are about
twice as broad as they are long. Each of these lobes is in turn divided by a median cleft, and
there are also two slight notches upon the bell-margin on either side of the median cleft;
the margin, therefore, displays 32 indentations, between which there are 32 lappets. The
margin of the bell is sharp-edged for the gelatinous substance, which is quite thick at the
center of the disk, becomes very thin as one approaches the periphery. The 8 marginal sense-
organs are found at the bottom of the median niches of the 8 principal lobes of the disk. Each
sense-organ is elongate and club-shaped, and protected above by a web which stretches between
the adjacent lappets; proximal half of club quite thick, with a well-developed swelling upon
its lower (subumbrella) side; this swelling is covered with wart-like elevations and provided
with one or two papilla. Distal to this swollen region the club extends outward as a cylindrical
tube which terminates in a swollen knob-like part containing an entodermal mass of crystalline
concretions, but no ocelli. “Two open pits project downward from the floor of the exumbrella
on either side of the base of the sensory-club. The structure of the sense-organ in Cyanea
has been studied by L. Agassiz, 1862; Eimer, 1878; and Fewkes, 1881. About 800 long ten-
tacles arise from 8 adradial, crescentic regions on the floor of the subumbrella, about midway
between the periphery and the center. The horns of these crescentic areas point outward
and the tentacles are arranged in about 5 concentric rows in each crescent, the oldest and
longest tentacles being on the innermost row. The tentacles are hollow and highly contractile;
when fully expanded they attain a length of about 25 times the bell-diameter; their surfaces
are thickly covered with nematocysts. Mouth 4-cornered and situated at center of subum-
brella; it is provided with 4 long perradial mouth-arms, the margins of which are greatly folded,
forming the curtain-like lips or oral fringes which hang downward in the water. Mouth-arms
about as long as bell-diameter, and with their fringes appear as a complexly folded, contractile
mass of curtain-like appendages hanging from the oral floor of the bell.

EAE, CORON
< es

SS —~ y

Fic. 382.—Cyanea postelsi, according to Brandt, after Vanhéffen, in Nordisches Plankton.

Gonads occupy 4 complexly-folded pouches which project from subumbrella floor at the
4 interradial sides of the stomach. Numerous clusters of small gastral cirri project from the
bases of the gonads into the stomach-cavity; these are far more prominent in the young
medusa than in the adult, for in the mature medusa they become hidden away at the bases
of the pendant, pouch-like folds of the genital organs. There is a very powerful and con-
spicuous system of circular muscles in the subumbrella; these muscles occupy a zone about
one-eighth as broad as bell-radius and which lies adjacent to and centrifugal from the gonads.
This zone of muscles is composed of 16 trapezia, the 8 in the rhopalar radii being only half
as wide as those in the adradit. 16 strands of radiating muscles extend from the outer side of the
zone of circular muscles and pass outward on either side of the sense-organs.

The central stomach is a wide, lenticular space in the center of the disk; peripherally it
gives rise to 16 radiating pouches, the outer edges giving numerous branched canals which
ramify through the lappets without anastomosing. The 8 pouches in the radii of the sense-
organs are less than half as wide as the 8 in the radii of the tentacles. The tentacles and the
stalks of the sense-organs are hollow and in direct connection with the gastrovascular space of

SEMAEOSTOMEA—CYANBA. 599

the medusa. There is no ring-canal. The gonads are great hollow bags forming part of the
gastric system of the ode There are deep clefts in the aboral floor of the stomach ¢ giving
it a reticulate appearance (see fig. 3, plate 65).

The gelatinous substance of the bell is translucent with a slightly bluish or yellowish tinge.
The entoderm of the gastrovascular system is of a rich brownish- -purple and the mouth-arms
and oral curtains are chocolate-purple. The gonads and tentacles are either yellowish or
reddish-brown, and the muscular system of the subumbrella is brown or yellow.

This species extends from the southern coast of New England northward to the Arctic
Ocean. It thrives best in the colder waters, and specimens found south of Cape Cod are
usually of small size. It is worthy of notice, also, that south of Cape Cod the medusz dis-
appear about the middle of June, while in the cold waters of the coast of Maine the mature
ones are most abundant in August and September. In Europe it is abundant off the coasts
from France to Northern Russia, and is found at Spitzbergen in August.

Cyanea arctica appears to be identical with the so-called C. lepine a of the North Pacific;
and C. postelsir of the Pacific is a closely allied form.

The embryonic and larval stages have been studied by L. Agassiz, 1862; Fewkes, 1881;
Hamann, 1890; MacMurrich, 1891; and Ida Hyde, 1894. Agassiz gives a series of igeies
illustrating the general developmental stages of the planula and scyphostoma, while Hy de
gives a very Copies: account of the histology of the early stages. The eggs are orange- -colored
and provided with a membrane, and are dehisced from tas ovaries into the gastric cavity,
where they undergo segmentation among the folds of the mouth-arms and ‘finally escape
through the mouth of the parent medusa as free-swimming planule. The segmentation is
total but unequal, the cells at one pole being smaller than those at the opposite pole. A
blastula is formed in which there is a large central blastoccel. The gastrula results from the
rapid divisions of one or two small cells at the small-cell pole, which form a layer that
invaginates. Hyde finds no wandering inward of free cells, but McMurrich records this
condition. The blastopore then closes over and the entoderm becomes entirely enveloped by
the ectoderm. In this condition the larva becomes a pear-shaped, ciliated planula and
swims actively through the water, the posterior, narrow end being that at which the gastrula
mouth had developed. One sometimes observes nematocysts in both ectoderm and entoderm
at this narrow hinder end of the planula. The next stage in development is instituted by the
formation of a shallow, crater-like, glandular invagination of the ectoderm at the broad,
anterior pole of the pear-shaped planula, and then the animal sinks down and attaches itself
to the bottom by this forward end. A cup-like depression of ectoderm then presses down
upon the entodermal sac at the narrow posterior end and finally fuses with it, and eventually
the mouth breaks through at this point.

The first pair of the radial pouches is formed from the entoderm, the second, in part at
least, from the ectoderm of the crater. MacMurrich, 1891, and Hargitt, 1902, observed that
planula in confinement encysted themselves during this stage, remaining thus for several
days until the mouth is about to break through, when the embryos emerge from the cyst
through a circular aperture at the center of its free, convex surface. Hyde, 1894, observed
this, however, only in one embryo and it is possibly an abnormal condition due to unfavorable
surroundings. Simultaneous with the formation of the mouth 4 tentacles make their appear-
ance, and the scyphostoma finally acquires 15 to 20 tentacles. Hargitt, 1902, finds that lateral
stolons are sometimes produced by the scyphostoma, and secondary scyphostomz bud out
from these stolons. A number of ephyra result from strobilization of the scyphostoma, and
this may occur in 18 to 20 days after the planula has attached itself but this period varies
considerably.

The young ephyra 3.5 mm. in diameter (plate 65, fig. 4) has a simple 4-cornered mouth
at the center of the subumbrella, and 4 smooth-edged, slightly raised lips. The 8 tentacular
notches in the margin are much wider and deeper than the notches of the sense-organs. The
tentacles arise from the bell-margin, but as the animal grows the margin extends beyond
them and they thus come to project from the subumbrella floor of the disk. 4 short, ento-
dermal gastric cirri (gt plate 67, figs. 2, 3) are found upon the oral floor of the subumbrella
near the interradial corners of the mouth and project into the stomach-cavity. The gastric
system in this stage consists of a wide, lenticular, central stomach from which there efend
outward 16 simple, radiating pouches in the radii of the tentacles and sense-organs. In

600 MEDUS-£ OF THE WORLD.

later stages the young medusa develops an increasing number of tentacles and the lips form
long curtain- like folds surrounding the 4-cornered mouth: When the young medusa is about
7 mm. in diameter there are a manibee of slender papilla upon the exumbrella and these are
clustered especially at the aboral apex. In this stage the medusa rarely comes to the surface,
but frequently spreads its oral fringes out over the bottom or sides of the aquarium and
remains sedentary. The same bavi is exhibited by the closely allied “Cyanea fulva” which
is represented in figs. 1 to 7, plate 66, and figs. 1 to 3, plate 67; and it is probably due to
some such habit that the young are rarely to ie found upon the surface while the large and
mature medusz are very abundant. The scyphostoma and young medusa feed upon pro-
tozoa, starfish, and mollusk larve.

Macallum, 1903, studied the composition of the body-juices of Cyanea arctica and found
them to be as follows:

]
| Na. | Ca. | K. | Mg.
== : z |
| Contentsiof sea-water's<..,eteis/ciieis ais scolarisiein <)>) eteagiot eel trae lip adatOe “ 3-66 | 11.99
| Content of body-juices of Cyanea arctica ............+...- | 100 | 2250) |e 707, | 11.31

The SO, is less in Cyanea than in sea-water by about 32 to 36 per cent and the medusa
contains more iron and less iodine than does sea-water.

M’Kendrick, 1881, studied the chemical composition of the coloring matter of Cyanea
and found that the blue pigment of Cyanea and Aurellia is in the form of granules surrounded
by clear protoplasm. This pigment is soluble in acids, but is precipitated in neutral or acid
solutions. Hence when the medusa becomes acid through decomposition after death the pig-
ment dissolyes out into the water, but during life it remains stable. This pigment matter of
Cyanea shows two absorption bands in the spectrum, one in the red and one in the orange,
very much as in Stentor ceruleus.

Holt, 1902, finds that in the North Sea this medusa is accompanied by young whiting.

Cyanea capillata var. fulva, L. Agassiz.
Plate 66, figs. 1 to 7; plate 67, figs. 1 to 3.
Cyanea fulva, Agassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 119, 162.—Agassiz, A., 1865, North Amer. Acal., p. 46.

This southern variety is distinguished from the northern C. arctica by the light yellow or
yellow- -brown color of the entoderm of its gastrovascular system, which is never rich brown,
as in the northern C. arctica. It is also much smaller, being rarely over 200 mm. in diameter.
The lappet notches are more uniform than in C. arctica and the tentacles are much less numer-
ous. The oral fringes, also, are less voluminous and by no means so complexly folded as
in C. arctica. This variety appears in great numbers early in May on the southern coast
of New England, and the medusz arrive at maturity about the middle of June, after which
they suddenly disappear. We have, however, met with swarms of them about 20 miles off
Barnegat Bay, New Jersey, early in August. This variety has not been taken north of Cape
Cod. The development is similar to that of the closely allied C. arctica.

We may regard this as a local race of C. arctica, which ranges from Cape Cod, Massa-
chusetts, southward to the Carolinas; where it is replaced by a still more southerly variety,
C. arctica var. versicolor.

Cyanea capillata var. versicolor L. Agassiz.
Plate 65, figs. 1 and 2.
Cyanea versicolor, Agassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 119, 162.—Agassiz, A., 1865, North Amer. Acal., p. 46.

This form bears the same relation to Cyanea arctica var. fulva as fulva does to the northern
C. arctica. It is smaller than fulva, but is distinguished especially by its pink coloration.
Mature medusz are about 110 mm. in diameter and are found in swarms off the coast between
Cape Hatteras, North Carolina, and Cape Canaveral, Florida. They are practically confined
to pure open water and do not frequent the harbors. The mature meduse bear many ball-

ii

PLatTeE 66.

All figures are Cyanea capillata var. fulva.

. Planula ro days old; enlarged view. Figure 1’. Egg drawn to the

same relative size.

. Scyphostoma with 8 tentacles, drawn in the act of feeding. 1 mm.

high.

. Scyphostoma with 14 tentacles, 24 days old. 1.5 mm. high.
. Scyphostoma, oral view of a specimen with ro tentacles.
. Young medusa with bell 8 mm. wide; showing the exumbrella

papilla. There are 3 tentacles in each adradial cluster.

. Young medusa with bell 1o mm. wide; showing the lips expanded

and spread out oyer the bottom of the aquarium. There are 5
tentacles in each adradial cluster.

Fig. 7. Marginal sense-organ of the medusa shown in figure 6.

Drawn from life, by the author, at the Agassiz Laboratory, Newport,

Rhode Island, June 8 to July 9, 1895.

PLATE 66

ARoena te.

Se

-

PLATE 67.

Fig. 1. Cyanea capillata var. fulva, young medusa g mm. in diameter.
Showing its habit of resting with lips spread out over bottom of
aquarium and its tentacles elevated, while at the same time the
bell pulsates vigorously. Agassiz Laboratory, Newport, Rhode
Island, June, 1895.
Fig. 2. Cyanea capillata var. fulva. Oral view of quadrant of bell of a young
medusa 10 mm. in diameter. gf. internal gastric cirri. Agassiz
Laboratory, Newport, Rhode Island, June 10, 1895.
Fig. 3. Cyanea capillata var. fulva. Oral view of a medusa 15 mm. in
diameter showing gastrovascular pouches (light blue) of subum-
brella. Newport, Rhode Island, June, 1895.

. 4. Aurellia aurita, young medusa 7 mm. in diameter, showing develop-
ment of radial-canal system during formation of ring-canal.
Tortugas, Florida, June 4, 1907.

Fi

og

Drawn from life, by the author.

PLATE 67

MAYER

SEMAEOSTOMEA—CYANEA. 601

like clusters of developing planula gathered into the peripheral canals of the gastric space.
The gelatinous substance of the disk is translucent milky-blue in color, while the gastro-
vascular space, gonads, radial and circular muscles of the subumbrella and the entodermal
cores of the tentacles are purplish-pink. The outer parts of the veil-like folds of the palps are
amber-brown, while the parts adjacent to the mouth are pink. The concretions of the 8
sense-organs are reddish-brown. The planulz are yellow, but the ephyra is pink.

abe curtain-like oral fringes are relatively smaller than in Cyanea arctica. However,
the chief distinction of C. SEraie ATO is its peculiar pink coloration. Even in the young ephyra
only 2.5 mm. in diameter, the stomach-cavity displays a deep purplish-pink, very different
from the pale yellow-colored ephyra of the southern C. fulva.

Mature medusz of C. versicolor occur in the winter months along our southern coast.

Among thousands observed by the author during the winter of 1904-05 not more than a
dozen lacked the pink coloration and these resembledi the variety C. capillata var. fulua. They
were, however, swimming among swarms of the typical pink versicolor meduse. The variety
versicolor appears to be a well- Peneed local race of Cyanea capillata.

Cyanea capillata var. nozakii Kishinouye.
Cyanea nozaki1, Kisutnovuye, 1891, In Japanese, 3 pp-, 1 plate.

Kishinouye’s paper upon this medusa is in Japanese, but with a German abstract, and
accompanied by two clear figures of the animal. ‘The bell is flat and shield-shaped, 5 times
as wide as high, 160 to 260 mm. wide. The bell-radius 1s 3 times as wide as the radius of the
central proracli 16 rounded ephyra lappets, twice as wide as long. The ocular stomach-
pouches are nearly rectangular. The tentacular stomach-pouches are twice as wide at their
bases, and at the zone of the sense-organs 2.5 times as wide as the ocular pouches. Color,
milk-white. Found in the Inland Sea of Japan.

This medusa is distinguished from Cyanea capillata var. fulva only by its color.

Cyanea annaskala von Lendenfeld.

Cyanea annaskala, von LENDENFELD, 1882, Zeit. fiir wissen. Zool., Bd. 37, p- 465, taf. 27-33, 78 fign.; 1884, Proc. Linnean Soc.
New South Wales, vol. 9, p. 275, var. marginata, Ibid., p. 928, var. purpura, Ibid., p. 928; 1883, Annals and Mag. Nat.
Hist., ser. 5, vol. 12, p. 261 (nettling cells); 1887, Descript. Cat. Australian Mus. Sydney, Medus, part 1, pp. 20, 21.

Cyanea muellerianthe, Haacke, 1887, Jena. Zeit. fiir Naturwissen., Bd. 20, p. 605, taf. 36, fign. 1-4; 1888, Biol. Centralblatt, Bd.
8, P- 358.

Desmonema rosea, AGassiz, A., and Mayer, 1898, Bull. Mus. Comp. Zool. at Harvard College, vol. 32, p. 15, plate 1, fig. 1 (young
medusa in the Desmonema stage).

Cyanea mullerianthe=C. annaskala, VON LENDENFELD, 1888, Biol. Centralblatt, Bd. 8, p. 218.

Desmonema annasethe (? young medusa), Harcke1, 1880, Syst. der Medusen, Pp. 526, taf. 30, fign. 1-4.

Cyanea rosea (? young medusa), Quoy et Gaimarp, 1824, Voy ge de ’ Urante, Zool., p. 570, planche 85, figs. 1, 2.

() Cyanea purpura, KisHinouye, 1910, Journal College Sci. Tokyo, vol. 27, art. 9, p. 18, plate 4, figs. 18, 19.

Umbrella flat, shield-shaped, 70 to 200 mm. wide and about 12 to 25 mm. thick, with
a few protruding nettling-warts at the middle of the exumbrella; elsewhere smooth. 8 marginal
sense-organs which lack ocelli, and with 32 marginal lappets divided into 8 main flaps of 4
lappets each. These lappets are evenly rounded and not wider at the end than at their bases;
the 16 ocular lappets are about half as wide, as also are the 16 velar lappets adjacent to them.
There are 8 U-shaped clusters of long tentacles arising from the floor of the subumbrella,
with the concavity of each U directed Buaward. These ‘tentacles are very numerous and are
arranged in 3 to 4 crowded rows in each U; they are filiform and when extended are about
300 mm. long. The 4 complexly folded, curtain- -like lips are about as long as the bell-radius.
The 4 protrusive gonads are large and complexly folded. The 8 Gena radial pouches of
the central ome are only anette half as wide as the 8 velar pouches. All of the pouches
break up into blindly- -ending, branched, non-anastomosing vessels in the lappets. There 1s
no ring-canal. The gelatinous substance of the disk and the tentacles are colorless. The
entoderm of the gastral cavity is brown. Curtain-like lips intensely purple. Genital organs
of the male are rose-colored; those of the female are orange-yellow. The medusa is distin-
guished from the Cyaneas of the northern hemisphere mainly by its brilliant coloration. It
appears to be more closely related to C. versicolor than to any other form, and it is interesting
to observe that C. versicolor is the most southerly in its range of any of the northern Cyaneas.

602 MEDUS.E OF THE WORLD.

This species is abundant along the temperate coasts of Australia and is found in Port
Philip, Victoria, in large numbers from January to March. It is described in great detail by
von Lendenfeld, who finds that the embryos remain attached to the mouth-arms until “they
are nearly matured to young scyphostomz”; they then afhx themselves to bodies in the
water and produce a long stalk with a chitinous perisare and 8 arms (tentacles?). Accord-
ing to von Lendenfeld the ephyra develops into an adult medusa by a complicated meta-
morphosis. The lappets of the umbrella are said to be produced by fission, but this state-
ment probably applies only to the ocular lappets, not to the 8 primary ephyra lobes.

In the variety purpura from Melbourne Harbor, Australia, the mouth-curtains are rich
purple throughout, and in the variety marginata from Sydney their free-margins are purple,
but elsewhere they are colorless.

Fic. 383.—Drymonena “victoria,” after Haeckel, in Deep-sea Medusx of the Challenger Expedition.

Cyanea mullerianthe Haacke, from St. Vincent Gulf, South Australia, is a delicately
pink-colored variety of this medusa, and Desmonema rosea Agassiz and Mayer is the same
medusa when young and in the stage wherein the tentacles of each cluster arise in a single row.

Desmonema annasethe Haeckel, 1880, may be a young contracted specimen of this medusa.
The 16 so-called feathered, radiating ribs of the exumbrella present the appearance of being
due to unnatural contraction. The tentacles arise in 8 U-shaped groups with 13 to 17 ten-
tacles in each crescent. This form is described by Haeckel from a preserved specimen found
off the west coast of South Africa. Color (? )

Dendy, 1889 (Proc. Royal Soc. Victoria, p. 112), describes parasitic actinian larvae found
upon the mouth-curtains of the Cyanea of Port Phillip, Victoria.

SEMAEOSTOMEAS—DRYMONEMA. 603

Genus DRYMONEMA Haeckel, 1880.

Drymonema, HarcxEt, 1880, Sitzungsber. Jena. Gesell. fiir Med. und Naturw., Jahrg. 1880, Feb. 20; 1880, Syst. der Medusen,
p- 633; 1881, Report on Deep-sea Medusx, Challenger Expedition, Zool., vol. 4, p. 124.—Micter, F., 1883, Zool
Anzeiger, Jahrg. 6, p. 220.

The type species is Drymonema dalmatina Haeckel, 1880, of the Mediterranean. The
same species was renamed D. “victoria” by Haeckel in 1881. D. gorgo is a closely related
form from the coast of Brazil.

GENERIC CHARACTERS.

Cyaneide with 8 marginal sense-organs. The tentacles are not grouped in separated
clusters, but arise diffusely in a wide annulus from the subumbrella. The central stomach
gives rise to 16 radial pouches (8 ocular and 8 interocular) which branch dichotomously, but
do not anastomose. No ring-canal. Marginal lappets numerous. Development unknown.

Fic. 384.—Drymonema “victoria,” after Haeckel, in Deep-sea Meduse of the Challenger Expedition.

This genus is distinguished from Cyanea and Desmonema by the fact that its tentacles arise
not in 8 separate clusters, but from a wide annular zone in the subumbrella. Moreover, the 16
dichotomously branched radial-canals, numerous velar lappets, and the radial furrows of the
exumbrella are all distinctive of Drymonema.

Drymonema dalmatina Haeckel.
Drymonema dalmatina, HarcKE1, 1880, Syst. der Medusen, p. 642= D. victoria, 1881, Report Deep-sea Meduse Challenger Exped.,
Zool., vol. 4, p. 125, plates 30, 31.—GraerFe, 1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 342-
Drymonema cordelio, ANTIPA, 1892, Jena. Zeitsch. fiir Naturwissen, Bd. 27, p. 339, taf. 9, fign. 1-3.
Haeckel, 1880, had four small specimens of this medusa from the Dalmatian coast,
Mediterranean. They ranged from 120 to 160 mm. in width and had only 64 radial furrows

604 MEDUS® OF THE WORLD.

upon the exumbrella and g double lappets per octant, 144 in all. In Antipa’s specimens from
the Gulf of Smyrna, Mediterranean, the radial furrows had increased to be 144 and thus
corresponded in number with the lappets. Haeckel’s specimens had only 80 terminal gastric
canals, while Antipa’s specimens had 144.

The following is a description of Antipa’s specimens, these being the more mature:

Bell flatly rounded, shield-shaped, 500 to 1,000 mm. wide, 144 radial furrows on the
exumbrella, and between them 144 marginal lappets. 8 marginal sense-organs in deep niches.
4 perradial, veil-like oral palps, each ending in 2 points, and thus the palps have 8 adradial
points. These palps are more than 1.5 times as long as the disk-radius and their outer edges
are complexly folded. The numerous tentacles arise from the middle zone of the subumbrella,
halfway between the center and margin line, and are 3 to 6 times as long as diameter of the bell.
There are 4 interradial protruding horseshoe-shaped gonads. Mouth-opening wide. Wall
of mouth-tube thickened at 8 subradial places. The 16 stomach-pouches terminate in 144
dichotomous ramuli (128 tentacular and 16 ocular). Color reddish-white (pink ?). Gulf of
Smyrna, coast of Asia Minor, Mediterranean; Trieste, Adriatic Sea, Graeffe; Straits of
Gibralter (?) Haeckel.

Drymonema gorgo F. Miiller.

Drymonema gorgo, Miter, F., 1883, Zool. Anzeiger, Jahrg. 6, p. 220.

,

The disk is 20 to 500 (usually 300) mm. wide. “‘Mouth-arms” or palps longer than
diameter of umbrella, whereas they are only about half as long as this in D. dalmatina. The
8 ocular stomach-pouches fork once, thus giving 16 marginal diverticula as in D. dalmatina.
The 8 velar stomach-pouches branch dichotomously 4 times, as in the mature D. dalmatina,
but the fifth, sixth, eleventh, and twelfth branches in D. gorgo branch dichotomously a fifth
time, and thus each of the 8 ocular stomach-pouches gives rise to 20 terminal branches in
the lappets. There are thus 8 20+16=176 dichotomous terminal gastrovascular canals in
D. gorgo and only 8X 16+16=144 in D. dalmatina.

Found at St. Catharina Island, coast of Brazil. Rare.

This may prove to be a variety of D. dalmatina.

Genus (?) PATERA Lesson, 1843; DONACOSTOMA L. Agassiz, 1862.

Patera, Lesson, 1843, Hist. Zooph. Acaléphes, p. 322.—HAEcKEL, 1880, Syst. der Medusen, p. 533-

(2) Melusina, Harcket, 1880, loc. cit., p. §34; 1881, Report Deep-sea Meduse Challenger Expedition, Zool., vol. 4, p. 1 (abandoned
by Haeckel himself).

(?) Donacostoma, Agassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 118, 163.

The type species of this problematic genus is Patera cerebriformis, first described as
Dianea cerebriformis by Lesson, 1829, Voyage de la Coquille, Zooph., p. 124, planche ro.
The description and figure are evidently so inaccurate as to be all but worthless. Dzanea,
Lamarck, 1816 (Hist. Anim. sans Vert., tome 2, p. 504), is a synonym of Geryonza.

GENERIC CHARACTERS.

Cyaneidz which are said to have 16 rhopalia. 16 clusters of tentacles arise from the
subumbrella alternating in position with the sense-organs. Agassiz’s Donacostoma has only
8 rhopalia, but 16 rows of tentacles.

Patera cerebriformis is said to come from near the Cape Verde Islands, tropical Atlantic;
and another species Donacostoma wood: L. Agassiz, 1862 (Cont. Nat. Hist. U. S., vol. 4, pp.
118, 163), is from the China Sea, and is said to have 8 “eyes,’’ but with 16 branches of
tentacles like Patera, arranged in a single row in each Jobe. Neither is well enough known
to be worthy of description here, and indeed it is possible that both belong to the genus Dry-
monema or even to Cyanea itself.

Family ULMARIDZ Haeckel, 1880, sens. ampl.

Flosculide+ Ulmarida, Hatcken, 1880, Syst. der Medusen, pp. 535, 539-
Ulmaridz, VANuOFFEN, 1906, Nordisches Plankton, Nr. 11, p- 56.

FAMILY CHARACTERS.

Semzostomez with simple or branched radial-canals and a ring-canal. With hollow
tentacles. 4 interradial gonads. 4 mouth-arms with folded, curtain-like margins.

SEMAEOSTOMEAS—FLORESCA, 605

The medusz of this family are closely related to the Cyaneide, but differ in that their
radial-canals are placed in intercommunication by means of a marginal circular canal;
moreover, th radial-canals anastomose in some of the genera, and this is never the case in

the Cyaneide.

The genera of the Ulmaridz are as follows

1- SuBrAMILY UMBROSIN#A.

The tentacles arise singly from the margin in the clefts between the lappets. 4 evaginated, sac-like gonads without subgenital
pits. 8 or 16 marginal sense-organs. 4 unbranched mouth-arms.

(?) Floresca (immature ?) = Floscula+ Floresca Harcxet, 1880. 8 rhopalia, 24 tentacles, 32 lappets 16 simple unbranched
radial-canals.

Discomedusa, Craus, 1877= Ulmaris+ Umbrosa Harcket, 1880. 8 rhopalia, 24 tentacles, 32 lappets. The 8 perradial and
interradial canals are branched, the 8 adradial canals are simple. 4 gonads.

Parumbrosa, Kisuinouy®, 1910. Similar to Discomedusa but with 64 instead of 32 lappets. (See Appendix to this
volume.)

Undosa HarcKet, 1880. 8 rhopalia, 4o tentacles, 48 lappets. Radial-canals as in Discomedusa.

Diplulmaris, Maas, 1908= Ulmaropsis, VANHOFFEN, 1908. 16 rhopalia, 16 tentacles, 64 lappets, 32 radial-canals arise from
stomach. The 16 rhopalar canals branch and the 16 tentacular canals are simple. All canals are merged in a marginal
network of anastomosing vessels.

2. SUBFAMILY STHENONINAE.

Tentacles arise from floor of subumbrella. Evaginated, sac-like gonads without subgenital pits. 8 to 16 marginal sense-
organs. 4 unbranched mouth-arms.

Sthenonia, EscuscHotz, 1829. 8 marginal sense-organs. 8 adradial linear clusters of tentacles. Perradial and interradial
canals branched. Adradial canals some simple and some branched.

Phacellophora, Branpt, 1835. 16 marginal sense-organs alternating with 16 clusters of tentacles. Radial-canals in the
rhopalar radii are branched, in the tentacular radii simple. 4 gonads.

Poralia, VANHGFFEN, 1902. Numerous straight, unbranched radial-canals. Numerous gonads in a ring around the
stomach-margin. =

3- SuBramity AURELIN#&.

The tentacles and lappets arise from the sides of the disk above the margin. Invaginated gonads with external subgenital
pits. 8 marginal sense-organs. 4 simple or bifurcated mouth-arms. Numerous tentacles and lappets.

Aurellia, Peron anv Lesueur, 1809. 4 simple non-bifurcated mouth-arms. Some or all of the radial-canals give rise to
anastomosing branches. 4 gonads.

Aurosa, Harcket, 1880. Similar to Aure//ia but with 4 bifurcated mouth-arms.

Subfamily UMBROSIN&.
SUBFAMILY CHARACTERS.

The tentacles arise singly from the bell-margin in clefts between lappets. 4 protrusive,
bag-like gonads without subgenital pits. 4 unbranched mouth-arms.

(?) Genus FLORESCA Haeckel, 1880, sens. ampl.
Floscula + Floresca, HArcKeL, 1880, Syst. der Medusen, p. 537, 538, 643-

GENERIC CHARACTERS.

Haeckel’s genera Floscula and Floresca may possibly be the young stages of some medusz
of the Ulmaridz. In these forms we find that the central stomach gives rise to simple radial-
canals which connect with one another by means of a marginal ring-canal. The tentacles
arise from the clefts between the lappets, not from the floor of the subumbrella. We must
bear in mind, however, that in the young of Cyanea the tentacles first appear in the clefts
between the lappets and later the margin grows outward, leaving them to project from the
subumbrella. Indeed all tentacles in Sey phomedusz are structures of the subumbrella.

Floscula has 8 tentacles and 16 marginal lappets; Floresca has 24 tentacles and 32
marginal lappets. Indeed Floresca presents every appearance of being immature and only
an advanced stage of “‘ Floscula,” both being one and the same species. Both come from
the tropical Indian and Pacific Oceans.

A brief description of these medus# may be of service should they prove to be mature
forms. Haeckelis the only naturalist who has seen them.

“Floresca parthenia” Haeckel.
Floresca parthenia+F. pallada, Harcket, 1880, Syst. der Medusen, pp. 538, 539, taf. 32, fign. 5-8.
Floscula promethea (younger stage), Harcket, loc. cit., p. 537, taf. 32, fign. 1-4. F. pandora, Ibid., p. 643.
Bell rounded, 50 mm. wide, 30 high, with a 16-rayed pigmented star-like marking in
the perradii, interradii, and adradii of the exumbrella. 8 rhopalia, (8X 4) 32 tongue-shaped

606 MEDUS® OF THE WORLD.

lappets. 24 (83) hollow tentacles 2 to 3 times as long as the bell-diameter. Throat-tube
I.5 times as long as the bell-radius and as the 4 complexly folded, leaf-shaped lips. The
central stomach gives rise to 16 unbranched radial-canals which are joined one to another by
a marginal ring-canal. 4 interradial crescentic gonads with their convexities outward.
These project from the floor of the subumbrella. There are no subgenital pits. The gonads
are lined on their inner, concave sides by a row of gastric cirri. Color (?) Found at New
Caledonia, New Guinea, and the Cocos Islands in the tropical Pacific and Indian Oceans.
Another specimen called “Floscula pandora” by Haeckel, 1880 (p. 643), is from the tropical
Pacific. The bell is 30 mm. wide, without a star-like marking upon the exumbrella, and with
a short throat-tube hardly one-fourth as long as the long, narrow mouth-arms. Tentacles as
long as the bell-radius. Lappets oval, sharp pointed, 1.5 times as long as broad.
.«» These meduse appear to be immature, although Haeckel states that the gonads contained
ripe eggs, but he studied only preserved specimens and cut no sections.

Fic. 385.—Floscula promethea.” Fic. 386.—“Floresca parthenia.”

After Haeckel, in Das Syst. der Medusen.

Genus DISCOMEDUSA Claus, 1877.

Discomedusa, Craus, 1877, Denkschrift, Wien. Acad., Bd. 38, p. 42.
Ulmaris+ Umbrosa, Harcket, 1880, Syst. der Medusen, p. 545.
Umbrosa, Maas, 1908, Expédition Antarctique Frangaise, Meduses, p. 9.

The type species is Discomedusa lobata Claus, 1877, of the Mediterranean. This may
prove to be an arrested variety of Haeckel’s Undosa undulata of the west coast of tropical
Africa. In Undosa, however, there are 40 tentacles and 48 lappets, whereas in Discomedusa
there are 24 tentacles and 32 lappets.

GENERIC CHARACTERS.

Ulmaride with 24 (38) tentacles, 32 (48) lappets, and 8 sense-organs. The ten-
tacles arise from the clefts between the marginal lappets. There are 8 simple, unbranched
adradial canals, 8 branched, perradial and interradial canals, and a marginal ring-canal.

Haeckel’s genus Ulmaris is apparently only the young of Discomedusa in a stage wherein
there are only 8 tentacles and 16 lappets. His genus Undosa bears the same relation to Disco-
medusa that Dactylometra does to Chrysaora, the tentacles and lappets having increased from

SEMA EOSTOME.2—DISCOMEDUSA, 607

24 and 32 to 40 and 48, respectively. Thus Discomedusa is a growth-stage in the development
of Undosa. The medusa of Trieste, Adriatic, appears to become mature, however, with
only 24 tentacles, and it is interesting to observe that under unfavorable conditions in brackish
water Dactylometra also becomes mature with 24 instead of 40 tentacles.

Discomedusa lobata Claus.

(?) Medusa stelligera EHRENBERG, 1835, Abhandl. Acad. Berlin, p. 260.—1836, Akalephen des Rothen Meeres, p. 82.
Discomedusa lobata, Ciaus, 1877, Denkschr. Wien. Acad., Bd. 38, p. 42, taf. 8, 9 —Grarrrr, 1884, Arbeit. Zool. Inst. Wien.,

Bd. 5, p- 343-
Umbrosa lobata, Harcke1, 1880, Syst. der Medusen, p. 546.
Ulmaris prototypus (? young medusa), Harcker, 1880, Joc. cit., p. 545, taf. 33, fign. 1-4.

Bell shield-shaped, flatter than a hemisphere, 150 mm. wide, 40 mm. high. The 16
ocular lappets are as wide as, but somewhat longer than, the 16 tentacular lappets. “There
are 24 tentacles, and 8 adradial ones are about as long as bell-diameter and twice as long
and thick as the 16 secondary tentacles. The 4 mouth-arms are wide, tapering, somewhat
longer than bell-radius and with curtain-like, folded margins provided with numerous ten-
tacula. Mouth cruciform. Genital radius somewhat wider than half the bell-radius. The 4
crescentic gonads are convex outwardly
and their ends nearly touch in the 4 per-
radii. Centripetal to these gonads are 4
lines of long, numerous, gastric cirri, one
row for each gonad.

The central stomach gives rise to 8
unbranched, adradial canals and to 8 per-
radial and interradial vessels, each of
which gives rise to a pair of branched
and anastomosing side-branches, the ter-
minal ramifications of which fuse with the
adradial canals and with the marginal
ring-canal.

This medusa is found at Trieste, Adri-
atic Sea, from December to March, the
young being common in January and the
adult in February and March. Claus gives
the best description of it. It may be 1den-
tical with the imperfectly described Medusa
stelligera Ehrenberg, from the harbor of
Alexandria, Egypt, in October. Ehren-
berg’s medusa, however, appears to have
about 40 short tentacles, all of equal
length, and may therefore belong to Haeckel’s genus Undosa.

The young medusa of Discomedusa lobata passes through a stage wherein there are only
16 lappets and 8 adradial tentacles. The 8 adradial canals are simple and the 8 perradial
and interradial ones branched.

Fic. 387.—“Ulmaris prototypus,” after Haeckel, in
Das Syst. der Medusen.

Discomedusa philippina, sp. nov.

This form is allied to D. lobata of the Mediterranean, but it appears to be smaller and to
differ in the bluntness of its lappets, in having no fusions between the adradial and perradial
and interradial canals, and above all by the blind branches on the outer side of the ring-
canal.

Bell 29 mm. wide, flatter than a hemisphere, evenly rounded, exumbrella thickly be-
sprinkled with prominent wart-like projections. Gelatinous substance fairly thick at center,
thin at bell-margin. 8 rhopalia. 32 oval lappets all similar each to each, thus differing from
D. lobata. 24 tentacles, the 8 adradial being longer and stouter than the 16 intermediate ten-
tacles. The tentacles were all broken off in the specimens obtained by the Albatross so that
their lengths could not be determined. 4 simple, crenulated lips at the end of a 4-cornered

60S MEDUS.2 OF THE WORLD.

manubrium about as long as the bell-radius. 4 interradial, crescentic gonads with their con-
vexities outward and with a single row of simple unbranched gastric cirri along their inner
sides. These gonads are about 5 times as wide as the perradial interspaces between them.
Central stomach 15 mm. wide. 8 simple, unbranched, adradial canals arise from central
stomach and proceed straight outward to ring-canal. Trident-like, pitch-fork-shaped, anas-
tomosing canals arise from the 8 (perradial and interradial) sides of the stomach and break
up into a network of vessels which fuse with the ring-canal. ‘These networks, however, do
not fuse with the 8 adradial canals, in this differing from D. lobata. On its outer side the
ring-canal gives rise to 64 simple, unbranched, blindly-ending diverticula; a pair in each
lappet. In formalin the gelatinous substance is transparent and the entodermal canal-system
dull milky. The gonads appear to be mature, but the preservation is such that I can not be

Fic, 388.—Discomedusa philippina, sp. nov. Drawn by the author, from speci-
mens obtained by the U. S. Fisheries Bureau steamer d/batross in
Catingan Bay, Philippine Islands, April 20, 1908.
certain that this is the case. Six specimens were found by the U. S. Fisheries Bureau steamer
Albatross, in Catingan Bay, Philippine Islands, on April 20, 1908. This medusa may possibly
be the young of Parumbrosa polylobata (see Appendix) but the large size of its gonads and
complexly branched canals renders this improbable.

Genus UNDOSA Haeckel, 1880.

Undosa, Harcker, 1880, Syst. der Medusen, p. 546.—Maas, 1908, Expédition Antarctique Frangaise, Meduses, p. 9.
Ulmaris (young medusa), loc. cit., p. 545.

The type species is Undosa undulata Haeckel, of the tropical Atlantic coast of Africa.
It is possible that Ehrenberg’s Medusa stelligera of the Mediterranean may be identical with
this species, but it is so imperfectly described that it is unrecognizable.

GENERIC CHARACTERS.

Ulmaridz with 8 sense-organs, 48 (8 x 6) marginal lappets, and 40 (58) tentacles which
arise from the clefts between the lappets. 4 interradial protrusive gonads without subgenital
ostia. 8 simple adradial canals, 8 branched perradial and interradial canals, and a ring-canal.

The genus is derived from Discomedusa through the multiplication of lappets and ten-
tacles. During the growth of the medusa they pass through a “Discomedusa” stage.

SEMAEOSTOME J

UNDOSA, DIPLULMARIS.

Undosa undulata Haeckel.

Undosa undulata, Harcxet, 1880, Syst. der Medusen, p. 546, taf. 33, fign. 5, 6.

Bell 120 mm. wide, 40 mm. high, flatly rounded. A brown, 16-rayed, star-like figure on
exumbrella composed of pigmented nettling ridges. 48 (68) sharp-pointed, oval lappets,
somewhat longer than wide and projecting prominently; the 16 ocular lappets are somewhat
longer than the others. Of the 40 tentacles, the 8 adradial are 2 to 3 times as long as bell-
radius, twice as long as the 16 secondary, and 3 times as long as the 16 tertiary tentacles.
All tentacles are hollow. The 4 mouth-arms resemble those of Aurelia aurita, but their
margins are more curtain-like and more folded and provided with numerous tentacular
filaments. The 4 gutters of the mouth-arms are very deep. The 4 folded, interradial, bag-
like gonads project from the floor of the subumbrella. There are no subgenital ostia. The
genital radius is about half that of the bell itself. Margin of central stomach circular. 8
simple, unbranched, adradial canals and 8 (perradial and interradial) canals, which give
rise to numerous side branches which in turn anastomose, forming a network of vessels con-
necting all 16 canals one with another and with the marginal ring-canal.

The general color is bluish, and
the star-like marking on the exum-
brella dark-brown. Found at Fer-
nando Po, coast of Guinea, Africa.

Haeckel’s “ Ulmaris prototypus se
from St. Helena is probably the
young of this species (Joc. cit., 1880,
p- 545, taf. 33, fign. 1-4).

Genus DIPLULMARIS MAAS, 1908.

Diplulmarts, Maas, 1908, Expédition Antarctique
Frangaise, Meduses, p. 9.
Ulmaropsis, VANHOFFEN, 1908, Deutsche Siidpolar
Expedition, Bd. 10, Zool. 2, p. 45.
The type species is Diplulmaris
antarctica, Maas, 1908, from the
Antarctic Ocean.

GENERIC CHARACTERS.

Ulmaride with 16 tentacles, 64

- marginal lappets, and 16 marginal

sense-organs. 16 branched radial-

canals in the radii of the sense-organs

and 16 simple straight radial-canals

in the tentacular radii. All 32 of the canals are joined by an anastomosing network of vessels
near the bell-margin.

Fic. 389.—Undosa undulata, after Haeckel is Das Syst. der Medusen.

This medusa bears the same relation to Undosa that Sanderia does to Dactylometra.
Indeed, there is a remarkable case of parallelism of mutation in the Pelagide and Ulmaride
as is shown in the accompanying table.

Family Pelagide.

Family Ulmaride.

|

]

8 rhopalia, 8 tentacles, 16 lappets...........-.-- Pelapial. cpepiatc sisi Ulmaris (immature ?)
8 rhopalia, (8 X 3) tentacles, 32 lappets.......-.- Chrysaoras -<c\e- cies | Discomedusa

8 rhopalia, (8 X 5) tentacles, 48 lappets.......... Dactylometra | Undosa .

16 rhopalia, (8 Xz) tentacles, 64 lappets. .....-.-. Sanderiaey fe satsle ace == | Diplulmaris

Diplulmaris is a connecting link between the subfamilies Umbrosidi1, in which the ten-
tacles arise from clefts between the lappets, and Sthenonidi1, in which they arise from the
subumbrella floor; for in the young Diplulmaris the tentacles arise from the subumbrella,

610 MEDUS-E OF THE WORLD.

but later the bell-margin becomes cleft so that the tentacles project from the clefts. Dzplul-
maris resembles the genus Phacellophora in having 16 rhopalia.

Vanhoffen, 1908, describes this medusa under the name Ulmaropsts drygalski1. Some
of his specimens were more advanced than those seen by Maas, and they had 64 marginal
lappets instead of only 32, as in those of Maas.

Diplulmaris antarctica Maas.

Diplulmaris antarctica, Maas, 1908, Expéd. Antarctique Frangaise, Meduses, p. 12, 1 taf.
Ulmaropsis drygalskii, VANHGFFEN, 1908, Deutsche Siidpolar Expédition, 1901-03, Bd. 10, Zool. 2, p. 45, fign. 10-12.
Ulmaropsis antartica, VANHOFFEN, 1909, Ibid., Bd. 10, Zool. 2, p. vi.

Fic. 390.—Diplulmaris antarctica, young medusa, after Maas, in Meduses Expédition Antarctique Frangaise.

The largest specimen is described by Vanhéffen, whose publication appeared only a few
weeks after that of Maas, and is as follows:

Bell 42 mm. wide, 16 marginal sense-organs alternating with 16 simple, hollow tentacles.
The sense-organs and tentacles are flanked by 64 slender, pointed lappets, the 32 lappets
flanking the sense-organs being wider and longer than those flanking the tentacles. 32 radial-
canals arise from the central stomach these being in the tentacular and ocular radii. The
tentacular canals are simple and unbranched, but each of the ocular canals gives off 2 pairs

SEMAEOSTOME.A—DIPLULMARIS, STHENONIA, 611

of side branches and thus 96 canals radiate outward toward the margin, before reaching
which they are all connected by a network of anastomosing vessels. There are 4 lips and 4.
gonads. g ephyre and young medusz of this species were found by the German Antarctic
Expedition between January and March. They ranged from 4 to 22 mm. in diameter. The
species appears to be quite variable, for only 6 of Thon were 16-rayed, while the 3 others were
12, 15, and 17 rayed respectively.

Two immature specimens were studied by Maas. ‘The bell of larger was 35 to 40 mm.
wide. 16 marginal sense-organs flanked by only 32 bluntly-pointed “lappets. 16 hollow,
tapering tentacles, Eomewhat shorter than the bell-radius. These tentacles arise from the
inter-rhopalar clefts between the lappets. The rhopalar clefts are only half as deep as the
inter-rhopalar. Ventral stomach circular, nearly as wide as bell- radius. 16 trident-shaped
radial-canals arise from the stomach-margin in the rhopalar radii and alternate with 16
straight, narrower, unforked canals in the tentacular radii. All the radial-canals give off
anastomosing side branches in the outer parts of their lengths near the ring-canal. The ring-
canal is at the zone of the origins of the tentacles, and 16 ‘slightly branched radiating diverti-
cula extend outward from it to the sense-organs. The 4 lips are bordered by curtain-like
fringes. 4 interradial gonads with clusters of gastric cirri. Canal system yellowish- brown,
the tentacles dull purple. The specimen was immature.

Maas also describes a smaller specimen only 15 mm. in diameter (fig. 390). This had
8 long tentacles alternating with 8 short. The short tentacles ile from the floor of the
subumbrella at some distance inward from the clefts between the lappets, and it appears that
during growth the clefts extend inward until they meet the tentacles at the zone of the ring-
canal. There are 16 simple, unbranched radial-canals in the radii of the tentacles and 16
trident-like canals in the rhopalar radii. None of these canals gives off side-branches close
to the ring-canal, as in later life. 4 interradial oval gonads are now visible, and the short
throat-tube expands into 4 pointed lips with folded margins. Central stomach circular,
about as wide as bell-radius.

This medusa is found in the Antarctic Ocean from January to March where it lives
along the edge of the Antarctic continent having been taken at Cape Adare, off Kaiser Wilhelm
II Land, and at other places.

Subfamily STHENONIN A.
SUBFAMILY CHARACTERS.

The tentacles arise in linear clusters from the floor of the subumbrella. 8 to 16 marginal
sense-organs. 4 protrusive, bag-like gonads without subgenital pits. 4 unbranched mouth-
arms. Some single and some branched radial-canals.

Genus STHENONIA Eschscholtz, 1829.

Sthenonta, Escuscuoxtz, 1829, Syst. der Acalephen, p. 59—Harcxet, 1880, Syst. der Medusen, p. 548.—VANHOFFEN, 1906,
Nordisches Plankton, Nr. 11, p. 56.

The type species and only known form is Sthenonia albida of Awatscha Bay, coast of
Kamtschatka.

GENERIC CHARACTERS.

Ulmaridz with 8 rhopalia, 16 ocular lappets, 8 bifurcated velar lappets, and 8 adradial
clusters of tentacles which arise from the subumbrella. 8 branched radial-canals in the rho-
palar radii. Numerous simple or branched radial-canals in the radii of the velar lappets and a
ring-canal. No subgenital pits.

Sthenonia albida Eschscholtz.

Sthenonia albida, Escuscuortz, 1829, Syst. der Acalephen, p.59, taf. 4.—pe BrarnviLte, 1834, Man.d’Actinologie, p.291, planche
36, fig. 1—Harcxer, 1880, Syst. der Medusen, p. 548.—VaNuGrreN, E., 1906, Nordisches Plankton, Nr. 11, p. 56, fig. 2

This form has not been seen since Eschscholtz described it.

Bell about 300 mm. wide, flat, and shield-shaped. 8 marginal sense-organs. 16 ocular
and 8 velar lappets with evenly rounded, reentrant markings. Each ocular lappet has a small
pointed projection into which the gastrovascular system extends, and each velar lappet has

612 MEDUS OF THE WORLD.

a pair of these projections. 8 adradial rows of tentacles arise from the subumbrella in the
intervals between the sense-organs; these rows being somewhat shorter than the intervals
berween them. The 4 mouth-arms are only one-third as long as bell-radius. The central
stomach is less than one-fifth as wide as the bell, and there are 4 interradial clusters of
gastric cirri. 8 radial-canals, each of which gives rise to several side branches, arise from
the central stomach in the radii of the sense-organs. A simple and a forked canal arise in
each of the 8 radii of the yelar lappets. These radial-canals anastomose to some extent and
fuse with a wide ring-canal in the zone of the sense-organs. On its outer side the ring-canal

iy “
Ay
Ata)
HAN

iy

WN

i i

if
yy

Fic. 391.—Sthenonia albida, according to Eschscholtz, after Vanhéffen
in Nordisches Plankton.

gives off a trident-shaped vessel in each rhopalar-radius and a pair of vessels in each velar
lappet-radius. Bell whitish, gonads and canals milk-white.
Found by Eschscholtz in Awatscha Bay, coast of Kamtschatka, Siberia.

Genus PHACELLOPHORA Brandt, 1835.

Phacellophora, Branvt, 1835, Mém. Acad. Impériale des Sci. St. Pétersbourg, sér 4, tome 2, p. 224; 1838, Mém. Acad. Impériale
des Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 365.—Harcket, 1880, Syst. der Medusen, p. 549.—VANHOFFEN, 1906,
Nordisches Plankton, Nr. 11, p. 58.—Brownr, 1908, Trans. Royal Soc. Edinburgh, vol. 46, p. 247.

Heccadecomma, Branpt, Ibid., p. 380.

Callinema, Verrii1, 1869, American Journ. Sci., ser. 2, vol. 48, p. 117; Ann. and Mag. Nat. Hist., vol. 4, p. 161.—Ferwkes, 1888,
Bull. Mus. Comp. Zool. at Harvard College, vol. 13, p. 235.

Phacellophora= Heccadecomma, Maas, 1906, Fauna Arctica, Bd. 4, Lfg. 3, pp. 506, 512, Jena.

The type species is P. camtschatica, described by Brandt, 1838, from the North Pacific.

SEMA EOSTOME 2—PHACELLOPHORA, 613

GENERIC CHARACTERS.

Ulmaride with 16 marginal sense-organs and numerous marginal lappets. The ten-
tacles arise in 16 simple, linear clusters from the floor of the subumbrella, centripetal to the
margin. Central mouth surrounded by 4 mouth-arms bearing curtain-like lips. Gonads
are 4 complexly folded sacs which project outward in the 4 interradii from the floor of the
subumbrella. No subgenital pits. The central stomach gives rise to numerous radiating
canals, some of which anastomose. There is a ring-canal at the bases of the marginal lappets.
Tentacles hollow.

Synopsts of the Species of Phacellophora.

| P. ornata.t

Name. P. camtschatica. P. sicula.* | P. ambigua.+
} _ — | =
Diameter of disk in mm.| 500 to 600 155 150 to 200 | 350
Shape and number of | 16 trident-shaped lap- | 32 narrow, rounded | (4X16) 64 lappets, all | (416) to (6 X16) lap-
marginal lappets. pets in rhopalar radii. | rhopalar lappets. 16 similar each to each, | pets all similar each to
7 small lappets in each| simple velar lobes. and evenly rounded. each and evenly
of 16 semicircular, rounded.
velar lobes.

Shape of mouth-arms. | Long, narrow, resemb- | As in P. ambigua. | Wide, curtain-like, and | As in P. ambigua.

ling those of Aurelia. resembling those of
Cyanea.

Number of radial-canals.. 16 branched rhopalar | As in P. ambigua. | 16 branched, rhopalar | 16 branched rhopalar
canals, and 5 X16 sim- | and 3to5 X16 simple,| and (2 X16) to (5 X16)
ple unbranched radial- unbranched canals. simple canals.
canals. |

Number of tentacles in | 20 to 24 9 to 15 | 9 5 tog

each cluster. |

Where found. North Pacific, Siberia to

Mediterranean, Naples, | Pacific coast of North
California.

Messina, and off the America, Washington. |
coast of Japan. |

North Atlantic, northern
coast of Maine.
Montevideo, South
America.

*Intermediate in character between P. camtschatica and P. ornata.
Closely allied, probably identical.

Phacellophora camtschatica Brandt.

Phacellophora camtschatica, BRanvt, 1838, Mém. Acad. Sci. St. Pétersbourg, tome 4, Sci. Nat., sér. 6, p- 366, taf. 8—Acassiz, A.,

1865, North Amer. Acal., p. 44.—Harcket, 1880, Syst. der Medusen, p. 549 —VANHOFFEN, 1906, Nordisches Plankton,

Nr. 11, p. 58, fig. 23.

Bell flat, 500 to 600 mm. wide. 16 marginal sense-organs each beneath a large, projecting,
covering scale, which is flanked by 2 narrow, rhopalar lappets, each about half as wide as the
covering-scale itself. 16 wide, nearly semicircular, velar lappets, each cleft into 7 marginal
lappets. 16 crescent-shaped linear clusters of tentacles arise from subumbrella floor of velar
lappets; the convexities of these crescents are inward and their horns outward, and each
consists of 20 to 24 tentacles. Tentacles about as long as bell-radius. The 4 mouth-arms are
wide, tapering, and thick, and about 1.5 times as long as bell-radius; their lips are folded
in a curtain-like manner. There are 4 interradial, sac-like gonads with narrow, perradial inter-
spaces between them. The central stomach gives rise to 16 rhopalar canals which send out
lateral branches, and also to 80 (5 16) simple unbranched velar canals. There is a ring-canal
at the zone of the tentacles, on the outer margin of which there arise 7 blindly ending, simple
diverticula in each velar lappet and a trident-shaped branch in the radius of each sense-organ.

Bell colorless to bluish, gonads reddish-brown, canals yellow, tentacles light-violet.

This medusa ranges along the shores of the North Pacific from Kamtschatka to San
Francisco, California. It has not been figured since Mertens studied it.

Phacellophora sicula Haeckel.

Phacellophora camtschatica, Hertwic, O. unp R., 1878, Nervensyst. und Sinnesorgane Medusen, pp. 113, 114, taf. 9, fig. 15;
taf. 10, fig. 16.—Haercket, 1880, Syst. der Medusen, p. 551-
Phacellophora ambigua, Kisu1Novuye, 1910, Journal College of Sci. Tokyo, vol. 27, art. 9, p. 21, 1 fig.

The brothers Hertwig describe the marginal sense-organs of this exceedingly rare
medusa and give a very diagrammatic figure of a part of the bell-margin. They were appar-

614 MEDUS.E OF THE WORLD.

ently under the impression that it was identical with P. ornata Brandt, from the Pacific, but
Haeckel rightly distinguished it as a distinct species.

The following description is based upon my study of a single good specimen of this medusa
collected by Dr. S. Lobianco at Naples, Italy, on January 11, 1901, and now preserved in
formalin at the Naples Zoological Station.

Disk 155 mm. in diameter, flatter than a hemisphere, being only 55 mm. high. Exum-
brella surface finely granular, being covered with small, thickly clustered nematocyst-warts.
16 marginal sense-organs, 4 perradial, 4 interradial, and 8 adradial. Sense-organs set at
bottom of deep, narrow clefts in bell-margin. The sense-club has no ocellus, merely a terminal
mass of entodermal concretions. No sensory pit in exumbrella above the sense-club. 32
narrow, rhopalar lappets are separated by shallow clefts from the 16 wide, velar, simple lappets.

Fic. 392.—Phacellophora sicula, drawn by the author, from a specimen found at Naples by
Dr. S. Lobianco, January 11, 1901. B, enlarged view of part of one of the tentacles.

The tentacles are arranged in 16 clusters and arise in a single row from the inwardly-
arched outer margin of the ring-canal, on the subumbrella side of the 16 velar lappets. Each
cluster consists of about g to 15 tentacles. In the Naples specimen these tentacles are about
half (75 mm.) as long as diameter of disk and are set inward at a maximum distance of 17.5
mm. from the bell-margin. A narrow canal extends throughout the length of each tentacle
on the inner (centripetal) side to its tip. A double row of mammiform, nematocyst-bearing
papillae extends along the inner side of each tentacle close to the tentacular canal, which sends
off lateral diverticula into the papilla. The outer (centrifugal) side of each tentacle is pro-
vided with circular muscle-fibers, which are interrupted along the line of the papillz.

SEMA EOSTOME.A2—PHACELLOPHORA. 615

The genital cross is about one-third (55 mm.) as wide as the bell itself. The 4 extruded,
pouch-like gonads with their swollen, folded, wart-like genital-sacs resemble those of Cyanea.
The 4 gonads are separated by very narrow intervals in the 4 perradii. The 4 wide, curtain-
like lips also resemble those of Cyanea and are not quite as long as the radius of the bell.
The central stomach gives rise to 16 rhopalar radial vessels which fork outwardly; and also
to 48 (316) inter-rhopalar radial-canals which are simple and do not fork. Lateral anasto-
moses between these radial-canals are very rare. Ring-canal very well developed, about 3
mm. wide, while the radial-canals are each about 2.5 mm. wide, being about as wide as the
spaces between them. The ring-canal gives rise to from 5 to 8 straight, simple, blindly-ending,
centrifugal vessels in each velar lappet.

(

\ Wie Qa
Oem) )
WOMAN /Y

WAYn 5 iy)

SS iy 1)

Fic. 393.—Phacellophora ambigua, according to Brandt, after Vanhoffen, in Nordisches Plankton.

In formalin the exumbrella is yellowish-milky in color, the canal-system milky and
translucent, the gonads dull orange to ocher, and the lips of a lighter hue of the same color.
Found at Naples and Messina, Mediterranean, and off the coast of Japan (Kishinouye).
This species is closely related to P. ambigua, but is distinguished by having only 16 wide,
simple, velar lappets instead of 32 narrow ones, as in P. ambigua. P. sicula is probably
only an arrested variety of P. ambigua in which the velar lobes remain entire and uncleft.

Phacellophora ambigua Haeckel.

Haccadecomma ambiguum, Branpvr, 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 380, taf. 27, 28.—AGassiz,
A., 1865, North Amer. Acal., p. 43-
Phacellophora ambigua, HAEcKEL, 1880, Syst. der Medusen, p. 550.-—VANHOFFEN, 1906, Nordisches Plankton, Nr. 11, p. 58, fig. 24.

Bell flatter than a hemisphere, 150 to 200 mm. wide. There are 64 (416) evenly
rounded, marginal lappets all similar each to each. The sense-organs are sunken within deep,
narrow clefts. Mouth-arms broad, curtain-like, similiar to those of Cyanea instead of being
Aurellia-like, as in P. camtschatica. About g tentacles in each of the 16 linear clusters. The
16 rhopalar canals are forked and between them are 48 (316) simple, unbranched, radial

616 MEDUSA OF THE WORLD.

vessels. The ring-canal gives rise centripetally to a short diverticulum in the radius of each
sense-organ and to 8X16 inter-rhopalar diverticula. The gonads are 4 interradial, sac-like,
protruding pouches separated by narrow intervals in the perradii.

This species is closely related to P. stcula of the Mediterranean and Pacific, but is dis-
tinguished by having 32 instead of 16 velar lappets. It is found along the Pacific coast of
North America, Port Townsend and Straits of Fuca, Washington.

Phacellophora ornata Haeckel.

Callinema ornata, V erritt, 1869, American Journ. Sci., ser. 2, vol. 48, p.117; Ann. and Mag. Nat. Hist., vol. 4, p. 161.—Fewkes,
1888, Bull. Mus. Comp. Zool. at Harvard College, vol. 13, No. 7, p. 235, plate 6, 4 figs.; 1888, Report U.S. Expedition to
Lady Franklin Bay, vol. 2, p. 40.

Phacellophora ornata, Hackett, 1880, Syst. der Medusen, p. 643.—Harait?, 1904, Bull. U.S. Bureau of Fisheries, vol. 24, p. 68.—
Vanuorren, 1906, Nordisches Plankton, Nr. 11, p. 59, fign. 25-26.—Browne, 1908, Trans. Royal Soc. Edinburgh, vol. 46,
Pp- 247, plate 2, figs. 3, 4.

Disk quite flat, with a slight dome-shaped aboral apex; it is about 350 mm. in diameter,
and the marginal lappets droop vertically. The surface of the exumbrella is covered with
small nematocyst-warts. Gelati-
nous substance of disk quite thick
and rigid. 16 marginal sense-
organs are set within niches be-
tween the 32 ocular lappets.
Ocular lappets about twice as long
as velar ones, but not so numerous,
there being 2 to 4 velar between
each successive pair of ocular lap-
pets. The clefts separating the
ocular lappets are deeper and
more distinct than those separat-
ing the velar lappets. Over 100
long tentacles arise from the floor
of the subumbrella in a broken
circle in 16 inter-rhopalar clusters
at a short distance inward from
the bases of the marginal lappets.
The 5 tog tentacles between each
successive pair of sense-organs
vary considerably in size, the long-
est being about equal to bell-
diameter; they are hollow and flat,
and there is a wavy double thick-
ening along the centripetal narrow
edge, which is covered with nema-
tocysts. Mouth simple and 4-
cornered, situated at center of
subumbrella. The 4 mouth-arms
are each about as long as bell-
diameter, their free edges much
folded. They are highly flexible
and contractile. The gonads are found in 4 interradial, crumpled sacs which project out-
ward from the floor of the subumbrella at the sides of the mouth. Central stomach 4-lobed,
being extended outward in the radii of the 4 genital organs, very much as is the case in
Aurellia. A large number of radiating canals run outward from the periphery of the central
stomach to the circular canal, which lies at a considerable distance inward from bell-margin.
The radiating canals in the radii of the sense-organs branch and anastomose, while those in
the tentacular radii are simple and slender. There are about 2 to 5 of these simple canals
between each successive pair of anastomosing canals. Circular canal broad, somewhat sinuous,
and it lies under the insertions of the ring of tentacles. Outwardly it gives rise to a blind canal

Fic. 394.—Phacellophora ornata.

SEMAEOSTOMEA]—PHACELLOPHORA, PORALIA. 617

in each velar lappet and a trident-shaped canal to each sense-organ and its adjacent lappets.
Usually the outer ends of these blind canals are simple, but occasionally they bifurcate
(fig. 395).

Disk transparent, the radiating and circular canals slightly brown in color. Sense-organs
glistening white. Nematocyst-bearing edges of tentacles white. Central stomach orange-
yellow, the mouth-arms citron-yellow, the gonads yellowish-brown.

This species is found at Eastport, Maine, and in the Bay of Fundy. It is very rare, and
has been taken there only by Verrill and Fewkes. In 1908 Browne describes a closely allied
or identical species from the South Atlantic about 200 miles east of Montevideo, South America.

I am gratefully indebted to Professor Verrill for permitting me to make drawings (figs.
394, 395) of the type specimen preserved in the Peabody Museum at Yale University.

This species is closely related to P. ambigua, but is distinguished by its greater number of
velar lappets and radial-canals. The velar lappets are also shorter than in P. ambigua.

Genus PORALIA Vanhoffen, 1902.

Poralia, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 4o.—Bicetow, H.B.,
1909, Mem. Museum Comp. Zool. at Harvard College, vol. 37, p. 45, plate 13, figs. 1-5.

GENERIC CHARACTERS.

Ulmaridz closely related to Phacellophora. With numerous simple radial-canals, and a
ring-canal, which on its outer side gives rise to blindly-ending vessels. The gonads form a ring

Fic. 395.—Phacellophora ornata. Portion of bell-rim.

of outpocketings in the lateral wall of the stomach extending entirely around the base of the
stomach, but interrupted at frequent intervals by vertical thickenings of the stomach wall.
Tentacles (?) There are no subgenital pits in the floor of the subumbrella. The mouth parts
are imperfectly preserved but appear to resemble those of Cyanea or Phacellophora. —

This genus was founded by Vanhoffen, but his single specimen was imperfect and imma-
ture, and our knowledge of it is chiefly due to the studies of Bigelow upon the more perfect
specimens found by the Albatross in the eastern part of the tropical Pacific.

The type species is Poralia rufescens Vanhoffen, from the Indian Ocean and tropical
Pacific.

Poralia rufescens Vanhoffen.
Poralia rufescens, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p- 41, taf. 4,

fign. 15-16; 1908, Deutsche Siidpolar Expedition, Bd. 10, Zool. 2, p. 47-—Bicetow, H. B., 1909, Mem. Museum Comp.
Zool. at Harvard College, vol. 37, p- 45, plate 13.

In Bigelow’s largest specimen the bell was 250 mm. in diameter. There were apparently
16 rhopalia, although some of these were destroyed so that the exact number was not deter-
mined with certainty. The rhopalia resemble those of Phacellophora and are set within deep

618 MEDUS2 OF THE WORLD.

niches. The sense-club is covered by a prominent scale, beneath which it stands in an almost
vertical position. There is a deep exumbrella sensory-pit above each sense-club. There
appear to be no distinct velar lappets, the bell-margins being only slightly wavy, excepting for
the deep, rhopalar clefts. In Bigelow’s large medusa 41 radial-canals arise from the periphery
of the circular central stomach, but two of these anastomose so that only 40 extend to the ring-
canal centripetal to the zone of the rhopalia. The mng-canal gives rise to a trident-shaped
diverticulum in the radius of each sense-club and to one or two simple, blindly-ending divertic-
ula in the inter-rhopalar spaces. The canal-system, like the rhopalia, bears a striking resem-
blance to Phacellophora. The mouth parts appear also to be similar to those of Phacellophora
but are not well preserved in any specimen yet captured. The gonads form a nearly continuous
ring around the periphery of the subumbrella floor of the stomach. This ring is not truly con-

Fic. 396.—Poralia rufescens, after H. B. Bigelow, in Mem. Mus. Comp.
Zool. at Harvard College.

tinuous, however, but is interrupted by 18 or 19 thickened, vertical ridges in the stomach-wall.
The gonads themselves thus consist of 18 or 19 outpocketings of the stomach-wall project-
ing outward. The numerous, simple, gastric cirri are arranged in a single line arising from the
stomach-wall on the inner side of the genital organs. The subumbrella is reddish-brown, the
gonads being paler. The tentacles were lost in all of the specimens so that we know nothing
of them.

Vanhoffen’s specimen came from a depth of about 350 fathoms between Queen Emma
Harbor and Siberut Island, Indian Ocean, and the two described by Bigelow were found by
the Albatross in the eastern part of the tropical Pacific.

The radial-canals in this medusa appear toincrease in number with growth, for Vanhéffen’s
specimen which was only about 60 mm. wide had 21 canals, while Bigelow’s 250 mm. wide
specimen had 41. Vanhoffen’s medusa had 7 or 8 (?) gonads, and both he and Bigelow believe
that the young medusa is probably octoradial.

Subfamily AURELINZ L, Agassiz, 1862.

The numerous tentacles and lappets arise from the sides of the exumbrella above the
margin. Gonads are invaginated sacs with external subgenital cavities. 4 simple or bifurcated
mouth-arms.

SEMAEOSTOMEA—AURELLIA. 619

Genus AURELLIA Péron and Lesueur, 1809.

(?) Evagora, Pron ev Lesueur, 1809, Ann. du Mus. Hist. Nat., Paris, tome 14, P- 343-

Aurellia, Ibid., p. 357-

Ocyroé, Ibid., p. 355-

Orythia, Lamarck, 1816, Hist. Anim. sans. Vert., tome 2, p. 502. Aurelia, Ibid. p. 512.

Medusa, Escuscuorrz, 1829, Syst. der Acalephen, p. 61.

Monoacraspedon + Diplocraspedon, Branvt, 1838, Mém. Acad. Impériale des Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, pp-

70; 372.

piblity tesco 1843, Hist. Nat. des Zoophytes, p. 339.

Claustra, Ibid., p. 378.

Aurelia, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, pp. 11, 72, 78, 159.—Acassiz, A., 1865, North Amer. Acal., p. 41.—
Craus, 1877, Denks. Akad. Math. eee + Wien., Bd. 38, p. 19.—Harcket, 1880, Syst. der Medusen, pp. 551, 644;
1881, Metagenesis und Hypogenesis von Aurela aria Jena, 36 pp., 2 taf.

Aurelia, von LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 279.—Goertre, A., 1887, Abhandl. zur Ent-
wickelungsgesch. Thiere, Heft 4, Leipzig (embryology).—Acassiz anv Mayer, 1899, Bull. Mus. Comp. Zool. at Harvard
College, vol. 32, p. 171.—VANnuOFrFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft 3, p. 23.—Bareson, 1895, Materials for
the Study of Variation, p. 428 (variations).—Browne, 1894, Quart. Journ. Micros. Sci., vol. 37, p. 245.—Hype, 1894,
Zeit. fiir wissen. Zool., Bd. 58, p. §35-—VANHOFFEN, 1888, Bibliotheca Zoologica, Heft 3, p. 19; 1902, Wissen. Ergeb.
deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 41; 1906, Nordisches Plankton, Nr. 11, p. 60.—Maas,
1906, Fauna Arctica, Bd. 4, Lfg. 3, p. 507 (discussion of literature); 1903, Scyphomedusen der Siboga Expedition, Monog
11, p. 26.—FR1IEDMANN, 1902, Zeit. fiir wissen. Zool., Bd. 71, p. 227-—Harairr, 1905, Journ. Exper. Zool., vol. 2, p. 548.

escomas HaeckeEL, 1880, Joc. cit., pp. 633, 644 (an Annan Aurellia with 16 sense-organs).

GENERIC CHARACTERS.

Ulmaridz with a simple, central mouth-opening which is surrounded by 4 well-developed,
radially situated, unbranched mouth-arms or palps. 8 marginal sense-organs. The tentacles
are small and alternate with an equal number of short lappets. Both tentacles and lappets
arise from the sides of the exumbrella a short distance above bell-margin. The bell-margin
is divided into 8 or 16 broad, velar lobes. The central stomach gives rise to a number of
branched, radiating canals which anastomose and are connected by a marginal ring-canal.
There are 4 interradial gonads and 4 well-developed, subgenital pits.

The name “‘ Evagora” which takes precedence over “Aurellia” was applied to Forskal’s
medusa persea which i is wholly unrecognizable, as is also ““Ocyroé,’ and these names must
therefore yield to “Aurellia,’ which was first proposed by Péron and Lesueur for durellia
aurita of Europe. They spell the generic name Aurellia.

The species of this genus are among the most widely distributed of Scyphomedusz, being
found in all oceans and all latitudes. They are most abundant along the shores of continents
and large islands and are comparatively rare in the open ocean far from land. It is possible
that the fossil Medusina costata from the lower Cambrian of Sweden is an Aurellia.

Although fully a dozen species of Aurellia have been described I believe that there are but
3 reasonably well-defined types. Of these 4. aurita is of world-wide distribution. 4. labiata
is found in the Pacific, and a third fairly well-defined species is 4. maldivensts, described by
H. B. Bigelow from the atolls of the Maldive Islands in the Indian Ocean. The distinctions
between many of the “species” are not well ascertained, and there are numerous varieties or
local races. A. aurita is subject to great individual variation and some of these chance varia-
tions have been described as species.

The species of Aurellra display much individual variability, and studies upon this subject
have been carried out by Ehrenberg (1835), Romanes (1876-77), Browne (1894-95), Duncker,
Sorby, Herdman, Unthank (1894), "Ballowitz (1898), and Hargitt (1905). Good reviews of the
results of the earlier of these investigations are given by Bateson, 1895 (Materials for the Study
of Variations, p. 426), and by Agassiz and W oodworth, 1896 (Bull. Mus. Comp. Zool. at
Harvard College, vol. 30, No. 2). The abnormal individuals of Aurellia aurita are peculiar in
that they generally preserve the radial symmetry of the disk, even though the number of
segments be changed. Radially symmetrical abnormalities appear to be about twice as
numerous as are irregular ones. This law applies also to the variations of the Leptomedusa
Pseudoclytia pentata (see vol. II, p. 278).

Browne, 1894 and 1895, discovered that congenitally abnormal ephyre of Aurellia survived
fully as well as normal ones, and also that the abnormal ephyre were no more abundant in
1892 than they were when Ehrenberg studied them at the same place in 1834. Evidently the
abnormal individuals do not acquire any fixed tendency to perpetuate their own peculiarities

620

MEDUSA OF THE WORLD.

rather than those of their normal parents, and thus the race as a whole maintains itself

unchanged.

than 8 marginal sense-organs.

Most interesting physiological studies have been carried out upon Aurellia by Romanes,
1885 (International Scientific Series, vol. 49, etc.), and also by Eimer, 1878. These studies
show that the marginal sense-organs of Aurellia are locomotor centers which control the
rhythmical pulsation. These marginal sense-organs tend to send out impulses to pulsation
at various rates, but the fastest working sense-organ controls all the others and forces them
to beat at its own rate. Parts of the subumbrella deprived of marginal sense-organs will still

Shape of umbrella.

Width in mm.
Height in mm.

Form of margin of
umbrella.

Number of velar
lobes.

Form of mouth-
arms.

Synopsis of the Races or Species of Aurellia.

About 22 per cent of the ephyra and of adult Aurellia have either more or less

Aurellia aurita and its varieties.

Length of mouth-
arms in terms of
radius of um-

brella (r).

Length of genital-
radius in terms of|
radius of um-
brella.

Number of pri-
mary branches
arising from each
interradial geni-
tal sinus.

Character of ad-

radial canals.

Color.

Where found.

Remarks.

A. aurita Péron et
Lesueur.

A. cruciata Haeckel.
(This is only a
variety of A.aurita.)

A. colpota Brandt
=A. coerulea von
Lendenfeld.

A. flavidula Péron
et Lesueur (this is
only a variety of A.
aurita)=A, haban-
ensis Mayer.

A. hyalina Brandt.

Flatly rounded to
hemispherical.

50 to 400

20 to 125

The 8 marginal
sense-organs set in
shallow clefts, 8

simple entire, velar
lobes.

8 simple.

Small,lancet-shaped
with complexly
folded margins, but
without lateral
lappets.

—

One-third r.

5 to 7, quite wide.

Simple, unbranched.

Very variable.
Milky to light-vio-
let, rose-red, or
almost colorless.

Atlantic coast of
Europe and in
Mediterranean.
Vast swarms.

Development
through alterna-
tions by strobiliza-
tion.

Flatly rounded to
hemispherical.

50 to 400
20 to 125

Sense-organs set in
deep clefts.

8 simple.

Small,lancet-shaped,
with complexly
folded margins, but
without lateral
lappets.

Only a little longer
than o.5r.

Half r.

5to7

Simple, unbranched

Very variable.
Milky to light-violet
rose-red, or
almost colorless.

Atlantic coast of
Spain and Medi-
terranean.

Flatly rounded to
hemispherical.

100 to 120
40 to 50

Sense-organs set in
shallow clefts.

8 slightly notched in
middle.

Large, complexly
folded, with lateral
lappets. Very wide
at their bases.

Tata

Half r.

Anastomosing with
other canals, or
simple and un-
branched.

Light rose-red ten-
tacles and gonads
deeper in color.

Indian Ocean to
Pacific.

Flatter than a hemi-
sphere.

140 to 250

50 to go

Sense-organs set in
clefts which may
either be deep or

shallow, being very
variable.

8 simple.

Thick, lancet-
shaped, with com-
plexly folded
edges.

Cae

One-third to half r.

Simple, unbranched.

Variable as in A.
aurita.

Atlantic coast of
North America,
Greenland to
Florida and West
Indies.

Development as in
A. aurita.

Flatter than a hemi-
sphere.

60 to 80
20 to 30

Sense-organs set in
wide, deep clefts.

8 simple.

Similar to A. aurita,

One-third to one-
fourth r.

All canals branched.

Variable. Trans-
parent to reddish.

North Pacific.

SEMAEOSTOMEZ—AURELLIA.

621

respond momentarily by contractions to all sorts of stimuli, electrical, chemical, thermal, o1

mechanical.

slowly than large ones.

Small parts of the disk with sense-organs attached pulsate somewhat more
Romanes succeeded in maintaining the rhythmical movement of the

medusa in parts of the disk without marginal sense-organs by stimulating weakly with a con-
stant or alternating current of electricity.

Romanes also found that a stimulus too weak to cause a response would if repeated
eventually give rise to a contraction.
and it is interesting to see that Lee and Morse, 1910 (Proc. Soc. Experimental Biology and
Medicine, New York, vol. 7, p. 38), find that this effect may be due to a rise in irritability,
brought about by the action on the living substance of small quantities of certain products
of metabolism, especially carbon dioxide and lactic acid, the same substances which in greater
concentration are important factors in fatigue.

Synopsts of the Races or Spectes of Aurellia—Continued.

This phenomenon is known as the summation of stimuli,

Aurellia aurita and its varieties.

A. dubia Vanhoffen.

A. vitiana Agassiz and
Mayer (immature).

A. marginalis L. Agassiz,
(This is only a variety
of A. flavidula.)

Shape of umbrella.

Width in mm.
Height in mm.

Form of margin of um-
brella.

Number of velar lobes.

Form of mouth-arms.

Length of mouth-arms
in terms of radius of
umbrella (r)

Length of genital-radius
in terms of radius of
umbrella.

Number of primary
branches arising from
each interradial genital
sinus.

Character of adradial
canals.

Color.

Where found.

Remarks.

Flatter than a hemi-
sphere.

130

3
As in A. flavidula.

8 simple.

Mouth-arms give rise to
lateral lappets.

Two-thirds r.

One-third r.

All canals branched.

Persian Gulf.

Described from a single
specimen.

Hemispherical.

80

40
As in A. flavidula.

8 simple.
Simple lancet-shaped.

Half to two-thirds r.

One-fourth to one-third r,

5 to 7, all very slender.

Simple, unbranched.

Gonads, mouth-arms,
and tentacles lilac.
Other parts colorless.

Fiji and Tonga Islands,
South Pacific.

Swarms in harbors.

Flatter than a hemi-
sphere.

160 to 300
60 to 120
As in A. flavidula.

8 simple.
As in A. flavidula.

Half r+.

7. As in A. flavidula.

Simple, unbranched.

Gonads often light rose-
red, or blue, but colors
variable as in A. flavi-
dula.

Florida Keys, Key West,

Havana to coast of
Maine.

(?) Development
slightly different from
that of A. flavidula
(see Hyde 1894).

A. solida Browne.

Hemispherical.

80

40

As in A. aurita but
sense-organs are set in
deep clefts and point
upward toward exum-
brella, not toward bell-
margin. The 8 sense-
clubs are thus di-
rected go° away from
direction assumed by
A. aurita.

8 simple.

Closely similar to those
of A. aurita. The sub-
genital ostia are very
small circular openings|
only 2 mm. wide.

rae

Four-fifths r.

As in A. aurita.

As in A. aurita.

Gonads salmon-colored,
other parts translucent
whitish.

Maldive Islands, Indian
Ocean.

Distinguished by its
peculiar sense-organs.

622 MEDUSA OF THE WORLD.

Krukenberg, 1880, finds that durellia aurita contains 95.34 per cent of water and only
4.66 per cent of solid matter.

The planula larva commonly develops into a scyphostoma which gives rise to a number
of ephyre through strobilization. In aquaria, however, Haeckel, 1881, finds that the planula
may develop directly into a single medusa without passing through the scyphostoma-stage.
In this case the ocular lobes and tentacles grow outward around the gastrula mouth, and the
gelatinous substance of the planula becomes that of the medusa. In other cases the scypho-
stoma develops into a single medusa which remains attached by a pedicel formed of the basal
part of the scyphostoma, recalling the condition observed in the Stauromedusz.

Hérouard, 1907 (Comptes Rendus, Paris, tome 145, p. 601, [bid., 1908, tome 147, p. 1336),
finds a peculiar scyphostoma in an aquarium at Roscoff, which may possibly be that oF Aurel-
lia affected by adverse conditions of confinement, although he calls it T aeniolhydra roscof-

fensis.

Synopsis of the Races or Species of Aurellta—Continued.

It develops lateral buds, and in addition to these peculiar cysts on its pedal zone.

Aurellia labiata and its varieties.

A. labiata Chamisso and

Eysenhardt *

A. “clausa” Lesson=
young of A. labiata.*

A. limbata Brandt.

A. maldivensis Bigclow.*

Shape of umbrella.

Width in mm.

Height in mm.

Form of margin of um-
brella.

Number of velar lobes.

Form of mouth-arms.

Length of mouth-arms
in terms of radius of

umbrella (r).

Length of genital-radius
in terms of radius of
umbrella (r)

Number of primary
branches arising from
each interradial genital
sinus.

Character of adradial
canals.

Color.

Where found.

Remarks.

Hemispherical or flatter.

200 to 300
100 to 200

Margin with 16 deep
clefts, 8 ocular, 8 inter-
ocular, 2 velar lobes in
each octant.

16, two in each octant.

Thick, pyramidal, short,
folded when mature,
but slender and simple,
as in A, aurita when
young.

‘Two-thirds to three-
fourths r.

One-fourth r.

As in A, aurita.

Usually simple, but may
branch dendritically
(without anastomos-
ing).

Light violet, gonads
darker in color.

Pacific coast of North
America to Malay
Archipelago.

Hemispherical or flatter.

80 to 100
40 to 50

Margin with 16 clefts,
2 velar lobes in each
octant.

16, two in each octant.

Small, thin, folded,

~

~

Ovaries, canals, tenta-
cles rose-red to wine-

red.

South Pacific coast, New
Ireland, Australia.

Too imperfectly known
for determination.

Hemispherical or flatter.

200 to 300
100 to 150

Margin with 16 deep
clefts, 2 velar lobes in
each octant.

16, two in each octant.

Triangular.

Half r.

All canals anastomose.

Umbrella bluish, ten-
tacles and margin
orange-brown. Velar
lappets brownish-
black.

North Pacific coast of
Siberia.

Flatter than a hemi-
sphere.

250
go

16 velar lobes with very
shallow interocular
clefts.

16, two in each octant.
Very shallow clefts be-
tween lappets.

Lips large, wide, flexible,
curtain-like and folded
with their free edges
lined by small tenta-
cles.

cos

Cne-fourth to one-third
—s

8 to 10 branched and
anastomosing. Al-
together about 48
canals arise from cen-
tral stomach.

Simple, unbranched.

Variable. Bell delicate
lilac. Canals and ten-
tacles violet-pink.
Gonads violet or blue.

Maldive Islands, Indian
Ocean, in January.

This form is chiefly
distinguished by its

Probably identical with curtain-like lips re-
A. limbata or A. abit calling those of Cyanea.

*These are probably one and the same species and should be called Aurellia labiata

Pate 68.

Fig. 1. Aurellia aurita. Eastport, Maine, August 24, 1897. Natural size.

Fig. 2. Aurellia aurita. Natural size of quadrant of bell. Tortugas, Florida,
May 4, 1905.

Fig. 3. Aurellia aurita. Showing a marginal sense-organ seen from the
exumbrella side. Showing tentacles arising from exumbrella
side of bell-margin. Tortugas, Florida.

Fig. 4. Aurellia aurita. The bell-margin from the subumbrella side showing
the marginal lappets. Tortugas, Florida.

Drawn from life, by the author.

PLATE 68

MAYER

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/) (eM,

Tae

rr

\
((

AN

AW Say mn (Ke LU)

\

Se a AN
SON wSy
IRQ AAS

V\\ A ROARS ARS
\ JA WSS
\'— \\S

\
\ \ S
aN \ SS SS
\ \ ~ S
] NARS \ NS \

WY DARA SS
SS \\\\

SEMAEROSTOMEZ—AURELLIA. 623

These cysts or statoblasts are formed during resting periods in about 15 days and are incased
ina chitinous envelope. After one of these statoblasts is formed the scyphostoma moyes a short
distance and leaves it behind. The chitinous envelope may then burst and the cyst develops into
a polyp with tentacles. When fed upon ovary of the sea-urchin this scyphostoma strobilated.

The early development of the planula and scyphostoma has been elaborately studied by
Goette, Claus, Hyde, Smith, Friedmann, and others and the results of these researches will be
reviewed under Aurellia aurita.

Friedmann, 1902, finds that in the scyphostoma the tentacles develop in the order of
4, 8, 16, 24; and that the 12 and 20-tentacle conditions are intermediate phases.

Aurellia aurita Lamarck.
Plate 67, fig. 4; plate 68, figs. 1 to 4.

Medusa aurita, Linnf, 1746, Fauna suecica, No. 1287; 1747, Westgita Resa, tab. 3, fig. 2; 1758, Syst. Nat., Ed. 10, tomus 1
p- 660.—GMeLIN, 1788, Linne’s Syst. Natura, tomus 1, pars 6, p. 3153-

Medusa, sp., Bortasr, 1758, Nat. Hist. of Cornwall, p. 257, plate 25, figs. 1x and x.

Aurelia aurita, Lamarck, 1816, Hist. Anim. sans. Vert., tome 2, p. 513—EnRenzeERG, 1834, Abhandl. Kgl. Akad. Wissen.,
Berlin, p. 99 (variations).—Detre Cutaje, 1841, Animali senza Vert., tome 7, Napoli, tav. 144, fig. 1; tav. 145, fig. 2.—

» Romanes, G. J., 1875, Nature, vol. 2, p. 29; 1876, Journal Linnean Soc. London, Zool., vol. 12, p. 528; vol. 13, P- 190,

plates 15, 16 (variations); 1885, Jellyfish, Star-fish and Sea-urchins, etc., International Scientific Series, vol. 49 (physiology
of pulsation).—Harcket, 1880, Syst. der Medusen, p. 552 (important list of literature)—Mortus, 1880, Zool. Anzeiger,
Jahrg. 3, p. 67 (medusa killed by freezing )—KruKenBeErG, 1880, Zool. Anzeiger, Jahrg. 3, p. 306 (water content of body).—
Harcket, 1881, Metagenesis und Hypogenesis von Aurelia aurita, Jena, 36 pp., 2 taf; Kosmos, Bd. 5, p.29.—M’Kenprick,
1881, Journ. Anat. and Physiol., vol. 15, p. 261 (coloring matter).—Morivs, 1882, Zool.Anzeiger, Jahrg 5, p. 586. (Aurellia
from Kiel contains 97.92 per cent of water.)—Craus, 1883, Untersuch. tiber Organization und Entwicklung Medusen,
Prag. Leipzig, pp. 1-23, taf. 1, fign. 1-20 (development).—Graerre, 1884, Arbeit Zool. Inst. Wien, Bd. 5, p. 343.—Gorrr,
1885, Zool. Anzeiger, Jahrg. 8, p. 554 (development); 1887, Abhandl. Entwickelungsgeschichte der Thiere, Heft 4, Leipzig.
(development).—Marsua tt, 1888, Quart. Journal Microscop. Sci., vol. 28, p. 84, (striped muscles).—Scnewiakorr, 1889,
Morphol., Jahrb., Bd. 15, H, 1 (structure of sense-club).—Muncuin, 1889, Proc. Zool. Soc. London, p. 583, 2 plates (brood-
pouches of the oral arms).—Craus, 1890, Arbeit. Zool. Inst. Wien, Bd. 9, p. 85 (development).—G6rrer, 1891, Claus und
die Entwicklung der Scyphomedusen, Leipzig. 64 pp-, 24 fign. (controversy over the manner of development).—-CLaus,
1892, Arbeit. Zool. Inst. Wien., Bd. 10, p. 20 (development of the scyphostoma).—Duncker, 1895, Archiv. fiir Naturgesch.,
Jahrg. 60, p. 7, plate 1 (variations).—Browne, 1894, Nature, vol. 50, p. 524 (variations).—Sorsy, 1894, Nature, vol. 50,
p- 476 (variations) —HerpMan, 1894, Nature, vol. 50, p. 426 (variations).—Browne, 1895, Quart. Journal Microscop.
Sci., vol. 37, p- 245, plate 25, figs. 1-7; 1901, Journal of Biometrica, vol. 1, p. 90, 3 diagrams (comparative variability of
ephyre and of adult meduse).—Haacke, 1897, Zool. Garten, Jahrg. 37, p- 357-—Battow11z, 1899, Archiv. Entwickelungs-
mech., Bd. 8, p. 239, taf. 5 (variations).—Hen, 1900, Zeit. fiir wissen. Zool., Bd. 67, p. 401, 5 text-figs., taf.24, 25 (develop-
ment).—Gdorre, 1900, Zool. Anzeiger, Bd. 23, p. 559 (development).—FR1EDMANN, 1902, Zeit. fiir wissen. Zool., Bd. 71,
p- 227, taf. 12, 13 (development of tentacles in scyphostoma).—He1n, 1902, Zool. Anzeiger., Bd. 25, p. 637 (development).—
VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Dampfer Valdivia, Bd. 3, Lfg. 1, pp. 43, 44 (world-wide dis-
tribution of 4. aurita); 1906, Nordisches Plankton, Nr. 11, p. 60, fign. 27-31.—Detap, 1906, Fisheries Ireland, Sci. Invert.,
1905, No. 7, p. 22, 1 plate (development).—Goopey, 1909, Proc. Zool. Soc. London, part 1, p. 78, plate 24.—HansTrIn,
1907, Naturgesch. des Tierreichs., p. 332, fig. 446.

Aurelia cerulea, VoN LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 280.

Aurelia colpota, G6rrTE, 1886, Sitzungsber. Akad. Wissen. Berlin, Jahrg. 1886, p. 836.

Aurelia aurita var. colpota, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 26, taf. 1, fig. 9 —Browne, 1905,
Fauna and Geog. Maldive and Laccadive Archipelagoes, vol. 2, p. 959.

Aurelia japonica, Kisu1novuye, 1891, Zool. Magazine, Tokyo, vol. 3, No. 33, p- 289, plate 7.

Aurelia vitiana, Acassiz, A.. anv Mayer, 1899, Bull. Museum Comp. Zool. at Harvard College, vol. 32, p. 171, plate 10, fig. 35;
1902, Memoirs, Jbid., vol. 26, p. 159.

Aurelia dubia, VANHGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 20, 24.

Aurelia cerulea, VON LENDENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 280.

Aurelia cruciata, HAECKEL, 1880, loc. cit., p. 644.

Aurellia aurita from America.
Plate 67, fig. 4; plate 68, figs. 1 to 4.

Medusa aurita, Fanricius, 1780, Fauna Groenlandica, p. 363, Nr. 356.

Aurellia flavidula, Pixon er Lesurur, 1809, Ann. du Mus. Hist. Nat. Paris, tome 14, p. 359, No. 92.

Aurelia flavidula, Lesson, 1843, Hist. Zooph. Acal., p. 376.—Acassiz, L., 1860-62, Cont. Nat. Hist. U. S., vol. 4, pp. 10, 12, 51,
160; vol.3, plates 6-9, 11a, 12b; plate 10, figs. 18, 22, 31, 32, 36; plate 10a, figs. 4b, 13, 15a, 16-41; plate 11, figs. 1-13
(development).—Pacxarp, 1863, Canadian Nat. and Geol., vol. 8 —Acassiz, A., 1865, North Amer. Acal., p. 42, figs. 65,
66.—HaeckEL, 1880, Syst. der Medusen, p.555.—Fewkes, 1881, Bull. Mus. Comp. Zool. at Harvard College, vol. 8, p.172,
plate 7, figs. 2-4, 6; 1884, Mem. Mus. Comp. Zool. at Harvard College, vol. 10, No. 3, plate 8.—Smitn, 1891, Bull. Mus.
Comp. Zool. at Harvard College, vol.22, p. 115, plates 1, 2, figs. 1-12 (development).—HypeE, 1894, Zeit. fiir wissen. Zool.,
Bd. 58, p. 535, taf. 24, figs. 36-53 (development)—Verritt, 1873, Report U.S. Commiss. Fish and Fisheries for 1871-72,
P- 723-—Macatium, 1903, Journal of Physiology, Cambridge, England, vol. 29, pp. 213-241 (inorganic composition).—
Harcitr, 1904, Bull. U.S. Bureau of Fisheries, vol. 24, p. 67 (scyphostome with stolons).—1905, Journal Exper. Zool.,
vol. 2, p. 548, figs. and plate (variations).

Medusa aurita, Escuscuoxtz, 1829, Syst. der Acal., p. 61.

Aurelia flavidula, A. aurita, Ephyra octolobata, Goutp, 1841, Report Invert. Massachusetts, p. 348.

Aurelia sex-ovalis, Morcu, 1857, Beskriv. af Gronland, p. 95.

Aurelia aurita, Morcn, 1857, Beskriv. af Gronland, p. 95.—Stimrson, 1853, Marine Invert. Grand Manan, p. 11.

Aurelia habanensis, Mayer, 1900, Bull. Mus. Comp. Zool. at Harvard College, vol. 37, p. 69, plate 24, figs. 73, 74; plate 26,
fig. 86 (Cuba to Tortugas).

624 MEDUS® OF THE WORLD.

Dimensions in mm. of a mature specimen of Aurellia aurita from Naples, Italy, captured
May 30, 1900. Bell-radius, 85; length of mouth-arms, 75; width of genital cross, 51. (See
text-hgure 397.)

The following is a description of the American form of Aurellia aurita, which has been
commonly called ‘4. flavidula”’:

Adult medusa.—Disk 140 to 250 mm. wide, 50 to go mm. high. When expanded it is
flatter than a hemisphere, but when contracted it becomes hemispherical. Gelatinous sub-
stance tough, thick at center, but thin at edge of disk. 8 marginal sense-organs at the bottom
of shallow niches between the 8 broad velar lappets. Each sense-club is blunt and contains
an ectodermal, proximal ocellus on the exumbrella side, a cup-like pigmented eye composed
of both ectoderm and entoderm on the subumbrella side, and a distal entodermal mass of
crystalline concretions. It is partially protected above by a bridge-like web stretching between
the adjacent lappets. Lappets of considerable size on both sides of the club.

A single median pit projects downward from exumbrella surface just above the sense-
club. Small and numerous tentacles arise from sides of disk at a slight distance above margin
and alternate with an equal number of small, elongate, marginal lappets which similarly arise
from the sides of the disk above the projecting margin. Tentacles hollow, with longitudinal
strands of muscles down their subumbrella sides, and with broken rings of nematocyst-cells

on their exumbrella sides. Margin of disk

entire and simple except at places of the 8

sense-organs, where it is broken by notches.

It forms a narrow, velum-like structure

% lying below the tentacles and marginal lap-

~e pets. Central mouth-opening 4-sided, sur-
A
a

A QUAsttA00 Fy MUU
RC May
RC | yy

rounded by 4 thick, stiff, gelatinous mouth-
arms, the 8 free edges of which are much
convoluted and provided with a row of
numerous, small tentacles. These free
edges inclose a median trough or gutter,

Nh)

=e
== which extends down the middle of the lower
= side of each mouth-arm. These mouth-
2 arms are each about as long as the radius
S) of the disk; at their bases they are broad

‘s and their free margins are here greatly

indented and folded in sinuous lines bor-

dered by small tentacles. The 4 interradial

gonads are horseshoe-shaped and alternate

Ai with the mouth-arms, and their position

Fic. Sala aurita. = by the author, from a specimen is marked on the floor of the subumbrella
ound at Naples, Italy, by Dr. S. Lobianco at the Naples : . E

Zoological Station, May 30, 1900. by 4 thick, horseshoe-shaped thickenings

of the gelatinous substance, in the center

of each of which there is a deeply sunken, subgenital pit. The radius of each of these horse-

shoe-like regions is about one-third that of the disk itself.

Goodey, 1909, finds 4 interrradial, canal-like grooves in the subumbrella floor of the
stomach leading from the gonads to the folds of the oral arms and serving to conduct the
genital products away from the gonads.

The central stomach occupies a 4-lobed space, its outline being determined by the periph-
eral edges of the 4 horseshoe-like genital cavities, and it gives rise to a complex system of
radiating canals extending from edges of stomach to circular canal at bases of marginal ten-
tacles. 8 separate, straight, non-anastomosing, adradial canals and 4 radial and 4 inter-
radial, pitchfork-shaped systems of anastomosing canals; each of these systems consists of
a single, straight, median canal and a pair of branching, lateral canals, which arise on either
side of the median canal very near the periphery of the central stomach-cavity. The lateral
canals each give rise to 3 to 5 radiating branches which anastomose sparingly and extend
outward to the circular canal. These branches decrease in caliber and anastomose more
frequently as they approach the circular canal. The gonads are found in 4 horseshoe-shaped

SEMAEOSTOMEA—AURELLIA. 62!

convoluted ridges on the subumbrella floor of the 4 genital cavities. The bases of the genital
ridges are beset with numerous, small, gastric cirri.

The gelatinous substance of the disk is of a translucent milky-white or yellowish-brown;
spermaries usually slightly pink. In old individuals the gonads in both sexes are white.

Common from Greenland to the West Indies. At Eastport, Maine, it is mature in Sep-
tember, and at Tortugas, Florida, 1 in May.

The American medusa is closely allied to Aurellia aurita of Europe and is at most
merely a variety of the latter. It may possibly differ from its European representative in
the thickness and rigidity of the mouth-arms, which are very broad at their bases and often
complexly convoluted at their free edges, but I have seen these same characters in Aurellia at
Naples and consider the American and European medusz to be identical.

A very complete description and numerous figures of the American medusa are given
by L. Agassiz, 1860-62, Cont. Nat. Hist. U. S., vols. 3 and 4.

Development.—The ova are dehisced from the gonads into the interradial grooves from
which they enter the median gutter of the mouth-arms and are here retained in small pouches
near margins of free edges of mouth-arms and finally set free in the planula stage. Minchin,
1889, gives a good description of these brood- -pouches. Segmentation total and unequal, anda
blastula is formed which has a large, central blastoceel. According to Hyde, 1894, the gastrula
may be formed in either one of two different ways: (1) by the invagination of a small part of
the blastula wall combined with the ingression of numerous cells from various parts of the wall
of the blastula; (2) by invagination of the wall of the blastula, aided only occasionally by the
ingression of cells from the blastula wall. According to Smith, 1891, however, the gastrula is
formed from a small invaginated region in the wall a the blastula, from which there dev elops
a single, continuous layer of cells, w hich layer finally completely fills the cleavage cavity, thus
giving rise to a 2-lay ered embryo with an open blastopore. Smith denies that this process 1s

aided in the least by the ingression of cells from the wall of the blastula into the blastoccel. He
finds, indeed, that a few cells are occasionally seen to wander into the blastula cavity, but these
always degenerate without taking any share in the formation of the entoderm. These varia-
tions in the mode of forming the gastrula have been seen in other Scyphomedusz, haying been
observed by Conklin in Diaiche, and Hyde and MeMurrich in Cyanea. The blastopore
then closes and the entoderm becomes a closed sac entirely env eloped by the ectoderm. The
larva then becomes ciliated and swims actively about as a pear- shaped planula, which soon
attaches itself to the bottom by the wide anterior end. A crater-like depression (cesophagus)
formed of ectodermal cells then appears at the narrow (now the upper) end of the animal, and
this presses down upon the entodermal sac. The first pair of radial stomach-pouches is formed
from the entodermal sac, while the second pair is formed, at least partially, from the ectoderm
of the cup-like depression. The mouth breaks through and 4 tentacles appear. The larva
then has 4 interradial, longitudinal, partial septa a formed of a fold of the entoderm sup-
ported by a central shelf of gelatinous substance. These septa extend from the margin of the
mouth to the lower end of the stomach-cavity. They form the 4 primary, gastric filaments
of the future ephyra, and there are no septa in the central stomach of the medusa.

As we have stated, it appears from the researches of Gotte and of Hyde that the two
original pairs of stomach- pouches are derived alternately from the ectoderm of the cesophagus
and from the entoderm of the primitive stomach, although Hyde shows that the lower aboral
floor of the 2 cesophagus pouches is formed at least partially from entoderm. Through
division of the 4 original stomach-pouches we have finally 24 pouches, 10 entodermal and 14
mainly ectodermal, as follows: 6 diammetrically opposite perradial, 4 interradial, and the 4
connecting adradial pouches are ectodermal. go° apart from these the 6 perradial and their 4
adjacent adradial pouches are entodermal. (See Gotte, 1887.)

One must remember that R. P. Bigelow, 1900, finds that the 4 primary stomach-pouches
of Casstopea xamachana are wholly entodermal, and HadZi, 1907, finds that this is also the case
in Chrysaora. Moreover, according to Hadzi there is no ectodermal invagination in Chrysaora
to form the mouth, but on the contrary the throat is evaginated and ined with entoderm.

The larva of Aurellia becomes a scy phostoma which finally attains a height of about 5 mm.
and acquires 8, 16, and finally 24 long tentacles. The ephyre are dev eloped through strobili-
zation of the scyphostoma. As many as 13 annular constrictions may develop below the zone of

626 MEDUS OF THE WORLD.
oral tentacles, and then an additional set of tentacles usually develops below the last con-
striction. As many as 12 disk-like ephyra may be cast off one by one, and finally the scypho-

stoma is left greatly reduced in size, but still provided with a corona of tentacles. After all
of the ephyre have been cast off through this terminal budding the scyphostoma frequently
develops irregular stolons from its sides and base. Haeckel’s observations of other modes
of development are discussed in the description of the genus Aurellra.

The young ephyra has 8 marginal sense-organs flanked by 16 lappets, the ocular clefts
of which are only about half as wide and deep as the 8 alternating clefts. The throat is at
first a simple 4-cornered tube and the lenticular central stomach gives rise to 16 simple,
separate, radiating canals which extend outward in the radii of the tentacles and sense-organs.
The tentacles then begin to develop (first one, then others laterally) in the 8 adradial spaces.
The lips elongate at the 4 perradial corners and form the mouth-arms, and a peripheral
ring-canal is formed by the radial-canals becoming T-shaped at their free, distal ends, and the
sides of each adjacent T fusing. As the animal increases in size, blind fale aaeel centrip-
etally inward from the ring-canal even before the ring-canal is complete, and fuse with the 8
perradial and interradial canals, which thus become pitchfork-shaped. (See hg. 4, plate 67.)

Full descriptions and very complete figures of the development of the various stages in
Aurellia flavidula are given by L. Agassiz, 1860-62. Hyde, 1894, gives a detailed and careful
account of the development of the planula and of the early stages of the scyphostoma, and
Smith has investigated the process of formation of the gastrula. Claus and Gotte, whose
views are at variance, studied the development of the scyphostoma and its gastral pouches.

Macallum, 1903, finds that, in Aurellia flavidula living in brackish water, the salinity
of the water may undergo considerable change during the day and yet the amount of NaCl
within the body of the Aurellia remain practically constant. The medusa contains slightly
less sodium and considerably more potassium than does normal sea-water. It contains
also about the same amount of calcium as is found in sea-water, but less magnesium and 32
to 36 per cent less SO. He gives the composition of Aurellra, Cyanea and of sea-water as
follows:

, “A. flavidula,” “
A. aurita Lamarck, this name being the oldest.
A. aurita is distributed over all warm and temperate oceans.

Na. Ca. K. Mg.
Contents of sea-water............-.- 100 3.84 3.66 11.99
Contents of body-juice of Aurellia... 100 4-13 5.18 11.43
Contents of Cyanea arctica........- 100 3-86 7-67 11.31

27 mature specimens of Aurellia from Tortugas, Florida, all collected at random from a
single swarm on May 4, 1906, were of dimensions and proportions as follows:

Armong 27 spectasns= Radius of Length of Radius of
umbrella. mouth-arms. | genital cross.

EA PERE SPOCLMIEN <1 in,ainlalalctelsivists iol taiaiercietcraieiaraistereeteiatetateiy 92 mm. go mm. 32 mm.
Smallest: specimens scresinys nelly s.eeielee encore esr eeienatatst 59 mm. 61 mm. 25 mm.
IS fafa teller ering doplenicnacancindgas cagasnessenr 78 mm. 74 mm. 33 mm.
Proportions of specimen with longest mouth-arms. ...... r 1.047

With: amallest mouthearmis e's ccs sacle sie ain/sets olem)atare r -89r

Average length of mouth-arms...........-+++e+e+-e0e r 957

Proportions of specimen with largest gonads..........-. ir o.5ir
Smallest gonads a6 core ate lateic sie clei tiesweleletarsiclevielels ice eters r 0.36r
Average ponaden ences nelaate atta r 0.42Fr

A. marginalis,”

and “4. habanensis”;

Thus individuals among these 27 specimens displayed all of the characteristics of Aurellia
and all should be called
I wholly agree with Vanhéffen, 1902, that

SEMABOSTOMEAS—AURELLIA. 627

Aurellia aurita forma ‘“marginalis.”

Aurelia marginalis, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, pp. 86, 160.—AGassiz, A., 1865, North Amer. Acal. rP- 43° -—=
Harcxet, 1880, Syst. der Medusen, p. 556. Sst 1894, Zeit. fiir wissen. Zool., Bd. 58, pp. 532, 544, taf. 32, 33,
fign. I-35. “
This variety is larger than 4. flavidula, being often more than 300 mm. in diameter.

Mouth-arms smaller than in 4. flavidula, being less than bell-radius in length. Genital

pouches fully half as wide as bell-radius, instead of being about one-third this width as in

A. flavidula. he gonads are of a pale rose color in both sexes.

Professor Hyde has made a careful study of the development up to the scyphostoma
stage. The gastrula results from a peculiar process of delamination. Some of the cells of
the one-layered blastula divide and their inner halves thus become free and wander into the
blastula cavity where they eventually form the entodermal layer.

According to Hyde, however, the development of 4. flavidula is itself subject to much
variation and the gastrula in this form also results in some cases from delamination. Hence
the peculiar features of the development in 4. marginalis are different only in degree from
those observed by Hyde in 4. flavidula and are not of specific importance. Moreover, we
must bear in mind that Smith finds that the gastrula of durellia flavidula is formed by invagina-
tion, and according to Gotte and to Hein, 1900, the gastrula of 4. aurita is also formed by
invagination. Staleness of the water in ordinary aquaria may profoundly alter the normal
course of development.

I am convinced that “ Aurellia marginalis” is only a variety of ‘A. flavidula,” which is
itself specifically identical with 4. aurita. For example, among 27 mature specimens of
Aurellia found in a single swarm at Tortugas, Florida, on May 4, 1906, if we call r the radius
of the umbrella, the length of the mouth-arms ranged from 0.89 to 1.04 r, the average being
0.95 7; also the dae of the genital cross ranged from 0.36 to 0.51 r, the average being 0.42 r.
Agassiz called specimens of Anvellia with arms longer than r and the genital Sardi less than
0.5 r “A. flavidula’’; those having arms less than r and genital radii more than 0.5 r he
would call “‘4. marginalis.” It is evident, however, that the two forms intergrade, and this
is true not only along the Florida reef, but also in the harbor of Eastport, Vieune: where I
found some individual Aurellias that conform to the proportions of “A. marginalis.” It is
safe to conclude that “‘4. marginalis” is merely a manuscript species and anatihl disappear
henceforth. L. Agassiz described it from the Florida reefs.

Aurellia aurita =“‘Aurellia dubia” Vanhoffen.
Aurelia dubia, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 20, 24.

The disk is 130 mm. wide and 43 mm. thick. The 8 marginal sense-organs are set in
deep niches, as in 4. flavidula, and there are 8 broad, marginal lappets. he 8 mouth-arms
are only two-thirds as long as the disk-radius. Genital- ins one-third of disk-radius; 7
radiating canals extend outward from each genital sinus. All of the canals fork; their branches
are narrow and elongate in the middle and small and numerous at the margin. Color ( ?)
Persian Gulf, March.

Described from a single specimen by Vanhoffen. Distinguished by its short mouth-
arms and its having only 8 velar lobes instead of 16 as in 4. labrata.

Aurellia solida Browne.
Aurelia solida, Browne, 1905, Fauna and Geog. Maldive and Laccadive Archipelagoes, vol. 2, plate 960, plate 94, figs. 1, 2;

1908, Trans. Royal Soc. Edinburgh, vol. 46, p. 249.

This medusa differs from Aurellia aurita in its marginal sense-organs, but in all other
respects the characters of 4. solida are well within the common ranges seen in A. aurita. In
Aurellia solida each marginal sense-organ arises from the inner end of a deep groove
which is open on the exumbrella side and bordered by the lateral lappets, but closed on the
subumbrella side. In Aurellia aurita the sense-club points outward toward the umbrella
margin. In Aurellia solida, however, the sense-club points upward toward the exumbrella
and eherefore at right angles to the position assumed by the sense- -club in Aurellta aurita.

In Aurellia aurita fiereli is a well-developed covering membrane or “hood” which extends
over and above the sense-club on the exumbrella side, but in Aurellia solida the “hood” is a

6258 MEDUS.Z OF THE WORLD.

mere ridge or ledge-like mass of tissue covering the concavity in which lies the sense-club.
The dorsal sensory-pit in Aurellra solida is a deep triangular funnel with a long, narrow mouth,
and is quite unlike the shallow exumbrella sensory-pit of Aurellza aurita.

The bell is 80 mm. wide and 40 mm. high. Canal-system similar to that of Aurellia
aurita, but the 4 circular, subgenital ostia are each only 2 mm. in diameter, whereas in 4. aurita
they are usually much larger. The oral arms are not quite as long as the radius of the
umbrella. The genital radius is about 0.4 that of the umbrella.

Maldive Islands, Indian Ocean, and 15° west of Madeira in the North Atlantic.

Aurellia labiata Chamisso and Eysenhardt.

Aurelia labiata, CHAmisso UND Eysennarpt, 1820, Nova Acta phys. med. Leop. Car., tome 10, p. 358, planche 28, figs. 1 A, B.

Aurellia labiata, pe Brainvitre, 1834, Manuel d’Actinologie, p. 294, planche 42, figs. 1, 2—Fewxes, 1889, American
Naturalist., vol. 23, p. 592, fig. 2; Bull. Essex. Institute, Salem, vol. 21, No. 7, p. 122, plate 5, fig. 2.

Diplocraspedon limbata, Branvr, 1838, Mém. Acad. Sci. St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 372, taf. 10.

Aurelia clausa (young medusa), Lesson, 1829, Voyage de la Coguille, Zool., p. 119.

Aurelia labiata+ A. clausa+ A. limbata, HaEcKEL, 1880, Syst. der Medusen, pp. 557, 558-

Aurelia limbata, Grrr, 1886, Sitzungsber. Akad. Wissen. Berlin, Jahrg. 1886, p. 836.—VaNnuOFFEN, 1906, Nordisches Plankton,
Nr. 11, p. 61, fig. 32.—Kisuinovuye, 1910, Journal College of Sci. Tokyo, vol. 27, art. 9, p. 22.

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Fic. 398.—Aurellia labrata. Drawn by the author, from specimens collected by the U. S. Bureau of

Fisheries steamer Albatross at Masbate Anchorage, Philippine Islands, April 21, 1908.

A, oral view of medusa with one mouth-arm cut off. B, bell-margin seen from exumbrella side showing
dorsalward migration of lappets and tentacles. C, section of bell-margin, the areas cut across
being dotted. c, circular vessel; ex, exumbrella; /, marginal lappet; rc, radial-canal; su, sub-
umbrella; v, velum-like bell-margin of bell; #, tentacle.

This species is distinguished by having 16 velar lobes separated by deep median clefts
instead of 8 simple lobes as in 4. aurita. The canal-system is similar to that of Aurellia aurita,
but there is probably a greater tendency for fusions to occur between the adradial and other
canals than in 4. aurita. The bell-margin projects downward from the subumbrella side as

SEMAEOSTOMEA[—AURELLIA, 629

8 plain-edged, velum-like folds spanning between the sense-organs. The tentacles and mar-
ginal lappets have migrated a considerable distance up the sides of the exumbrella, above the
velar margin. A longitudinal strand of muscle-fibers extends down the subumbrella side of
each tentacle and interrupts the rings of nematocysts which trend across its exumbrella
side. When the medusa is old the mouth-arms become much thickened and folded as in
Aurellia aurtta.

The dimensions of three specimens obtained by the U. S. Fisheries Bureau steamer
Albatross at Masbate Anchorage, Philippine Islands, on April 21, 1908, are as follows:

| |
Mm.| Mm.) Mm.
|

Diameter of umbrella. ................ 174 | 189 | 128
Diameter across zone of gonads........ | 57 53 | 42

Length of each mouth-arm............ 74 75 i) 52
|

Aurellia labiata is distinguished from 4. aurita by having 16 notches in its bell-margin,
by its peculiar velum-like, inter-rhopolar, subumbrella membranes representing the true bell-
margin, and by the very small size of its subgenital ostia. The mouth-arms are also shorter
than one commonly observes them to be in 4. aurita.

Aurellia maldivensis H. B. Bigelow.
Aurelia maldtvensis, Bicetow, H.B., 1904, Bull. Mus. Comp. Zool. at Harvard College, vol. 39, p. 261, plates 6, 8, figs. 22, 23,27.

Bell 250 mm. wide and about one-third as high. 8 marginal sense-organs, flanked by
small, pointed, ocular lappets. 8 wide interocular or velar lobes are each divided into 2 by

Fic. 399.—Aurellia maldivensis, after H. B. Bigelow, in Bull. Mus. Comp. Zool. at
Harvard College.

a very slight, central depression, thus forming 16 lobes as in 4. labiata. About 500 small
tentacles alternate with an equal number of small, dorsal lappets asin durellia aurita. The
4 mouth-arms, or palps, are large and curtain-like, recalling those of Cyanea; their lips are
complexly folded and bear numerous, short tentacles. About 48 radial-canals arise from
the central stomach, the 8 canals to the marginal sense-organs and the 8 adradial ones do
not branch, but all of the others branch, and occasionally anastomose, so that about 175
canals reach the circular vessel at the margin. The 8 canals to the sense-organs each give
off 2 side-branches in the immediate neighborhood of the sense-organ. These side branches
extend to the circular vessel. The 4 gonads are small and horseshoe-shaped and have
wide subgenital pits.

The bell is of a delicate lilac, the canals and tentacles pinkish-violet, and the mature
gonads bright violet. The color is, however, variable, some specimens being blue.

Abundant in the lagoons of the atolls of the Maldives, Indian Ocean, in January.

630 MEDUS‘® OF THE WORLD.

The comparative dimensions and other details are stated in the table giving a synopsis
of the species of Aurellra.

In its cleft, velar lobes it recalls 4. /abiata, while in its wide, curtain-like mouth-arms it
stands alone among Aurellias. Indeed its peculiar mouth-curtains are all that separate it from

A. labiata.
Genus AUROSA Haeckel, 1880.

Aurosa, Haecket, 1880, Syst. der Medusen, p. 559.

The type species and only known form is durosa furcata Haeckel, from the tropical
Indian Ocean.
GENERIC CHARACTERS.

The medusa is similar to 4urellia but the 4 mouth-arms are bifurcated at their outer
ends. ‘There are 24 anastomosing radial-canals and a marginal ring-canal. 4 interradial
gonads, 8 rhopalia, 8 velar lobes, and numerous small tentacles as in Aurellia.

Aurosa furcata Haeckel.

Aurosa furcata, Haecket, 1880, Syst. der Medusen, p. 559, taf. 33, fign. 7, 8.

Bell flat, shield-shaped, 80 mm. wide, 30 mm. high, 8 velar lappets as in Aurellia aurita.
8 marginal sense-organs. Tentacles small, numerous, arising from exumbrella side of margin

Fic. 400.—Aurosa furcata, after Haeckel, in Das Syst. der Medusen.

asin 4. aurita. Gonads asin 4. aurita. Genital radius half the bell-radius. Central stomach
gives rise to 24 radial-canals, all of which give forth anastomosing side-branches. There
are 4 perradial, 4 interradial, and 16 adradial canals. Thus each genital pouch gives rise to
5 radial-canals, and the perradial canals arise from the angles between the genital pouches.
The peripheral network of anastomosing canals becomes narrower in its mesh the nearer
the vessels are to the marginal ring-canal. The 4 mouth-arms resemble those of Aurellia
aurita, but each one bifurcates near its outer end. Their curtain-like margins are much
folded.

Found near Cocos Islands, Indian Ocean, southwest of Sumatra.

RHIZOSTOM. 631

Order RHIZOSTOM Cuvier, 1799.

Rhizostoma, Cuvier, 1799, Journal de Phys., tome 49, p. 436.—Harcket, 1880, Syst. der Medusen, p. 560.—HAMANN, 1881,
Jena. Zeit. fiir Naturwissen., Bd. 15, p. 243.—Craus, 1883, Untersuch. iiber Organisation und Entwick. der Medusen,
Leipzig, p. 60; 1886, Arbeit. Zool. Inst. Wien, Bd. 7, p. 110.—von LenpeNreELp, 1888, Zeit. fiir wissen. Zool., Bd. 47,
p- 208.

Rhizostomee, AGassiz , L., 1862, p. 208, Cont. Nat. Hist. U. S., vol. 4, p. 149.

Rhizostomata, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 39-—Maas, 1903, Scyphomedusen der Siboga Exped.,
Monog. 11, p. 88; 1906, Revue Suisse de Zool., tome 14, p. 100; 1907, Ergeb. und Fortschritte der Zoologie, Bd. 1, p. 201.

Rhizostomide, EscuscuHourz, 1829, Syst. der Acalephen, p. 42.

CHARACTERS OF THE ORDER.

Scyphomedusz without marginal tentacles*, and with numerous mouths which are borne
upon § adradial, fleshy, Heanched arm-like appendages which arise from the center of the
subumbrella. The lips of the numerous mouths are bordered by minute, constantly moving
tentacles.

The rhopalia and marginal lappets of the Rhizostomz are similar to those of Semaos-
tomee. The Rhizostomz are the most highly differentiated Scyphomedusz, and owing to
the generally tough consistency of their gelatinous substance and their large size they bee
often been found preserved as fossils, especially i in the lithographic slates OF Solenhofen and
Fischstadt. They are tropical forms and none are known from the polar regions. The
genus Rhizostoma is the only one which extends far into temperate regions, and the majority
of the genera are confined to the warm waters of the Indo-Pacific region. A few are found
in the tropical Atlantic and the Mediterranean. The Rhizostomz develop, in so far as we
know, through strobilization from scyphostomz. Phylogenetically they are derived from the
more Sale organized Semzostomez, but they have lost their marginal tentacles, though in
Lobonema the marginal lappets have become greatly elongated and may in some respects finchion
astentacles. The ephyra of the Rhizostomz has a simple cruciform, central mouth, as in the
Semzostomee, but the 4 rays of the cross soon fork at their outer ends and then grow outward
in the form of 8 adradial, fleshy, mouth-bearing appendages, which branch in a characteristic
manner and constitute the so-called ‘‘mouth-arms.”” The primitive central mouth may then be
obliterated by the coalescence of its lips, but numerous other mouth-openings remain in the
gutter-like grooves which extend down the ventral sides of the mouth-arms; and these mouths
may also extend over parts of the dorsal sides of the mouth-arms. The semzostomous genus
Aurosa with its 4 bifurcated mouth-arms, which in other respects resemble those of 4urellia,
appears to be a connecting link between the Semaostomez and the Rhizostome.
The gonads of the Rhizostome are inyaginated as are those of Aurellia, not protrusive
as in Cyanea. In the young meduse of all Rhizostoma and in the mature medusz of some
genera such as Cassiopea there are 4 sepa-
rate genital sacs which project into the
central stomach-cavity of the medusa. In
- many Rhizostomz such as in C otylorhiza,
C Mastigtas, etc., the inner ends of the 4
primitive, genital bags may coalesce and
their walls break down, forming a cruci-
A B C form cavity beneath the stomach of the
Fic. 401 -—Diagrams illustrating the fusion of the 4 primitive medusa and not connected with the gas-
genital cavities (a) to form a single cruciform cavity (c). : :
trovascular cavity, but opening to the
surrounding ocean through the 4 interradial genital ostia. The formation of this cruciform,
genital cavity, or subgenital porticus of Haeckel, is illustrated in the 3 diagrams of figure 401.
In A, we see 4 separate, interradial, genital sacs (G) projecting into ane Sromeche -cayity (S),
asin (Rhizestonta and Cassiopea. In B we see these 4 sacs fused at their inner ends as 1n some
species of Cephea (Netrostoma); and in C we find the walls broken down in the fused regions
forming a cruciform genital space lying beneath the stomach as in. ' Cotylorhiza, C ‘aie ssa,
Mastigias, etc.
The umbrella of the Rhizostome resembles that of their more simply organized ancestors
the Semzostomez. It is usually dome- shaped and covered with nematocyst-w arts. The bell-

*Excepting in Lobonema, gen. nov., wherein the marginal lobes are converted into long, tapering tentacle-like organs.

632 MEDUS.2 OF THE WORLD.
|appets and rhopalia are in all respects similar to those of the Semaostomez. The muscular
system of the subumbrella is well-developed and these forms are usually vigorous swimmers,
although in Cassiopea we find that the meduse# commonly remain upon the bottom with their
oral sides uppermost, and the pulsations of the umbrella serve mainly to stir up currents which
may bring food to the mouths.

~ At the center of the subumbrella we find a thick, disk-shaped, gelatinous projection called
the arm-disk, for the 8 adradial mouth-arms arise from its lower side. This arm-disk is
merely the lower wall of the stomach which has become thickened in order to give support
to the heavy gelatinous mouth-arms. In all forms, however, having a unitary, cruciform,
genital cavity, an open space lies between the arm-disk and the stomach so that the arm-
disk is suspended from the subumbrella by 4 thick perradial columns which are separated
one from another by the 4-rayed genital porticus, which opens to the outer world by 4 inter-
radial ostia which alternate with the columns.

The cruciform, central stomach dips downward into these perradial columns and 4
bifurcated or 8 simple canals arise from the stomach and extend downward into the 8
adradial mouth-arms, giving off numerous branches to the mouths. The 16 canals to the
scapulets, when these are present, arise from these 8 mouth-arm ducts, as do also the canals
to the arm-disk, which fuse into 4 and finally into a single, central duct at center of arm-disk.
The central stomach also gives rise to canals which radiate outward through the subum-
brella of the bell. These may be connected by one or more ring-canals, or by networks of
anastomosing vessels.

The facility with which some of these medusa may be maintained alive in aquaria has
permitted certain physiological work to be performed upon them. Bethe, 1903, 08, 09, studied
the rhythmical pulsation of Cotylorhiza and Rhizostoma, and Mayer, 1906, 08, carried out
experiments upon Cassiopea. Bethe finds that the pulsation resembles that of the verte-
brate heart in all important respects. The pulsation-stimulus is nervous in nature, and the
“all or none” principle applies to medusz as it does to the vertebrate heart, as does also the
phenomenon of the refractory stage of Marey, 1876. A definite interval of time elapses between
the passage of the nervous stimulus and the response of the muscles, and the pulsation is a
reflex due to a constantly present stimulus, the response to which is periodic, because after
the nerves have responded to the stimulus they become incapable of reacting to it until after a
definite interval of rest, this resting period being called the refractory stage.

Bethe, 1908, 09, in his study of Rhizostoma pulmo comes to conclusions in respect to the
effects of the ions of sea-water upon pulsation, which are in accord with those of Mayer, 1906.
(See Rhizostoma pulmo.)

Mayer, 1906, 1908, working upon Cassiopea, found that the sea-water is a balanced fluid,
neither stimulating nor inhibiting pulsation. This is due to the fact that the stimulating
effect of the sodium ion of sea-water is counterbalanced by the inhibiting influences of the
calcium, potassium, and magnesium. The stimulus which produces pulsation is due to the
constant maintenance of a slight excess of the sodium cation in the marginal sense-clubs, over
and above its concentration in the surrounding sea-water. This excess of sodium is main-
tained by the constant production of sodium oxalate in the terminal entoderm of the sense-
clubs. This oxalate precipitates calcium to form the calcic oxalate crystals of the sense-club
and sets free sodium chloride the sodium ion of which acts as a nervous stimulant. Details
of these researches upon pulsation are given in the accounts of Cotylorhiza tuberculata,
Rhizostoma pulmo, and Casstopea xamachana. j

Hargitt, Zeleny, and Stockard have studied regeneration in Rhizostome. Zeleny stated that
in Cassiopea the greater the number of arms removed up to 6 the more rapidly does each and
every arm regenerate, but this is refuted by Stockard, who further shows that the regenerating
tissue has a greater ability to absorb nutriment than have the normal, somatic body tissues,
and that in consequence of this the body shrinks in size in direct proportion to the growth of
the regenerating arms, the growing arms reducing the body as do cancer cells in their prolifer-
ation. Stockard also shows that cuts near the center regenerate more rapidly than those near
the margin of the disk, this being in accord with Morgan’s law that the deeper the level of the
cut the more rapid the rate of regeneration. In Rhizostoma pulmo Hargitt found that two
rhopalia sometimes regenerate in the place of one which he had removed, and I have observed

RHIZOSTOM.B. 633

the same thing in Casstopea. In Cassiopea xamachana R. P. Bigelow showed that the rhopalia
are derived from every alternate tentacle of the scyphostoma, the other tentacles degenerating
wholly. I find that inthis medusa when the rhopalium regenerates it gives rise to a short lateral
branch thus tending in an abortive manner to regenerate che contadle. from which it originally
came.

Many observations have been carried out upon the embryology of Rhizostome, and
reviews of these researches will be found in the descriptions of Casstopea xamachana, Cotyl-
orhiza tuberculata, Rhizostoma pulmo, Masti gas papua, Ph yllor hiza punctata, and Stomo-
lophus meleagris. Claus, Goette, R. P. Bigelow, Kowaleysky, von Lendenfeld, and Vanhoffen
have been especially active in these perches

Haeckel, 1880, considered the presence or absence of a unitary, cruciform, genital cavity
to be of great systematic importance and sought to separate families upon this distinction; but
Claus, von Lendenfield, Vanhoffen, Maas, and Browne have demonstrated that this is a matter
of no great import, for in different individuals of the same species we may find in some cases 4
separate genital sacs, while others have a cruciform genital cavity, and still others may have
a more or less complete coalescence and breaking apart of the partitions in some quadrants
and not in others. It is therefore evident, as was first clearly shown by Claus, 1883 (Organ-
isation und Entwick. Medusen), that the conditions exhibited by the genital sacs afford no
criteria for the distinction even of genera, much less of families. Indeed, Haeckel’s system
leads to the separation of closely related forms and the close approximation of remotely
related forms, and is quite artificial.

Claus, 1883, 1886, and Vanhoffen, 1888, have attempted to separate the families of
Rhizostome upon the distinctions afforded by the manner of branching of their mouth-arms.
Claus’s system somewhat modified by von Lendenfeld, 1888 (Zeit. fiir wissen. Zool., Bd. 47,
p- 208), distinguished nine families as follows:

Ruizostom#£: Scyphomeduse without marginal tentacles and with 8 adradial mouth-arms.

Archirihizide: Mouth-arms unbranched. Gastrovascular network simple. No central mouth.

Cassiopeide: Arm-disk flat. Arms long, irregularly branched with appendages. Radial-canals numerous. No central
mouth,

Craunostomide: Arm-disk wide, style-shaped, arms dichotomously forked. Arm-margins free, with clubs. Centripetal
canals end blindly. Subgenital porticus unitary. Central mouth-opening present.

Cepheide: Arm-disk wide and flat. Arms dichotomously forked, with 2 of the axial, terminal wings turned outwards.
With clubs. No direct central mouth-opening.

Lychnorhiztde: Arms 3-leaved or distally 3-winged. 8 or 16 radial-canals. Gastrovascular network simple. No central
mouth.

Stomolophide: Arm-disk style-shaped, elongate, with 8 pairs of lateral “shoulder ruffles” or “scapulets.” Proximal
parts of the arms fused into a tube, distal parts branched. 16 radial-canals, with well-developed net-work of con-
necting vessels. No central mouth.

Rhizostomide: Arm-disk style-shaped, elongate, with 8 pairs of lateral “scapulets” with clubs. Lower-arm three-
winged, with dorsal mouths. 16 radial-canals. Centripetal network of canals well-developed. No central mouth.

Catostylide: Arm-disk very wide, elongate, and style-shaped. Lower arm 3-winged with dorsal mouths. No centrip-
etal networks of canals. Subgenital porticus unitary. No central mouth.

Leptobrachide: Arm-disk wide and fused with the upper arms. Lower arms long, ribbon-shaped, and 3-winged.
Simple canal net spread over the entire subumbrella. Subgenital porticus unitary. No central mouth.

A simpler system is proposed by Vanhéffen, 1888 (Bibliotheca Zoologica, Heft 3), who
divides the Rhizostome into 7 families:

Rhizostomata simplicia: Mouth-arms simple and unbranched. All of these are apocryphal, having been seen only
by Haeckel and Fewkes.

Dichotoma: Mouth-arms dichotomously forked, with lateral expansions.

Pinnata: Elongate mouth-arms pinnately or irregularly branched.

Triptera: Mouth-arms 3-winged. Each mouth-arm with a ventral and 2 dorsal lamella which meet at a point at the
lower end of the arm.

Trigona: Identical with the Rhizostomata triptera.

Lorifera: Mouth-arms elongate, lash-like, and triangular in cross-section; with mouths developed along the 3 angles
of the arms.

Scapulata: Mouth-arms with simitar-shaped “scapulets” or “ruffles” projecting from their dorsal sides.

As was pointed out by Maas, 1903, Vanhoffen’s Triptera and Trigona are identical
and should be united, thus reducing his families to six. Schultze, 1808, showed that the
mouth-arms of the ‘‘Dichotoma” of Vanhoffen are not forked at their outer ends, but give
rise to 2 broad, longitudinal, Jateral lamella, which may branch secondarily. With these
modifications Vanhoffen’s system affords the readiest means of classifying the Rhizostome,

being based upon the mutations of the most conspicuous organs, the mouth-arms.

634 MEDUSZ OF THE WORLD.

Maas, 1903 (Syphomedusen der Siboga Expedition, p. 89), proposes another system
based upon the character of the muscle-system of the subumbrella, the presence or absence
of ocelli on the sense-clubs, the character of the canal-system and of the mouth-arms. Maas’s
system is as follows:

Arcadomyaria: The subumbrella muscles are arranged in feather-like arcs. Mouth-armselongate and irregularly pinnate
in their branching. Rhopalia with ocellus and without an exumbrella sensory pit. Radial-canals twice as numerous
as the rhopalia and connected by an anastomosing network of vessels. One or more ring-canals may or may not be
present. 4 separate genital sacs, with small, round, interradial ostia. There is only one family, the Cassiopeide.

Radiomyaria: Radial-muscles of the subumbrella better developed than the circular muscles. Mouth-arms bifurcated.
Rhopalia without ocelli and without exumbrella, sensory pits. 8 principal and other secondary radial-canals, all
connected by a marginal network. No definite ring-canal. Funnel-shaped genital ostia. A unitary subgenital
cavity may or may not be present. There is one family, the Cephetde.

Cyclomyaria: Circular muscles of the subumbrella better developed than the radial-muscles, the latter being often absent.
Mouth-arms 3-winged, or derived from this type. There are 3 groups of the Cyclomyaria, as follows:

(4) 16 radial-canals which extend from the stomach to the bell-margin, and between them a blindly-ending, anastomosing
network of vessels. Mouth-arms with scapulets. Genital ostia slit-like and divided by a median flap. Rhopalia
without ocelli, but with sensory pits with radiating furrows. Group 4 is equivalent to the Rhizostomide+ Stomo-
lophide of Claus, or to the Rhizostomata scapulata of Vanhoffen.

(B) 8 rhopalar canals extend to the bell-margin and 8 in the inter-rhopalar radii end in the ring-canal. On its outer
side, the ring-canal gives off a network of anastomosing vessels, and on its inner side it gives rise to another network
which ends blindly without connecting with the stomach.

(1) Mouth-arms 3-winged, usually with pinnate lateral branches. Genital ostia slitlike. Rhopalia with pigment
spots, and sensory pits with radiating furrows. This contains the family Lychnorhizide of Claus; including
the genera Lychnorhiza, Crambione, and Crambessa.

(2) Mouth-arms very elongate, triangular in cross-section. Genital ostia wide openings. Rhopalia with pigment
spots and furrowed sensory pits. This contains a part of Claus’s family Leptobrachide.

(C) The 8 rhopalar extend to the bell-margin, and (8 X) canals extend only to the ring-canal. On its outer side the
ring-canal gives off a narrow network and on its inner side is a network of wider mesh.

(1) Mouth-arms very elongate, triangular, file-shaped. Genital ostia wide slits. Genera: Thysanostoma, Lepto-
brachia, and Himanostoma of Claus’s Leptobrachide.

(2) Arms 3-winged, not elongate. Ostia wide. Rhopalia with ocelli and small sensory pits without furrows.
This is equivalent to Claus’ family Catostylide containing the genera Loborhiza, Crossostoma, and Mastigias.

The Arcadomyarta of Maas is only a new name for Vanhéffen’s Rhizostomata pinnata,
the Radiomyaria are equivalent to Vanhéffen’s Rhizostomata dichotoma, and the Cyclomyaria
includes rather confusedly the triptera+trigona+ scapulata+lorifera of Vanhoffen. Maas’s
system is erroneous in some respects; for example the rhopalia of Cassiopea xamachana have
pigment spots while those of Casstopea frondosa have none; similarly Crambessa tagi has
(“ocelli”) pigment spots but Crambessa mosaica has none. It is therefore evident that the
presence or absence of “‘ocelli” does not afford a suitable criterion for the separation even
of genera. The exumbrella sensory pits may have furrows in one species of a genus and
be simple in another, as in Rhopilema esculenta and R. verillit. In my opinion the older and
simpler system of Vanhéffen is to be preferred to this complex scheme proposed by Maas.

In view of the observations of Schultze and of Maas, we may amend Vanhiffen’s system
as follows: ;

Rhizostomata pinnata: Rhizostome with 8 separate, elongate, linear mouth-arms which give rise to pinnately or com-
plexly arranged side branches (figs. 4, 5, and 7, plate 69). The circular muscles of the subumbrella tend to be
bowed outward in a series of arcs on both sides of each radial-canal, the conyexities alternating with the canals. The
genera are as follows: Toreuma Harcxet, 1880, with 8 rhopalia.

Cassiopea PéRon AND Lesurur, 1809, with more than 8 rhopalia.

Rhizostomata dichotoma: 8 separate mouth-arms, the lower parts of each one of which is V-shaped in cross-section, the
apex of the V being centrad, and the rays directed outward (fig. 404, p. 650) The mouths are developed upon the
ventral sides of the mouth-arms. The radial-muscles are powerfully, and the ring-muscles weakly, developed. The
radial-canals are all connected by a marginal network of vessels, without a definite ring-canal. The genera are as
follows:

Cephea Péxon anv Lesurur, 1809. Exumbrella with a central dome bearing solid, wart-shaped protuberances.

Cotylorhiza L. AGassiz, 1862. With a smooth, simple dome at the center of the exumbrella.

Polyrhiza L. Acassiz, 1862. Exumbrella with a central depression and with radiating furrows.

Rhizostomata triptera: 8 separate mouth-arms the lower parts of each of which is Y-shaped in cross-section, due to the
development of 2 lateral, dorsal, and a median ventral, longitudinal lamella, all 3 of which taper to a point at the
lower end of the arm. The mouths are developed upon the free edges and partially over the sides of the 3 lamella
or wings of the mouth-arms (fig. 411, p. 664) The ring-muscles are powerfully and the radial-muscles weakly, devel-
oped. A ring-canal with a network of anastomosing vessels arising from its inner and outer sides is usually present.
Mouth arms without scapulets. The genera are very closely related and are as follows:

Catostylus L. AGassiz, 1862= T oxocl ytus + Crambessa Haecker. Mouth-arms without filaments, clubs, or other
appendages. The network of vessels arising from the inner side of the ring-canal ends blindly, without con-
necting with the stomach.

Lychnorhiza Harcxer, 1880. With filaments but without clubs upon the mouth-arms. In other respects similar
to Crambessa.

RHIZOSTOM4—TOREUMA. 635

Rhizostomata triptera, continued—

Crambione Maas, 1903. Similar to Crambessa, but with both clubs and filaments upon the mouth-arms.

Mastigias L. Acassiz, 1862= Mastigias+ Eucrambessa Harcket. Each mouth-arm terminates in a naked club.
Numerous clubs or filaments among the mouths. The network of vessels which arises from the inner side
of the ring-canal connects with the stomach.

Pseudorhiza von LenpENFELD, 1882. Similar to Mastigias but without lateral clubs or filaments upon the
mouth-arms. A terminal club present. The canals which arise from the circular vessel, between the radial-
canals end blindly without reaching the stomach.

Phyllorhiza L. Acassiz, 1862. Mouth-arms with lateral filaments, but without clubs as in Lychnorhiza. Canal-

system as in Mastigias.

Vessura, Harcxet, 1880. Mouth-arms with clubs and filaments as in Crambione. 4 perradial canals arise
directly from the stomach, but the 4 interradial canals result from the fusion of a network of vessels which
arise from each interradial side of the stomach. An outer and an inner zone of circular muscles with an
annular separation between them.

Lobonema, gen. nov. Marginal lappets elongated to form tentacles-like organs. Mouth-arm membranes per-
orated by window-like openings. Exumbrella covered profusely with papille.

Rhizostomata lorifera: 8 mouth-arms, very elongate, whip-like, and triangular in cross-section, with frilled mouths
developed along the angular edges of the arms (fig. 419, p.691) The cylindrical upper parts of the arms are rudimen-
tary and partially fused one to another by gelatinous arches spanning between them and connecting them with the
arm-disk. The genera are as follows:

Thysanostoma L. AGassiz, 1862. No terminal clubs on the mouth-arms. Mouths along the 3 angles through-
out length of arms.

Lorifera, HAecKet, 1880. Similar to Thysanostoma, but with a naked club at lower end of each arm.

Leptobrachta Branpt, 1838=Leptobrachia+ Leonura Harcxet. Mouths confined to the upper and lower ends
of the mouth-arms, leaving the middle part of the arm naked.

Rhizostomata scapulata: Each mouth-arm bears a pair of simitar-shaped appendages (scapulets) which arise from the
outer side near the base of the arm and bear frilled mouths (fig.421, p.697). The circular muscles of the subumbrella
are powerfully, and the radial-muscles weakly, developed, or even absent. The genera are as follows:

Rhizostoma Cuvier, 1800=Pilema Harcket, 1880. 8 free mouth-arms, the lower parts of which are Y-shaped,
or 3-winged, in cross-section. Each arm terminates in a naked club. There are no other clubs or filaments.

Rhopslema Harcxer, 1880. Similar to Rhizostoma but with numerous clubs or filaments upon the mouth-arms.

Euptlema Hartcket, 1880. Similar to Rhizostoma, but the arms have neither clubs nor filaments.

Stomolophus L. Acassiz, 1862 = Brachiolophus+ Stomolophus, Harcket, 1880. The 8 mouth-arms are fused
along their sides leaving only the lower ends free and forming an elongate throat-tube for the central mouth
which remains open. :

Rhizostomata simplicia: Rhizostome with unbranched mouth-arms. These apocryphal forms are described by Haeckel
and by Fewkes from alcoholic specimens of small size. They are apparently immature or injured specimens. No
naturalist has seen any of these forms since Fewkes described his “Stomotonema reticulatum” in 1884. It is probable
that we should drop these meduse from further consideration, but in the faint hope that some may be discovered
I have given descriptions of them based upon the statements of Haeckel and Fewkes.

RHIZOSTOMATA PINNATA Vanhiffen.

Rhizostomata pinnata, VANHGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 40.—Maas, 1903, Scyphomedusen der Siboga
Exped., Monog. 11, p. 38.
Cassiopetde, Craus, 1883, Organisation und Entwick, Medusen, Leipzig—von LenpENFELD, 1888, Zeit. fiir wissen. Zool., Bd.

47, P- 211.
Arcadomyaria, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 88; 1907, Ergeb. Fortschritte der Zool., Bd. 1,
p- 201; 1906, Revue Suisse de Zool., tome 14, p. 100.

Rhizostomous medusz with 8 linear, pinnately, or complexly branching mouth-arms.

GENERA.
Toreuma Haeckel, 1880. 8 rhopalia.
Casstopea Péron and Lesueur, 1809. More than 8 rhopalia.
Genus TOREUMA Haeckel, 1880.

Toreuma, HarcxeEL, 1880, Syst. der Medusen, p. 566.—VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 40.—Maas,
1903, Scyphomedusen, Siboga Exped., Monog. 11, p. 43-

The type species is Toreuma dieuphila, described by Péron and Lesueur from the Indian
Ocean.

GENERIC CHARACTERS.

Rhizostomata pinnata with 8 adradial, linear mouth-arms which branch pinnately or
complexly, and the main side branches also branch. 8 marginal sense-organs.

This genus is closely related to Casstopea and is distinguished only by having 8 rhopalia,
whereas Casstopea has more than 8. Haeckel is the only modern naturalist who has seen
any of these forms. They all come from the Indian Ocean.

636

MEDUS OF THE

WORLD.

Toreuma dieuphila.

Cassiopea dieuphila, Pinon et Lesueur, 1809, Annal du Mus. Hist. Nat. Paris, tome 14, p. 356.
Cassiopea theophila, pe Lamarck, 1816, Hist. Nat. Anim. sans Vert., tome 2, p. 511.
Rhizostoma theophila, Escuscnoutz, 1829, Syst. der Acalephen, p. 53.

Polyclomia theophila, Aassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 159.
Toreuma theophila+ T. thamnostoma+T. gegenbauri, HaeckeL, 1880, Syst. der Medusen, pp. 566, 567, 645.

It is probable that Haeckel’s T. “‘thamnostoma”
stages of Péron and Lesueur’s “

Casstopea”’

dieuphila.

and “TJ. gegenbauri”
I therefore present the descriptions

are only growth-

of the three forms side by side in order that they may readily be compared. Haeckel enjoyed
the opportunity of studying Péron and Lesueur’s original specimen preserved in Paris.

Toreuma dieuphila=Cassiopea
dieuphila Péron and Lesueur.

Toreuma “gegenbauri” Haeckel.

Toreuma “thamnostoma” Haeckel,

Diameter of bell in
mm.

Shape of bell.

Number of marginal
lappets.

Length of mouth-
arms in terms of
bell-radius (r).

Number of branches
of each mouth-arm.

Appendages upon
mouth-arms, be-
tween mouths.

Color.

Where found.

60 to 80

Hemispherical (contracted ?)
Exumbrella with coarse warts.

96. In each octant 10 short,
rectangular, velar, and 2 very
small, ocular lappets.

Less than r long (contracted ?)

6 to 8 wide, flat, main side-
branches.

Numerous small, and 10 to 20
large, club-shaped vesicles.

Bell brownish-red, with white
spots on the lappets. Gonads
and clubs white.

Northwest coast of Australia,
in the Indian Ocean.

60

Flatter than a hemisphere, without
papillx.

80. In each octant 8 short, rec-
tangular, velar, and 2 very small,
ocular lappets.

1too.5rlong.

8 to 12 flattened main side-
branches.

Numerous small clubs and a very
large one at base of each arm,
half as long as arm itself.

Bell brown (?) An elongated,
white spot upon lappet.

Tropical Indian Ocean.

go

Flatter than a hemisphere. Exum-

brella with small warts.

120 to 160. In each octant 14 to 18
short, rectangular, velar, and 2 very
small ocular lappets.

Nearly 2 r long.

12 to 16 cylindrical, main side-
branches.

Numerous club-shaped vesicles.
Smoother than in T. dieuphila,
and not longer than width of mar-
ginal lappets. (Large clubs lost?

Bell dark-brown with numerous
white spots. Abaxial surface of
arms yellowish-brown. Clubs
white.

Indian Ocean.

Genus CASSIOPEA Péron and Lesueur, 1809.

Cassiopea, Peron et Lesurur, 1809, Ann. du Mus. Hist. Nat. Paris, tome 14, genre 24, p. 356.—Escuscuovtz, 1829, Syst.
der Acal., p. 42.—Tirersius, 1834, Acad. Caes. Leop. Nova. Acta., tom. 15, p. 256.—Branpr, 1838, Mém. Acad. Impériale
des Sci., St. Pétersbourg, Sci. Nat., sér. 6, tome 4, p. 396.—Gurpy, 1883, Nature, vol. 27, p. 31 (habits).—von LenpENFELD,
1884, Proc. Linnean Soc. New South Wales, vol. 9, p.284.—Ctaus, 1883, Organisation und Entwick. der Medusen, p. 60.—
Corasanti, 1886, Atti Acad. Med. Roma, Anno. 12, 18 pp. (blue color). —Acassiz AND Mayer, 1899, Bull. Mus. Comp.
Zool. at Harvard College, vol. 32, p. 175.—Keter, C., 1883, Zeit. fiir wissen. Zool., Bd. 38, p. 632.—Browne, 1905, Fauna
and Geog. Maldive and Laccadive Archipelagoes, vol. 2, p. 966.—Maas, 1903, Scyphomedusen der Siboga Expedition,
Monog. 11, pp. 38, 80.—Scuuttze, L. S., 1898, Abhandl. Senckenberg. Gesell., Bd. 24, Heft. 2, p. 163.

Catsiopeat polyclania, Agassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, pp. 139, 155, 159-—Perkins, 1908, Papers from Tortugas
Laboratory of Carnegie Inst. of Washington, Publication 102, p. 150.

Polyclonia+ Cassiopea, Harcket, 1880, Syst. der Medusen, pp. 567, 568.

The type species is Medusa andromeda Forskal, called C. forskalca by Péron and Lesueur

(=C.

andromeda,

Eschscholtz) of the Red Sea and Indian Ocean.

cribed by Péron and Lesueur, 1809, is called Cassiopea dieuphila.
places this in the genus Toreuma.

GENERIC CHARACTERS.

The first species des-
Haeckel, 1880, however,

Rhizostomata pinnata with 8 (4 pairs of) adradial, complexly branched mouth-arms

the lower or ventral surfaces of which bear numerous mouth-openings and vesicles.
are 4 gonads and 4 separate subgenital cavities.
organs and twice as many radial-canals as sense-organs.
communication one with another by means of an anastomosing network of vessels.

ring-canal may or may not be present.

There are mo

There

re than 8 marginal sense-

The radial-canals are placed in

A definite

RHIZOSTOM®—CASSIOPEA, 637

Tilesius, 1834, figured 4 species of Casstopea and represented each of them as having
8 subgenital cavities. Relying upon the figures of Tilesius, L. Agassiz, 1862, separated the
genus Polyclonia, having but 4 subgenital cavities. Later pee ehes have demonstrated
that all the known species of these meduse have normally but 4 subgenital cavities, and
should therefore be placed in the genus Cassiopea. Haeckel, 1880, attempts to separate
Casstopea from Polyclonia by calling meduse with 16 marginal sense-organs Cassiopea,
while those with 12 of these organs are called Polyclonia. The number of marginal sense-
organs is, however, very variable, not only among different species of these meduse but also
among individuals of the same species, and therefore can not be used as a means of establish-
ing generic distinctions.

The medusz of this genus are all inhabitants of warm oceans, and are found in greatest
abundance in the tropical coral regions of the East Indies and Red Sea. R. P. Bigelow finds
that C. xamachana from the West Indies develops through the monodiscus strobilization
of a scyphostoma and the young ephyre of this species and of C. frondosa haye a simple,
central, 4-cornered mouth, thus recalling the adult condition in the Semaostomez, from which
forms the Rhizostomz Rave evidently een derived. The rhopalia of the ephyra are derived
from the bases of each alternate tentacle of the scyphostoma, the other tentacles degenerating.

The number of “species” of Cassiopea has been multiplied greatly, owing to the remark-
able color-range and variability in other respects of these medusa. These color types appear
to be local, ad the Cassiopea medusz of almost every new region of the tropics are nearly
certain to be described as “‘new species” based on color peculiarities. It is therefore impos-
sible, at present, to classify the forms of Casstopea with any degree of certainty.

The blue and amber-green coloration of these medusz is due to the presence of com-
mensal plant organisms. Colasanti, 1886, describes the blue pigment matter as zoocyanin,

Maas, 1903, attempts to separate the genus into two cohorts; one, consisting of C.mertenst,
C. mertensi var. ndrosta, polypoides, xamachana, ornata, and ornata vat. digitata, distinguished
by its long, cylindrical, pinnately branched mouth-arms. The other group consists of C.
andromeda and its varieties: C. depressa and C. depressa var. picta. This latter cohort has
irregularly branched, short, flat mouth-arms. An idea of the range in color-patterns of these
medusz may be obtained from an inspection of plates 70 to 72 which exhibit photographs of
a few of the varieties of Cassiopea xamachana, all taken in the course of an hour from the
moat of Fort Jefferson, Tortugas, Florida.

Stockard demonstrated that in C. xamachana the nearer the injury 1s made to the center
of the disk the more rapid the rate of regeneration. He also found that the more arms we
remove, the more does the central disk shrink during the growth of the regenerating arms, and
he thus finds that the regenerating tissue absorbs nutriment at the expense of the normal body
tissue as do cancerous tissues in their growth.

Mayer finds that the rhythmical pulsation in C. xamachana is due to a nervous stimulus,
and this stimulus is caused by the presence of a slight excess of sodium in the rhopalia over and
above the concentration of this ion in the surrounding sea-water. This excess of the sodium
ion is due to the constant formation of sodium oxalate in the sense-club, and this oxalate
precipitates the calcium chloride of the sea-water to form the calcium oxalate crystals of the
sense-club and sets free sodium chloride.

R. P. Bigelow finds that the vesicles between the mouths of C. frondosa serve to capture
prey and to thrust the food into the mouths.

Cassiopea andromeda Eschscholtz.

Medusa andromeda, ForskAt, 1775, Descript. que in Itinere Orientali Observavit, Haunia, p. 107, tab 31, 3 fign.

Cassiopea andromeda, Escuscuoutz, 1829, Syst. der Acalephen, p. 43.—Titestus, 1829, Nova Acta Phys. Men. N. C., tome 15,
p- 266, taf. 69, 70.—Mitnr-Epwarps, 1849, Cuvier’s Régne Animal Illustré, Zooph., plate 51, fig- 1—Harcker, 1880,
Syst. der Medusen, p. 569.—Ketter, 1888, Zool. Anzeiger, Bd. 11, pp. 359, 389-— —Hartlaub, 1909, Zoolog. Jahrbiicher,
Abth. Syst., Bd. 27, p. 467, taf. 23, fign. 1-8.

Cassiopea forskalca, PERon et Lesurur, 1809, Annal. du Mus. Hist. Nat., Paris, tome 14, p- 356.

Bell flat, shield-shaped, 100 to 120 mm. wide, 20 to 30 mm. high. 15 to 18, usually 16,
marginal sense-organs. A variable number of short, blunt lappets. In each paramere are I to
6, usually 3, velar flanked by 2 ocular lappets. 8 mouth-arms, wide, flat, and hardly as long as

—
Qo
1°.)

MEDUS OF THE WORLD.

Synopsis of the Described Forms of Casstopea.

C. andromeda. C. andromeda) C. andromeda | C. andromeda) C. andromeda | C. polypoides. | C. xamachana,
var. zanzi- var.malayen- var. maldiv- | var. acyclob-
barica. sis. ensis. lia.
Shape of bell. | Flat. Flat. Flat. Exumbrella With low cen- | Sucker-like Exumbrella
concave. tral dome. concavity on| concave.
exumbrella.
| Number of 12 to 18, usu- | 16 16 12 to 19, us- | 16 16 II to 23, us-
| rhopalia. ally 16. ually 16. ually about
16.

Number of §, occasionally) 8 3, 5,7,0Frg | 70 to 10 5 5 5
marginal lap- | 1 to 6.
pets in each
paramere.

Length of Less than r. | Less than. | Less than r. | Less than r. | 0.67 rt 1 to 0.257
mouth-arms
in terms of
bell-radius(r).

Vesicles and fil-| Many small; | Many small, | Many small, | Linear, hand-| 8 large, many | Large and Large and
aments among] 5 or more slargeclubs.| 2 to 3 large | shaped, and | small fila- small clubs small ribbon
mouths. large, club- Longer than | clubs. ribbon- ments. and filaments.) like filaments.

shaped vesi- | in C. andro- shaped ap-

cles. meda. pendages.
Some very
large.

Where found. | East coast of | Zanzibar Malay Archi- | Maldive Is- | Amboina, Coral flats of | West Indies to
Africa, Red coast, East pelago. lands, Indian) Malay Arch-| Red Sea. Florida.
Sea to the Africa. Ocean. ipelago.

Malay Arch-
ipelago.
C.frondosa. | C. ornata. C.ornata var.| C. depressa. | C. depressa | C. mertensii. | C. ndrosia.
digitata. var. picta.
Shape of bell. | Flat. Flat. Fiat. Fiat. Flat. Rounded with-|) Exumbrella
outaconcay-| concave.
ity on exum-
brella.

Number of 12 16 16 16 14 to 16 16 18 to 22
rhopalia.

Number of 5 5 Variable, 9 Variable, 5 to| 8 4
marginal lap- about 5. 12 usually.
pets in each
paramere.

Length of 0.757r tor. r+ I to 0.5 Less than. | Less than r. | 1 too.5 r 1too.5r
mouth-arms
in terms of
bell-radius(r).

Vesicles and fil-| Only flat leaf-| Very small Very small Very small Very small Very large Small leaf-
aments among) shaped vesi- | clubs. clubs. clubs. clubs. clubs. shaped vesi-
mouths. cles. cles.

Where found. | West Indiesto| Pelew Islands, Malay Arch- | Coast of Moz-| Red Sea. Caroline Is- | Fiji Islands.

Florida. New Guinea, ipelago. ambique, lands.
East Africa,
Madagascar.

bell-radius. 4 to 6 flat, short side branches arise from each arm in a tree-like manner and these
in turn give off side branchlets. Numerous small and 5 or more large, club-shaped vesicles on
each arm between the mouths. The largest 2 to 3 times as long as width of main branches of
arms. 4 small subgenital ostia.

Color very brilliant and variable.

Exumbrella is reddish-brown to violet-brown, with

milk-white spots, between which are dark, radial stripes. Bell-margin usually bluish or violet.
The milk-white spots on the exumbrella are disposed as follows: A large oval spot above
each sense-organ and a small, white spot upon each lappet. Thus there are 80 (5X 16) small

RHIZOSTOMA—CASSIOPEA. 639

spots and 16 large ones, 96 in all. Mouth-arms olive-green to reddish-brown, spotted
with white.

This East Indian species ranges from the Red Sea to Sumatra, giving rise to a number
of color varieties and local races, many of which have been described as distinct species.
Keller, 1888, records its having wandered into the Suez Canal from the Red Sea. Hartlaub
gives a good description of hig species from Dyjibuti, East Africa.

Cassiopea andromeda var. zanzibarica Chun.

Cassiopea andromeda var. zanzibarica, Cuun, 1896, Mittheil, Naturhistorischen Museum, Hamburg, Bd. 13, p. 17.

This resembles C. andromeda in most respects, being 40 to go mm. in diameter and with
a flat disk. But it is said to be distinguished from C. andromeda by having 6 velar and 2
ocular lappets between each successive pair of sense-organs instead of 5 lappets as in the typical
C. andromeda. The 5 large clubs or filaments of the arm-disk are also larger than in C. andro-
meda, being 10 mm. long. The color is also different, but quite variable. The exumbrella is
usually brownish-red with 16 to 17 smoky-gray or white radial streaks, which commence at the
outer edge of the central concavity of the exumbrella and fork over the lappets. Some speci-
mens have 3 white spots over the velar lappets between each successive pair of sense-organs.
Mouth-arms light whitish-red beset with small brownish-white clubs. The large central fila-
ments are black. Zanzibar coast, July to August. This is certainly identical with C. andromeda.

Cassiopea andromeda var. malayensis Maas.

Cassiopeja andromeda var. malayensis, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, pp- 40, 433 taf. 4, fign.

24-25, 27-29, 31-34; taf. 11, fign. 98, 102; taf. 12, fig. 112.

This medusa is said to be distinguished by having 1, 3, 5, 7, or more velar lappets between
each pair of ocular lappets. Mouth-arms are slightly shorter than bell-radius and compressed
dorso-ventrally. A few large, isolated, club-shaped appendages are between the frilled mouths,
but these are not commonly found upon all of the mouth-arms. Indeed, there are usually
only 2 to 3 of these large clubs upon the mouth-arms of any individual medusa. The umbrella
may become more than 200 mm. in diameter, although such large specimens are rare. There
are normally 16 small, marginal sense-organs, each containing an entodermal concretion and
a distal pigment spot. Young medusz have about 32 eeulai and 16 velar lappets, but as
growth proceeds the velar lappets increase by division so as to become 3, 5, 7, or more times
as many as in the young medusa. ‘The mouth-arms are compressed dorso-ventrally, those
of C. acyclobbra laterally. They branch quite irregularly, in a tree-like manner.

Both male and female medusz are described by Maas, who records numerous examples
ranging from 10 to 200 or more millimeters in diameter. The medusa appears to be widely
distributed among the islands of the Malay Archipelago. Color (7) It is evidently identical
with C. andromeda.

Cassiopea andromeda var. maldivensis.

Cassiopea andromeda var. maldivensis, BRowNe, 1905, Fauna and Geog. Maldive and Laccadive Archipelagoes, vol. 2, p. 962.

The disk is about 75 mm. wide with a central concavity on the exumbrella. The mar-
ginal sense-organs range from 12 to 19 but are usually about 16. The marginal lappets are
very indistinct and range from about 7 to 10 between each successive pair of sense-organs.

There are 4 pairs of complexly branching mouth-arms which project slightly beyond the
umbrella margin. The proximal branches of the mouth-arms are generally alternate in
position, while those of the outer branches are generally dichotomous. There are numerous
disk-shaped, flat, spatula-like appendages over the mouth-arms. These are adjacent to the
oscula. There are also cylindrical or somewhat flattened linear appendages and hand-shaped
appendages, which are almost leaf-like and may be 25 mm. long and g mm. wide. There
are not more than § to 6 of these cylindrical or hand-shaped appendages on each of the 8
mouth-arms. At the center of the mouth-arm disk there is a single long appendage which
may either be cylindrical or hand-shaped.

The 4 subgenital ostia are very small and triangular in shape, about 3 mm. wide. The
gonads are similar to those of Cassiopea Rimachanas abe peripheral canal-system consists

640 MEDUS.-E OF THE WORLD.

of a radial-canal to each sense-organ and an equal number of intermediate radial-canals.
These radial-canals are put into communication one with another by means of an anastomosing
network of vessels, without any distinct ring-canal.

The medusa is dark-green without spots or pattern of any sort. This green color is due
to an infesting alga, one of the Zooxanthelle. The chlorophyl is readily dissolved out in
alcohol.

This medusa is abundant in muddy bottoms in shallow bays and back waters at Febidu,
Maldive Islands, Indian Ocean. It is a well-marked variety of C. andromeda being distin-
guished by the hand-shaped appendages on its mouth-arms.

Cassiopea andromeda var. acycloblia Schultze.

Cassiopeja acycloblia, Scnuitze, L.S., 1898, Denkschrift. Med. Nat. Ges. Jena., Bd. 8, p. 459, taf. 33, fign.2, 4,6; taf. 34, fig. 15.
Cassiopeja andromeda var. cyclobalia, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 40.

Bell flat with low central dome (?) at middle of the exumbrella. 16 marginal sense-organs
and 80 (516) marginal lappets. Exumbrella flecked with 16 large, oval, radially placed,
white spots above the sense-organs. A wide ring of white binds these spots together. There
is also a small, white fleck on the exumbrella side of each lappet. Those of the interradial
lappets fuse with the white ring. Mouth-arms only three-fifths as long as bell-radius. Diam-
eter of arm-disk less than disk-radius. The mouth-arms branch dichotomously each with
one large and many small filaments. Amboina, Molucca Islands.

Cassiopea polypoides Keller.
Cassiopea polypoides, Ketrer, 1883, Zeit. fiir wissen. Zool., Bd. 38, p. 632, taf. 36, 37, fign. 6-23.

Disk flat, shield-shaped, 100 to 150 mm. wide. A well-developed, sucker-like depression
at center of exumbrella, the thickened, outer edge of which is 15 to 20 mm. inward from bell-
margin. Margin of umbrella is thin, but at the center there is a disk-like thickening of the
gelatinous substance of the exumbrella. There are 16 marginal sense-organs each with a
red-brown ectodermal pigment spot and a terminal entodermal mass of concretions. 80
short, rounded, marginal lappets, 3 velar and 2 ocular in each of the 16 parameres. The
thick mouth-arm disk is octagonal with 8 equal sides. The 8 mouth-arms are about as long
as radius of umbrella and project beyond the bell-margin; they are somewhat higher than
wide at their distal ends. Each arm usually gives rise to 3 pairs of alternately arranged,
pinnate branches, with frilled mouths on their under sides. There are numerous clubs and
filaments on the under sides of the mouth-arms, the largest filaments being over 30 mm.
long. The 4 subgenital ostia are small, oval, and elongated in a radial direction. Central
stomach 4-sided. 32 radiating canals extend outward from the stomach, 16 to the sense-
organs and 16 to intermediate parts of the margin; these 32 vessels connect one with another
by an anastomosing network of vessels, but there are no specialized ring-canals.

Exumbrella light-brown with a broad, white ring which gives out 16 cog-wheel-like,
forked branches on its outer edge in the radii of the 16 sense-organs. There is also a short,
white, radial spot on the exumbrella side of each marginal lappet. 16 dull white, spoke-like
pigment areas in the gelatinous substance of the exumbrella extend outward from the edges
of the central thickening to near the margin in the radii of the sense-organs. The suctorial
mouths are intense brown. Appendages and filaments of mouth-arms very variable in color,
being sky-blue, greenish-blue, horny-yellow, translucent white, or rose-red. In one variety
the exumbrella is olive colored with indistinct radiating spots.

Keller distinguishes five varieties based on color and differences in number of the mouth-
arm filaments as follows:

Cyanea. With 5 or 6 large, sky-blue or green-blue filaments on each mouth-arm. The white oral tufts are numerous,
the clubs rarer. This is the commonest form.

Flava. The large filaments are twisted, never flattened, their color is horny-yellow or translucent. Common.

Albida. The large filaments are twisted and white in color. Rare.

Rosea. Exumbrella olive colored, radial spots indistinct. Some of the filaments are round, some flat, and of rose-red
color. Club-shaped vesicles rare. Rare variety.

Herbacea. The mouth-arm appendages poorly developed. No filaments. The mouths of the arms large. Rare variety.

PLaTE 69.

. Cassiopea frondosa, young medusa, natural size. From the bottom

of the moat of Fort Jefferson, Tortugas, Florida, July 10, 1905.

. Casstopea frondosa. Half of exumbrella shown on the right, and half

of the subumbrella on the left. Natural size. Moat of Fort
Jefferson, Tortugas, Florida, June 23, 1907.

. Sense-organ of Cassiopea frondosa seen from subumbrella side.
. Cassiopea xamachana, side view, natural size. From moat of Fort

Jefferson, Tortugas, Florida, July, 1905.

. The mouth-arms of Cassiopea xamachana, showing side walls of

stomach cut across. The canal-system in the arms is shown in
green.

. Sagittal section of Cassiopea xamachana, showing stomach cavity.

The gastroyascular canals are shown in green.

. Casstopea xamachana. One of the mouth-arms cut off so as to show

the branching of the canal-system.

. Cassiopea xamachana. Cross-section of a mouth-arm near its base

showing the fringing tentacles of the mouths and the shapes of
the appendages.

Drawn from life, by the author.

PLATE 69

MAYER

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_

PLATE 70.

Fig. 1. A rare, small-sized variety of Casstopea xamachana. In its color-
pattern this resembles Casstopea ndrosia from the Fiji Islands.
Fig. 2. The common color-pattern of Casstopea xamachana.

Cassiopea xamachana photographed from life, from specimens
found in the moat at Fort Jefferson, Tortugas, Florida, July,
1905. Natural size.

MAYER

Puate 71.

Cassiopea xamachana, photographed from

* size, July, 1908. From the moat at Fort Jeffe
; and aboral views of the medusa.

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i Ly “
Da ee

A iF Wy
@ 7 4
ye is sd vi

RHIZOSTOMA.—CASSIOPBA, 641

This medusa was found in large swarms by Keller on the shallow coral flats of the southern
parts of the Red Sea. It differs from the ty pical C. andromeda 1n the thick-rimmed sucker of
the exumbrella, and the long, laterally compressed arms. Keller describes it in detail with good
hgures. In common with other Cassiopeide it les upon the bottom with its oral surface
uppermost. Keller draws comparisons between its habits and structure and those of actinians,
etc. This medusa is probably only a local variety of C. andromeda, but the thick, sucker-
like disk at the middle of the exumbrella appears to distinguish it.

Cassiopea xamachana R. P. Bigelow.
Plate 69, figs. 4 to 8; plates 70 and 71; plate 72, the seven lower figures.

Cassiopea xamachana, BicEe.ow, R. P., 1892, Zoolog. Anzeiger, Bd. 15, p- 212; Johns Hopkins University Circulars, 1892,
vol. 11, pp- 71,84; 1893, Journal Institute of Jamaica, vol. 1, p. 301, 1 plate; 1900, Memoirs Boston Soc. Nat. Hist., vol. 5,
No. 6, p. 191, figs. A to L, plates 31 to 38, 66 figs.—Perkins, 1905, Year Book of the Carnegie Institution, No. 4, p. 118.
Publication, No. 102, p. 150, plate 4.—Mayer, 1906, Year Book of the Carnegie Institution, No. 4, p. 117; Publication
of the Carnegie Institution, No. 47, 62 pp. (rhythmical pulsation); 1907, Year Book Carnegie Institution, No. 6, p. 1213
Ibid., 1908, No. 7, p. 123-—Maas, 1903, Scyphomedusen der Sboga Expedition, Monog. 11, p. 40.—SrockarD, 1907,
Year Book Carnegie Institution of Washington, No. 6, p. 119 (regeneration); Ibid., No. 7, 1908, p. 130.—Papers from
Tortugas Laboratory of Carnegie eae of Washington, vol. 2, p. 61, figs. 1-29; Journal of Experimental Zoology,
1909, vol. 6, p. 433, 8 figs —Zeteny, C., 1907, Journal Experimental Zoology, vol. 5, p- 265, 4 text-figs. (regeneration )—
DaHLGREN AND KFPNeER, 1908, Text- aor of ae of Animal Histology, p. 88, fig. 85 (histology of muscles)—Mayer,
1908, Papers from the Tortugas Laboratory of the Carnegie Institution of Washington Publication, No. 102, p. 113 (the
cause of rhythmical pulsation); Popular Science Monthly, vol. 73, pp. 481-487, 4 figs.; 1909, Report of a Interna-
tional Zoological Congress, 4 pp-—Harvey 1909, Year Book of the Carnegie Institution of Washington, No. 8, p. 129.

Casstopea frondosa, Fewxes, 1882, Bull. Museum Comp. Zool. at Harvard College, vol. 9, p. 254, plate 1, figs. 7-19; plate 2,
figs. 1, 2; plate 3, figs. 1-3, 9, 10; Ibid., 1883, p. 80, plate 1, fig. 16.

The disk is usually about 150 mm. in diameter, although Bigelow records one from
Jamaica 240 mm. wide. It is flat and with rounded edges. Pine is a well-marked con-
cavity at the middle of the exumbrella, the diameter of hich is about equal to the disk-
radius. It enables the medusa to cling firmly to any smooth surface as by a sucker.

The number of the marginal sense-organs ranges from II to 23, although there are usually
about 16. For example, in 25 meduse taken at random and ranging in size from 23 to 149
mm., one had 13 marginal sense-organs, 1 had 14, 1 had 15, 12 had 16, 5 had 17, 1 had 18,
2 had 19, 2 had 20. I have seen one medusa with 11 and one with 23 sense-organs. The
number is independent of the size of medusa, being determined at time of strobilization.

The sense-organs are short, blunt, and club-shaped, and are set within niches protected
above by a shelf-like membrane spanning the cleft between the adjacent lappets. There is
no exumbrella pit above the club. Each sense-organ contains a terminal mass of entodermal
crystals and an aboral cup-shaped ectodermal ocellus having reddish-brown pigment. There
are 5 short, blunt, rounded lappets between each successive pair of sense-organs; the 2 lappets
adjacent to the sense-organs are only about half as wide as the others. The mouth-arm disk,
which projects as a flat ‘plate from the center of the subumbrella, is only about two-thirds as
wide as the disk-radius. 4 pairs of adradial mouth-arms arise from this disk. Each of these
8 mouth-arms is about 1.25 times as long as radius of bell and projects somewhat beyond
bell-margin. These mouth-arms are triangular in cross-section, their aboral surface being
broad and flat; they each give rise to 10 to 15 alternate, primary branches, which in turn
give rise to secondary branches. These branches are commonly longer than in C. frondosa
and are also longer and stouter than in C. andromeda Eschscholtz and more slender and have
more primary branches than in C. polypoides Keller. In the axil of each primary branch of
the mouth-arms is a single, flat, ribbon-like filament, which varies in length with the size of
the adjacent branch. There are also 5 to 13 large, ribbon-shaped Fesreae upon the oral
surface of the mouth-arm disk. The largest flament is at the center and is fully one-fourth
as long as the bell-diameter. The Rlomene decrease successively in length out over the
mouth-arms; those at the tips of the arms being only about one-seventh as long as those at
the center. In addition to the filaments there are numerous short club-shaped, nematocyst-
bearing vesicles scattered among the mouths.

The mouths are found upon the oral (uppermost as the medusa lies upon the bottom)
sides of the primary and secondary branches of the mouth-arms, and to some extent upon
the oral sides of the 8 basal trunks of the arms. There are no mouths at the center of the

642 MEDUS OF THE WORLD.

J 2 ppt 2 ¢

Fic. 402.—Cassiopea xamachana, after R. P. Bigelow, in Mem. Boston Soc. Natural History.

b, bud growing from side of scyphostoma; cd, circum-oral disk or peristome; ¢/, interrhopalial lobe; /t,
interrhopalial tentacle; m, mouth; p, proboscis; rht, rhopalial tentacle; 1, tentacle.

25, strobila, in which the degeneration of the rhopalial tentacles is nearly complete and the interrhopalial
tentacles have begun todegencrate. 26, a complete strobila. The basal polyp bears a bud which
broke off and swam away while the drawing was being made. The ephyrula was detached during
the following night. The rhopalia are visible through the umbrella. At y is a pair of twin
rhopalia (compare y, fig. 30). 27, basal polyp of same specimen, a few hours after separation of
the ephyrula. 28, optical section of same. 29, an ephyrula recently set free; oral aspect, gastric
filaments visible through mouth; X 31. 30, specimen of about same age, showing variations of
margin at u, w, y and z; X 31.

RHIZOSTOM.8—CASSIOPBA. 643

mouth-arm disk in the full-grown medusa, although they are commonly found near the edges
of the disk. The mouths are fringed with a multitude of fine, waving tentacles.

There are 4 small, deep, oval-shaped, interradial subgenital pits, and 4 separate in-
vaginated genital sacs. Central stomach is cruciform, being encroached upon at the inter-
radial sides by the 4 sac-like gonads. The axial ducts of the 8 mouth-arms empty into this
central stomach at the 4 principal radu. The central stomach also gives rise to twice as many
radial vessels in the subumbrella as there are marginal sense-organs. Every alternate vessel
extends to a sense-organ, the others going to intermediate parts of the rim. All of these radial
vessels are put into communication one with the other by numerous anastomosing branches;
but there is no well-defined circular vessel such as is figured by Haeckel in Casstopea ornata.

There is a well-developed zone of circular muscle-fibers in the outer half-radius of the
subumbrella. These have a more and more cuspate trend as we near the center of the disk,
there being twice as many cusps as there are radial vessels, the outward convexities of the
cusps being between the vessels.

The general color of the medusa is greenish-gray-blue, the greenish color being due to
clusters of commensal plant-cells within the gelatinous substance of the disk near the surface.
If the medusa be maintained in darkness for a month this green color disappears, leaving
the animal a pale, translucent blue-gray. Around the outer edge of the central concavity
of the exumbrella is a wide, dull white circle, edged on its inner side with faint gray-brown.
A more or less Y-shaped, radial, white stripe extends outward from the broad ring in the
radii of the sense-organs, the sense-organ being in the center of the crotch of the Y._ In addition
a single, radial stripe extends outward down the middle of the exumbrella side of each marginal
lappet. Occasionally these radial stripes are more or less separated from the broad, white
circle. Conspicuous spoke-like, white stripes extend outward in the radii of the sense-organs.
These are white regions found in the gelatinous substance of the bell and extend half-way through
the gelatinous bibscinee from the subumbrella toward the exumbrella surface. The mouths,
AGH: and yesicles are olive or olive-brown, the vesicles and filaments being of a decided
green. Among the many color varieties there is a rare one in which the spoke- are dull white
spots are diamond-shaped, and there is no broad, white ring on the exumbrella. The whole
medusa is more translucent than are the more abundant medusz with the white ring. They
are also smaller than the common form. Curiously enough this color variety bears a striking
resemblance to Cassiopea ndrosia Agassiz and Mayer, from the Fiji Islands, South Bacihic.
Various forms of its color patterns are shown in the photographs i in plates 70 to 72 taken
from life by the author.

I find that Cassiopea can thrive well in darkness for more than a month, hence the
medusa is not dependent upon its commensal plant cells for the oxygen it requires. In this
connection Whitney, 1907 (Biol. Bulletin, vol. 13, No. 6, p. 291), finds that if green hydra
be placed temporarily in a 0.5 to 1.5 per cent solution of glycerin, the green alge (Chlorella
vulgaris) pass out throughthe mouth. Then if the /ydra be replaced in water it will grow nor-
mally, but remains clear and does not regain the green bodies even when placed in an aquar-
ium with alge.

This Seduce was discovered in great abundance by Dr. R. P. Bigelow in a salt-water
lagoon called the Great Salt Pond, near Port Henderson, Kingston Harbor Jamaica. It is also
exceedingly common in the salt-water moat of Fort Jefferson, Tortugas, Florida, where it is
found upon the weedy bottom throughout the summer; and it occurs in many semi-stagnant,
salt lagoons along the Florida eer as far north as Miami.

The early stages of the development of the egg into the scyphostoma are as yet unob-
served, but the process of the formation of asexual buds by the scy phostoma has been elabor-
ately studied by Bigelow and observed also by Perkins. The buds arise from the perradial
sides of the calyx of the scyphostoma near the point of origin of the stem. Scy phostomz are
never found with more than two buds attached. When two are present the older is always
attached to the apex of the younger bud. The bud is at first hemispherical, hernia-like; then
elongated, and finally spindle-shaped. The ectoderm, entoderm, and mesogloea of the bud
are produced from the corresponding layers of the parent scyphostoma, and the 4 ectodermal,
septal muscles of the bud are derived from out-growths of one or both of the septal muscles
of the parent which lie in the interradii adjoining the perradial area of bud formation. The

644 MEDUS4 OF THE WORLD.

bud is set free as a spindle-shaped larva which swims by means of cilia. After 2 or 3 days the
mouth breaks through at the pole which was at the proximal end of the bud while it was
attached to the parent, thus resembling the case of budding in Cotylorhiza. The mouth is
not formed by an inyagination of the ectoderm, but breaks through by the local disappear-
ance of the mesogloea and the fusion of ectoderm and entoderm at the posterior end of the
larva. The anterior end then elongates to form the stem of attachment, and in about 4 or 5
days after being set free the larva fastens itself to some solid object.

Fic. 403.—Cassiopea xamachana, after R. P. Bigelow, in Mem. Boston Soc. Natural History.

it, interrhopalial tentacle; oc, ocellus; rht, rhopalial tentacle; x, abnormal branched tentacle.

17, scyphostoma showing first traces of rhopalial structure. 18, small part of margin, more highly magni-
fied. 19, scyphostoma at slightly older stage. 20, small part of margin of similar larva. 21,
early stage in strobilization. 22, rhopalial tentacle of same specimen seen from side. 23, older
rhopalial tentacle. 24, strobila in which rhopalial tentacles have begun to degenerate.

RHIZOSTOMA®—CASSIOPEA. 645

Usually the 4 perradial tentacles are soon supplemented by the 4 interradial ones, and in
about 3 days after they first appear the 8 tentacles are as long as the proboscis of the scypho-
stoma. 8 adradial tentacles then develop. The number and arrangement of the tentacles is,
however, very variable but finally there are about 32. In any case there are twice as many
tentacles as there are to be rhopalia. The full-grown tentacles are tapering, slender, and
about 3 times as long as the body of the scyphostoma. Half of them are erect, and the alternate
half stretch more horizontally outward.

The 4 primary, gastric pouches are not formed by evagination from ectoderm and ento-
derm alternately, as in the sexually produced scyphostoma of Aurellia, etc., according to Goette,
but are wholly entodermal as Hadzi finds them to be in Chrysaora, and simply separated one
from another by the ingrowth of the 4 interradial septa. At first the septa are simple, entire
buttresses of entoderm with an axial sheet of mesogloea, but later each septum becomes perfor-
ated immediately under the interradial tentacles, thus forming a ring-sinus. There are 4 longi-
tudinal strands of septal muscles, 1 in the mesogloea of each septum.

The septa bear no definite relation to the exact position of the interradial tentacles, for
these may arise on either side of or in the plane of a septum. In this respect the scypho-
stomz resemble those of Aurellia and Cotylorhiza according to Claus, and differ from the
Anthozoa, in which the tentacles are invariably interseptal. Every alternate tentacle stands
erect while the others extend outwardly. When the scyphostoma disk is about 2 mm. wide,
conical enlargements which contain crystalline concretions are observed in the entoderm at the
bases of the erect tentacles. An ectodermal ocellus develops upon the aboral (lower) side
of each conical enlargement and the tentacle itself begins to degenerate, becoming finally
absorbed, leaving only the sense-club with its ectodermal ocellus and terminal mass of ento-
dermal concretions. When the tentacles begin to degenerate, slight pulsating movements of
the disk commence. The marginal lobes grow out while the rhopalia are being formed, and
finally the interrhopalial tentacles are also absorbed.

Strobilization is monodiscus, but the scyphostoma after setting free the ephyra develops
new tentacles and gastric pouches, and may strobilate a second time.

The young ephyra has the same number of marginal sense-organs as the adult medusa.
It has 4 simple lips and a central mouth-opening. Then the angles of the lips become extended
to form 8 oral arms, very much as in the adult 4urosa. Then there is a stage wherein the
oesophagus is divided into 4 tubes with 3 osculz and an oral vesicle on each arm. Rhizostoma
and Cotylorhiza go through a similar stage. The septal muscles and their funnel-cavities
disappear wholly in the ephyra, as do also the 4 interradial septa.

Mayer, 1906, 1907, 1908, finds that the stimulus which produces pulsation in Cassiopea
is nervous in nature and will pass over newly regenerated tissue which contains neryous,
but no muscular elements. Moreover, if the muscles be paralyzed by magnesium the pulsation-
stimulus still travels through the nervous network of the subumbrella, even though the muscles
can not respond to its presence by contraction.

If an annulus, or strip of any shape constituting a closed circuit, be cut from the sub-
umbrella and stimulated momentarily at any one point, 2 waves of contraction start in opposite
directions around the strip from this stimulated point. By pressing upon one side of the ring
we dampen and reduce the strength of the initial wave passing over that side, and when the
two waves meet the stronger wave overpowers and annuls the weak one. Thus a single
contraction-wave is entrapped in the ring-circuit and travels constantly around it at a uni-
form rate. The mechanical arrangement of the pulsating medusa in nature is such as
to prevent the formation of such continuous pulsation-waves—the pulsations are recurrent
and each contraction-wave is annulled as soon as it has produced a single contraction of
the medusa.

The sea-water is a balanced fluid for the medusa, neither stimulating nor inhibiting its
pulsations. This balance is due to the fact that the tonic sodrum of sea-water is a powerful
nervous and muscular stimulant, but the magnesium, calctum, and potassium are inhibitors
and annul the stimulus produced by the sodium. If calcium be absent the magnesium quickly
checks all pulsation. On the other hand, a slight increase in the sodium serves as a nervous
stimulus which overcomes the inhibiting tendency of the magnesium, calcium, and potassium
and produces contraction.

646 MEDUS® OF THE WORLD.

The pulsation-stimulus is engendered in the marginal sense-organs. A uric oxalate of
sodium is developed constantly in the entodermal cells of the outer end of each sense-club.
This oxalate precipitates calcium, thus forming the crystalline concretions which consist of
calcium uric oxalate, and at the same time it sets free such soluble stimulants as NaCl and
Na,So,. Thus we find that the sense-clubs are engaged in the maintenance of a slight con-
centration of sodium over and above that found in the sea-water itself. This slight excess of
the sodium ion is a stimulant to the nervous elements within the sense-club and the nervous
elements respond to it recurrently, producing the rhythmical contractions of the muscles.

If a disk without marginal sense-organs be set into pulsation and then disturbed by a
sudden current in the sea-water, etc., it displays excitement by markedly increasing the
amplitude of its pulsations. Hence its ability to display excitement is not dependent upon
the sense-organs, but upon the general nervous tissues of the subumbrella.

When the marginal sense-organs regenerate, each one appears with a short, hernia-like
side branch, which disappears later. In this connection it is interesting to see that the sense-
organs are normally formed as side buds from the bases of each alternate tentacle of the
scyphostoma, and then the tentacles themselves are absorbed. Thus when they regenerate
they display a tendency to replace the tentacle as well as the sense-club.

In 1909 I succeeded in grafting two individuals of C. xamachana, side by side, so that their
subumbrellas joined. The double-medusa then pulsated constantly at the rate of the faster
individual which initiated and controlled all of the rhythmical movements; but if one pinched
the controlled medusa its rate increased and it then assumed a temporary control of the
double animal. Hence the complex always pulsated at the rate of its fastest member. Hargitt
attained a similar result with 2 individuals of Gonionemus murbachi1, but in this case the
rims were attached around nearly their entire edges so that any movement of one medusa
must necessarily cause a corresponding movement of the other. In the two Cassiopeas,
however, the contact was at a single narrow bridge of tissue only, and indeed the medusz pul-
sated independently until the nerve-nets of their subumbrellas joined in the process of
regeneration.

The color of the umbrella of C. xamachana is mainly due to the presence of numerous
symbiotic alga, Zooxanthelle, which Bigelow finds contain starch, cellulose, and chlorophyl.
These plant cells are globular and occur in small clusters imbedded in the mesogloea and are
greenish-brown in color.

A well-marked, conical, pit-like depression is occasionally seen upon the aboral side of
each mouth-arm near its point of origin from the arm-disk, but more commonly in male than
in female medusz. The female meduse greatly outnumber the males. Perkins believes that
the medusz may be hermaphroditic, but of this we have no evidence. Pseudorhiza haeckelii
is, however, known to be hermaphroditic, the spermaries being in the gutters of the mouth-
arms.

Zeleny, 1907, finds that meduse maintained in pulsation appear to regenerate at about the
same rate as if the disk were at rest. Certainly the functional activity of pulsation seems to
be of no aid in accelerating regeneration, for Stockard also finds that the medusa regenerates
at practically the same rate whether it be pulsating or at rest.

Stockard, 1907, discovered that tissues removed from various parts of the subumbrella
regenerate more rapidly the nearer they are to the disk-center, and less rapidly as the periph-
ery is approached, thus according with the rule discovered by Morgan in the regenerating
fish’s fin—the deeper the level of the cut the more rapid the regeneration.

In 1908, Stockard made the interesting discovery that if the medusz be starved while they
regenerate lost arms the disk of the medusa shrinks during the process of regeneration, and its
rate of decrease is greater the greater the number of removed arms. The regenerating tissue
evidently possesses a greater capacity for absorbing nutriment than does the somatic tissue of
the disk itself, and in this respect the regenerating tissue behaves as does that of cancer which
grows rapidly even when the normal tissues surrounding it are wasting away. (See Year Book
of the Carnegie Institution of Washington, No. 7, p. 131, 1908.)

PLATE 72.

The three uppermost figures are of Cassiopea frondosa and show two varieties
of color pattern of the exumbrella and one view of the subumbrella
showing the mouth-arms.

The seven lower figures are of Cassiopea xamachana showing varieties in
color-pattern. The central figure shows the exumbrella of a medusa
with its mouth-arms spreading outward.

Photographed from life, by the author. The medusz are seen upon a
sandy bottom.

MAYER

RHIZOSTOM22—CASSIOPBA. 647

Cassiopea frondosa Lamarck.
Plate 69, figs. 1 to 3; plate 72, the 3 upper figures.

Medusa frondosa, Patras, 1774, Spicilegia Zoolog., fasc. 10, pp. 29, 30, plate 2, figs. 1-3.—Gexin, 1788, Linné’s Syst. Nature,
tomus I, pars 6, p. 3157.—Bosc, 1802, Hist. Nat. d Vers., tome 2, p. 170.

Casstopea frondosa, LAMARCK, 1816, Hist. Nat. Anim. sans Vert., tome 2, p. 512.—Escuscnoxmz, 1829, Syst. der Acalephen,
p- 43-—Titestus, 1834, Acad. Caes. Leop. Nova. eel aoe 15, pp- ee. 278, tab. 72, fign. 1-5.—Lesson, 1843, Zooph.
Acal., p. 405. —Mitne-Epwarps, 1849, Cuvier’s Régne Animal, Zooph., plate 51, fan 3-—Perkins, 1906, Year Book,
cares Institution of Washington, No. 4, p. 115; 1908, Publication No. 102, Carnegie Institution of Washington, p. 152,

Pak Péron et Lesueur, 1808, Annal. der Mus. Hist. Nat., Paris, tome 14, p. 357, Nr. 85.

Polyclonia frondosa, Acassiz, L., 1860, Cont. Nat. Hist. U. S., vol. 3, plates 13, 13a; 1862, Ibid., vol. 4, pp. 139-148, 159.—
Acassiz, A., 1865, North Amer. Acal., p. 41; 1881, ee vol. 24, p- 509. See also Archer, H, Ibid., p. 307.-—HakrckE1,
1880, Syst der Medusen, p. 568; 1881, Report H. M. S. Challenger, Zool., vol. 4, p. XVI11.—VANHOFFEN, 1888, Bibliotheca
Zoologica, Bd. 1, Heft. 3, p. 4o.—Bicetow, R. P., 1893, Johns Hopkins University Circulars, vol. 2, No. 16 p- 106 (habits,
physiclogy).—Pzrxins, 1906, Year Book, iGamere Institution of Washington, No. 4, p. 115; 1908, Publication No. 102,
Carnegie Institution of Washington, p. 152
Disk flat, with rounded edge, and about 120 to 260 mm. in diameter. There is no con-

cavity at the center of the exumbrella, such as is seen in Casstopea xamachana. There are

constantly 12 marginal sense-organs in C. frondosa, 4 perradial, 8 adradial, each of which
contains a terminal entodermal mass of crystalline concretions. ‘There are no ocelli. There
are 60 short, subrectangular, nearly straight- -edged, marginal lappets, 5 between each suc-
cessive pair of marginal sense-organs. ‘The lappets flanking the sense-organs are only half
as wide as the other lappets. The 4 pairs of mouth-arms arise from a Shallow: flat, ats
arm-disk at the center of the subumbrella; this arm-disk is not quite as wide as the semi-
diameter of the medusa. The mouth-arms, which are about three-fourths as longas bell-radius,
usually bifurcate at their free ends and give mse to short, pinnate side branches from their
oral sides; but occasionally the branches are quite long as in Cassiopea xamachana. The
numerous frilled mouths are found exclusively upon their lower sides, the upper sides of the
mouth-arms being smooth and without mouths. Scattered quite uniformly between the mouths
are 30 to 40 small, expanded, flat, leaf-shaped vesicles. “There are 4 small, round, interradial,
subgenital pits, ad 4 separate, invaginated genital sacs which project into the stomach- -cavity.

A duet extends from each of the 8 oe arms into the central stomach, and 24 radial-canals

pass outward from the stomach into the subumbrella and are put into communication one

with another by a network of anastomosing vessels. 12 radial-canals go to the marginal
sense-organs and 12 are intermediate in position.

General color of gelatinous substance amber-yellow, slightly olive, or greenish. Just
above each of the 12 marginal sense-organs 1s usually a single, large, bilateral: bean-shaped
white spot in the gelatinous substance bf the exumbrella. There is GO a smaller white spot
in each marginal lappet, and above this an irregular line of 3 to § smaller white spots between
each successive pair of marginal sense-organs. A more or less broken, axial, white line extends
through the length of Sa mouth-arm in the gelatinous substance. The frilled mouths are
of a cinnamon color and the leaf-like vesicles are opaque, dull white. The spots upon the
bell are very variable in number and arrangement (see plates 69 and 72).

Cassiopea frondosa is found throughout the West Indian region and the Florida Reefs.
Perkins observes that it lives upon sandy rather than weedy bottoms. In common with other
species of the genus it lies upon the bottom with the oral surface and mouth-arms uppermost.
In this position it remains for long intervals of time, slowly contracting its disk in a sluggish
rhythm. This movement serves not only to maintain the disk upon the bottom, but to create
a water-current over the mouth-arms. It prefers purer water than C. xamachana, and is usually
found in protected places among the mangroyes in the cuts between the Florida Keys.

In Kingston Harbor, Jamaica, this medusa is found upon the muddy bottoms of protected
lagoons, especially in those surrounded by mangroves, near the harbor entrance where the
water is quite pure. In Jamaica it attains to a far greater size than in Florida. A specimen
which I found in a mangrove lagoon near Port Royal in March, 1909, was of the following
dimensions in mm.: Bell 259 mide! arm-disk 95 wide, mouth-arms 129 long, pinnately and
complexly branched, and projecting beyond the rim of the bell. Color as in the Florida
specimens. J am told that the medusa becomes even larger in Jamaica.

L. Agassiz (1862, p. 147) showed that the young ephyra of this species possesses a central
mouth-opening which disappears in the adult.

648 MEDUS OF THE WORLD.

C. frondosa can at once be distinguished from C. xamachana by its amber color, the
absence of ocelli on its rhopalia, the absence of a sucker-like concavity upon its exumbrella,
and by the fact that it has constantly 12 marginal sense-organs, whereas C. xamachana has
11 to 23 (see plate 69). It is far less hardy in aquaria than C. xamachana,

According to Bigelow, 1893, in the adult female the mouths disappear from the oral
disk while at the same time the oral vesicles increase in number until they are closely crowded
together and completely cover it. The eggs are discharged from the ovaries into the stomach,
where cleavage begins; they then pass out on to the oral disk and are to be found there in large
numbers, cemented together in small, reticulated clusters at the bases of the vesicles; they
remain there until some time after they have become ciliated planula. Bigelow reared the
scyphostome of this species to the 8-tentacle stage. The young scyphostoma appears to be
entirely similar to those of other species of Caksinpen:

Dr. R. P. Bigelow has shown that while the vesicles on the oral surface of the disk serve
to protect the young, those of the mouth-arms serve to capture food. These vesicles usually
stand upright, but upon being struck by an unwary copepod they bend down and close the
mouth of the nearest funnel in the manner of a lid. The prey thus finds itself within one of
the mouths, tightly shut in by the overlying vesicle.

Cassiopea ornata Haeckel.

Cassiopea ornata, Hatcket, 1880, Syst. der Medusen, p. 570, taf. 37, fign. 1-8.—Hamann, 1881, Jena. Zeit. fiir Naturw., Bd. 15,
p- 248 (structure of the mouth-arms).

Bell 100 to 120 mm. wide, 30 to 40 mm. high, flat and shield-shaped. 16 rhopalia, 80
lappets, 96 white spots, as in C. andromeda. Mouth-arms cylindrical, slender, and somewhat
longer than bell-radius, not broad and flat as in the typical C. andromeda. ‘There are only
small, club-shaped vesicles between the mouths. The characteristic feature of this species 1s
said to be the presence of 2 distinct ring-canals. The inner ring-canal connects the 16 principal
radial-canals at an annulus some distance inward from the margin, while the outer ring-
canal is at the margin. The 16 inter-rhopalar radial-canals are narrower than the rhopalar,
and soon lose themselves in the network of anastomosing vessels of the subumbrella, whereas
the 16 rhopalar canals extend straight out to the sense-organs. The network of vessels becomes
fine-meshed on the inner side of the ring-canal, but on its outer side it gives off a wide-meshed
network, the meshes of which econ finer as they near the bell-margin, where there is a
marginal ring-canal. These hypothetical ring-canals are so peculiar and ‘unlike the simple net-
work seen in other species of C Jasstopea that the fact of their existence requires confirmation.
Haeckel alone has observed them. The medusa is from the Pelew Islands and New Guinea.

I find among the collections of the U. S. Fisheries Bureau steamer Albatross seven speci-
mens of a medusa which appears to be a closely related variety of, if not identical with, C.
ornata. None of these has the remarkable ring-canals figured by Haeckel, and this leads me
to doubt their existence in Haeckel’s medusa. The dimensions in mm. of the largest of these
medusz are as follows: Bell 76 wide; exumbrella flat, smooth and without an aboral sucker-
cavity; arm-disk 39 wide; mouth-arms 31 long, stout and flattened laterally, with g to 12 short,
stout, blunt, dentritically arranged side branches. A few very small, flat, club- like appendages
less than 1 mm. long scattered among the mouths of the mouth- -arms; bit these become larger
near the arm-disk, The arm-disk itself is thickly covered with irregularly shaped tuber-like, or
truffe-shaped, appendages, the largest being 3 to 4 mm. long. There are 16 rhopalia. 5X16
blunt, square-edged, marginal lappets. 32 tree-like radial-canals which give off an anasto-
mosing network, but no distinctly differentiated ring- -canal. These medusz were obtained in
the following localities in the Philippine Islands in 1908: 3 large specimens from near
shore at Tataan, Simaluc Islands, February 19 and 20; 3 from Subic Bay, January 7, in a
seine, and 1 from Catbalogan, Samar, on April 16.

Cassiopea ornata var. digitata Maas.

Cassiopea ornata var. digitata, Maas, 1093, Scyphomedusen der Siboga Expedition, Monog. 11, pp. 40, 45, taf. 4, fign. 26, 30.

Bell about 100 mm. wide, very flat, without an aboral concavity or a dome. 16 marginal
sense-organs, 32 rhopalar and 3X 16, or more, velar lappets which are, however, quite irregularly

RHIZOSTOM 8

649

arranged. 16 long, violet-colored, radial stripes upon the subumbrella. The mouth-arms
branch in a hand-shaped manner, the terminal branches resembling fingers in shape. These
mouth-arms are 1.5 times as long as the disk-radius. There are no large club-shaped appen-
dages between the mouths, all being very small. The mouths are brown to violet.

This variety is distinguished Gon the typical C. ornata by its color and by its finger-
shaped mouth-arms. It is found among the islands of the Malay Archipelago, at Saleyer,
and elsewhere.

Cassiopea depressa Haeckel.

Casstopea depressa, HarcKeL, 1880, Syst. der Medusen, p. 572.

Bell flat, shield-shaped, 100 to 120 mm. wide, 15 to 20 mm. high. Exumbrella smooth,
without aboral concavity or dome. 16 rhopalia, 144 wide, pointed, Bae not prominent lappets.
In each paramere 7 velar between 2 ocular lappets. 8 very wide, flat mouth-arms shorter
than the bell-radius and with 6 to 8 short, wide-spreading main-branches. Numerous very
small club-shaped vesicles between the mouths, hardly larger than the rhopalia, only 0.4 to
0.8 mm. long. No radial spots on the exumbrella.

Found at Madagascar and at the Querimba Islands off Mosambique, East Africa.
Described in detail by Haeckel.

Cassiopea depressa var. picta Vanhoffen.

Cassiopeia picta, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 26, taf. 2, fign. 1, 2.
Cassiopea depressa, var. picta, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 43.

Disk flat, 60 to 85 mm. wide. 14 (?) to 16 marginal sense-organs. 112 (716) velar
and 32 ocular lappets, all similar each to each, and blunt and small. The lappets are irregu-
larly developed in the two specimens described by Vanhoffen, and while there are usually
5 velar lappets between each successive pair of ocular lappets, in some parameres there are
0, 3, 8, or even 10 velar lappets. Arm-disk octagonal with 8 equal sides and half as wide as
bell-diameter. The free, projecting parts of the 8 mouth-arms are somewhat shorter than
the radius of the disk and project about one-third of their length beyond the bell-margin.
These mouth-arms are pinnately branched with short branches as in C. xamachana, and have
no appendages excepting small lancet-shaped ones, as in C. ndrosia and C. depressa. The
musculature is similar to that of C. ornata. The 32 radial-canals give off anastomosing side
branches which place them all in connection one with another. There are no distinctly
differentiated ring-canals.

Ground color of disk translucent opal. There are 16 large white spots over the 16 marginal
sense-organs, and in the large medusa these are fused into a ring of varying width, being
widest in the radi of the sense-organs and narrowest in intermediate positions. popu r
to this white ring are 144 white, ase oval streaks, one over each lappet; those over the 3
ocular lappets Smallest. those over the interocular lappets the longest. The 32 small a
over the ocular lappets are fused with the 16 large, white, radial spots. 16 white rays in the
subumbrella appear as large egg-shaped spots, their blunt ends inwards. They are in the
radi of the sense-organs aed ererd from the outer edges of the central stomach and gonads
outwards with their radial edges almost touching.

This species was Beeeped by Vanhoffen from two specimens found near Beibul in the
Red Sea, in December, 1884. It differs from other species of Cassropea in the large number
of its marginal lappets and its very wide arm-disk. ‘There is no raised central Scie on the
exumbrella and no large club- shaped vesicles on the mouth-arms, such as are seen in C.

polypoides.

Cassiopea mertensii Brandt.

Cassiopea mertensti, BranvT, 1838, Mém. Acad. Sci. St. Petersbourg., Sci. Nat., sér. 6, tome 4, p. 396, taf. 20-23.—Harcket,
1880, Syst. der Medusen, p. 572.
Cassiopeja mertensit, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 40.

Bell evenly rounded without an aboral concavity, 100 to 120 mm. wide, 30 to 40 mm.
high. 16 rhopalia. 128 small, tongue-shaped, prominently projecting lappets. In each
paramere 6 velar between 2 ocular lappets. 8 cylindrical mouth-arms 1.5 times as long a.

650 MEDUS OF THE WORLD.

bell-radius give off 8 to 12 main branches each, which also branch in a tree-like manner.
Numerous large club-shaped vesicles between the mouths, some half as long as bell-radius.
Bell yellowish, rusty-brown, lighter in the center. Radial streaks reddish-brown. There
are 2 white, half-moon-shaped spots over each rhopalium. Upper surfaces of mouth-arms
light-yellow. Mouths dark rusty-yellow. Vesicles white.
Found at Ualan, Caroline Islands, tropical Pacific.

Cassiopea ndrosia Agassiz and Mayer.

Cassiopea ndrosia, AGAssiz AND Mayer, 1899, Bull. Museum Comp. Zool. at Harvard College, vol. 32, p. 175, plate 14, figs.
Pena iaee var. ndrosia, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, pp- 40, 43-

Bell 50 mm. in diameter with a shallow concavity at the center of the exumbrella, similar
to that of C. xamachana. Rhopalia variable in number, 18 to 22. Marginal lappets very
indistinct ee there are 2 velar flanked by 2 ocular lappets in each paramere. Mouth-arms
cylindrical, 1.5 times as long as bell- radius, and branched in a tree-like manner. Each arm
gives off 6 to 12 main side (Mera nes. There are numerous small, flattened, expanded leaf-
shaped vesicles between the mouths, most numerous at center of arm-disk. No ribbon-
shaped filaments. 4 small, round, subgenital ostia. 4 separate genital sacs.

General color of bell grayish-brown, with bluish, inter-rhopalar, radiating streaks and
white radiations in the subumbrella in the rhopalar radu. A large, spearhead-shaped white
spot with its pointed end outward is found near the margin of the exumbrella above each
sense-organ; there are also 4 small, radially elongated, white spots near the margin in each
paramere—one above each of the rudimentary lappets. The aboral surfaces of the mouth-
arms are grayish-white, the mouths deep brown, and the vesicles olive-green.

Found upon muddy bottoms in Suva Harbor and at Komo Island, Fiji Islands, South
Pacific, in November.

C. ndrosta \acks the large vesicles of C. mertensi1 and has an aboral exumbrella concavity,
whereas the bell of C. mertensit.is, apparently, evenly rounded. It is most closely related to
C. xamachana of the West Indies, and resembles one of its color varieties, but lacks the ribbon-
like flaments of C. xamachana.

RHIZOSTOMATA DICHOTOMA Vanhiffen 1888.

Rhizostomata dichotoma, VANHGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 39.—Maas, 1903, Scyphomedusen der
Siboga Expedition., Monog. 11, p. 31.

Chaunostomide+ Cepheid, Craus, 1883, Organisation und Entwick. Medusen, Leipzig ——von LenpeNnFretp, 1888, Zeit. fiir
wissen. Zool., Bd. 47, p. 211.

Radiomyaria, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 89; 1907, Ergeb. und Fort. Zool., Bd. 1, p. 201.

CHARACTERS OF THE GROUP.

8 separate mouth-arms the lower ends of each one of which gives rise to 2 expanded,
leaf-like side-walls, or lateral membranes, the outer edges of which give rise to secondary
branches and bear the frilled mouths. Thus each arm is ye -shaped in cross-section (fig. 404).

f

Fic. 404.—Diagrammatic representation of the shape and position of the mouth-arms in the
Rhizostomata dichotoma. The figure on the right hand shows a section of one of the
mouth-arms. ‘The middle figure is an oral view of the bell.

RHIZOSTOMA2—CEPHEA. 651

There are no scapulets upon the mouth-arms. The radial-muscles are powerfully and the
ring muscles weakly developed. A description of the genera follows:

Cephea Péron anv Lesueur, 1809. Exumbrella with a central area bearing wart-shaped projections.

Cotylorhiza L. AGassiz, 1862. Exumbrella with a smooth central dome without wart-like elevations. Radial-canals
of the bell all similar each to each.

Polyrhiza L. Acassiz, 1862. Exumbrella with a central concavity and with radiating furrows.

Genus CEPHEA Péron and Lesueur, 1809.

Cephea, Péron er Lesurur, 1809, Annal du Mus. Hist. Nat., tome 14, p. 360.—Escuscuovtz, 1829, Syst. der Acalephen, p. 55.—
Lesson, 1843, Hist. Zooph. Acal., p. 410.—AGassiz, 1862, Cont. Nat. Hist. U. S., vol. 4, p. 155-—Harcker, 1880,
Syst. der Medusen, p. 573.-—VANHOFreEN, 1888, Bibliotheca Zoologica, Heft. 3, p. 39; 1902, Wissen. Ergeb. deutsch.
Tiefsee Exped. Valdivia, Bd. 3, Lfg. 1, p. 45-

Cephea+ Netrostoma, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, pp. 31; 325 35) 81, 89-

Netrostoma, Scuuttze, L. S., 1898, Denkschr. Med. Nat. Gesell. Jena., Bd. 8, p. 457.

Microstylus+ Perirhiza, KisHinovuyYe, 1902, Journal College Sci. Tokyo, Japan, vol. 17, Art. 7, pp. 11, 13-

Stylorhiza, HarcKer, 1880, Syst. der Medusen, p. 612.

Halipetasus, Scuuttze, 1898, Denkschr. Med. Nat. Gesell. Jena, Bd. 8, p. 458.

The oldest known species is ‘“‘ Medusa octostyla” of Forskal, and this may serve as the
type of the genus Cephea.

GENERIC CHARACTERS.

Rhizostomata dichotoma in which the 8 mouth-arms fork once dichotomously and each
fork gives rise to short dichotomous or dendritic branches. Solid, wart-shaped tubercles at
the center of the exumbrella. The central stomach gives rise to 8 rhopalar and numerous
inter-rhopalar radial-canals, all of which connect with a network of anastomosing vessels in
a wide zone near the margin. Rhopalia without ocelli and without exumbrella, sensory pits.
There is no definite ring-canal. Development unknown.

The described species of Cephea are all found in the tropical Indian Ocean and Pacific
region. Cephea cephea (Medusa cephea, Forskal) is apparently widely distributed over the
Indo-Pacific region and is distinguished by its numerous, long, tapering, conical, pointed
filaments; its deep rhopalar clefts in the bell-margin; oval velar lappets fused one to another
by a thin web, and its brown coloration. C. octostyla of the Red Sea—Malay Archipelago—
is distinguished by its very low exumbrella dome with very small tubercles. The marginal
lappets are indistinct. Also in C. comifera, C. dumokuroa, and C. cerulea the lappets are so
indistinct that the bell-margin is entire, save for the deep niches of the 8 rhopalia. In C. dumo-
kuroa and C. cwrulescens the central dome bears warts only on its sides, leaving its apex bare.
C. cwrulea has only 16 long filaments, whereas C. conifera has more than 100 and C. dumo-
kuroa none. In.Cephea cerulescens we find very small tubular and somewhat large spindle-
shaped filaments between the mouths, and the subgenital porticus is only partially differ-
entiated. In some quadrants the primitive genital sacs may have fused and the fused wall
broken down to form an opening, so that one may pass a probe into one subgenital ostia and
out through another without penetrating any tissue, the passage being continuous and actually
a part of the outside world. In other quadrants, however, the gonads may be quite separate one
from another or merely fused without any break in the area of fusion. In C. conifera and
C. dumokuroa, on the other hand, the subgenital porticus is unitary and cruciform, whereas
in C. setouchiana the 4 genital sacs are fused along their inner walls, but the walls remain
unbroken. In Cephea typhlodendrium the filaments are small and spindle-shaped, and con-
fined to the arm-disk.

It is evident that we have in the Red Sea, Indian Ocean, and western parts of the tropical
Pacific a large number of closely related forms of Cephea displaying many local variations.
I think there are only 2 well-marked forms and these are but the extremes of an intergrading
series: C. octostyla with low dome or flat exumbrella and small warts, and C. cephea with a
high dome and large warts.

L. S. Schultz, 1898, proposes a genus Netrostoma to include Rhizostoma dichotoma with
mouth-arms laterally compressed and several times dichotomously branched; no large fila-
ments between the mouths on the mouth-arms, although filaments may be found on the
mouth-arm-disk.

652 MEDUS OF THE WORLD.

As a matter of fact the mouth-arms of all known species of Cephea give rise to secondary
dichotomous, or dendritic, branches, and all are laterally compressed. Moreover, we can not
separate genera merely upon the relative size of the mouth-arm-filaments, for confusion is
certain to result.

“ Microstylus” of Kishinouye is evidently a Cephea closely allied to C. typhlodendrium.

I have therefore broadened the definition of the genus Cephea to include all Rhizostomata
dichotoma with a wart-bearing central area upon the exumbrella and with 8 forked mouth-
arms, the forms of which are themselves still further branched.

In the collection of Cepheas made by the U. S. Fish Commission steamer A/hatross in 1908,
I find an intergrading series among specimens of Cephea collected all at the same time on
the surface at Jolo, Philippine Islands. In some there is no central dome, the exumbrella
being quite flat, and in others there is a low but well defined dome. In some the exumbrella
warts are large and mammiform, while in others they are mere granules, often absent over
wide areas or leaving the center smooth. The filaments upon the mouth-arms and arm-disk
may be absent or long and filiform. Thus among these intergrading individuals (evidently
all of one and the same species) some are identical with Forskl’s “Medusa octostyla,” others
are similar to Schultze’s “‘Halipetasus scaber.”

Cephea octostyla L. Agassiz.

Medusa octostyla, Forskat, 1775, Descrip-Anim. Itin. Orient, p. 106, No. 18, Icon., tab. 29. af

Medusa cephea, Linné, (Gmelin) 1788, Systema Natura, Ed. 13, p. 3158. Non Medusa cephea, Forskal.

Cephea cyclophora, Mitne-Epwarps, 1849, Cuviér’s Régne animal illustré Zooph., planche 51, fig. 4.

Cephea octostyla, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 156.

Stylorhiza octostyla, HarcKet, 1880, Syst. der Medusen, p. 613.

(?) Stylorhiza polystyla, Harcket, 1880, Syst. der Medusen, p. 613.

Halipetasus scaber, Scuurtze, L. E., 1898, Denkschrift. Med. Nat. Gesell., Jena, Bd. 8, p. 458, taf. 33, fign. 5 und 7.

According to Haeckel and Forskal the bell is 300 mm. wide, flatter than a hemisphere.
Surface of exumbrella smooth, without a central dome, and without radiating furrows. 8
rhopalia 50 to 60 ( ?) wide, flatly-rounded, marginal lappets. Rhopalar clefts shallow. Arm-
disk wider than bell-radius. 4 small subgenital ostia. 8 bifurcated mouth-arms 1.25 times
as long as bell-radius; the forks of each arm as long as undivided upper part of arm. g long,
stout filaments of uniform caliber arise apparently from the arm-disk. These are 1.5 times
as long as diameter of bell and end simply, without terminal knobs. In addition to these
filaments there are about 12 short, stout, swollen, club-shaped appendages between the mouths
on the lower sides of the arms. The color is blue and hyaline. Arabian coast of the Red Sea.

Forskal’s description evidently refers to the medusa figured on his plate 29, not that of
plate 30 as stated by Niebuhr, who edited the plates of his work in 1776. While Forskal’s
figure is remarkably good for its period, his description is too brief and vague to be of value.
The description given above has therefore been mainly derived from a study of his figure.

Haeckel’s “Stylorhiza polystyla” from Singapore is described from a preserved and
damaged specimen. It is said to be 100 mm. wide with flatly rounded bell and 80 marginal
lappets. In each octant are 8 rectangular velar and 2 small oval ocular lappets. 4 subgenital
ostia hardly as wide as the columns between them. Mouth-arms nearly twice as long as bell-
radius; many times dichotomously branched, with 16 very long tubular appendages and
numerous vesicles upon long pedicels. 8 of the long appendages arise from the bases of the
arms and 8 from the crotches of the primary forks. This is probably identical with Forskal’s
medusa.

Many well-preserved specimens of Cephea octostyla were obtained by the U. S. Fisheries
Bureau steamer Albatross at Jolo Anchorage, Philippine Islands, tropical Pacific, in February
and March, 1908.

The bell of the large specimens is 90 mm. wide, exumbrella flat; rim vertical, 20 mm. high.
Neither central dome nor coronal furrow, but there is a zone of numerous, low, wart-like
protuberances upon the exumbrella, leaving the center free. This wart-covered zone is about
40 mm. in diameter and 15 mm. wide, leaving a circular area about 10 mm. in diameter free
of warts at the center of the exumbrella. Other parts of the exumbrella are smooth. There
are § rhopalia set within fairly deep niches. These lack ocelli and have no exumbrella sensory
pits. There are about 72 indistinct lappets, 7 velar and 2 ocular lappets in each octant. These

RHIZOSTOMA2—CEPHEBA, 653

lappets are similar each to each and are rectangular in outline, being separated by very slight
indentations which are spanned by a web. Fairly deep grooves extend up the vertical rim
of the exumbrella surface of the bell, between the lappets.

The arm-disk is as wide as the bell-radius and is 18 mm. thick. The free parts, upper,
of the 8 bifurcated mouth-arms are each about 20 mm. long. At the center of the arm-disk
we find 4 to 12 or more tapering, somewhat flattened, wart-covered filaments which are about
25 mm. long and terminate each in a simple, pointed end. There are also numerous shorter
filaments, ranging from about 15 to 5 mm. in length, all near the center of the arm-disk, and
many still shorter ones between the numerous frilled mouths of the 8 mouth-arms.

There are 4 small, oval, subgenital ostia only about 2 mm. wide. The ring-muscles of
the subumbrella are entire, but are very weakly developed. The central stomach gives rise
to 8 large rhopalar and about 56 (78) somewhat narrower radial-canals, all connected
one with another by a network of anastomosing vessels. The rhopalar canals proceed straight
through this network, giving off small side branches to the adjacent radial-canals, but the inter-
rhopalar canals tend to lose their identity in the network. There is no distinctly differentiated
ring-canal. There is a unitary, cruciform, genital cavity opening to the outer world by the
4 genital ostia. There are many fairly large, reddish-brown dots over the outer surface of
the arm-disk and reddish-brown streaks around the exumbrella warts. The color of the bell

Fic. 405.—Cephea octostyla. Drawn by the author, from specimens taken by the A/batross at Jolo Anchor-
age, Philippine Islands, in February and March, 1go8.

A, B, and C, side views of exumbrella, showing variations in development of warts; D, rhopalium seen
from subumbrella side.

is faded in formalin, but a color note states that the exumbrella bore numerous small, round,
ocherous spots. When the bell is 15 mm. wide there are a few, small, scattered warts near the
center of the exumbrella. There are 8 rhopalia set within shallow niches. The bell-margin
between the sense-clubs is entire and there are no lappets. There are 8 quite wide rhopalar
and 8 X 3 somewhat narrower inter-rhopalar canals, all set into communication by side branches
forming a marginal network. The 4 genital cavities are separate. The 8 bifurcated mouth-
arms lack appendages either upon the arm-disk or between the frilled mouths. This young
specimen was caught upon the surface under an electric light in Jolo Anchorage, Philippine

Islands, on February 13, 1908.

Cephea octostyla var. ccerulescens Maas.

Netrostoma cerulescens, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 35, taf. 5, fign. 37, 46; taf. 11, fign.
97, 103; taf. 12, fig. 109; 1906, Revue Suisse de Zool., tome 14, p. 101.—Browne, 1905, Fauna and Geog. Maldive and
Laccadive Archipelagoes, vol. 2, p. 967.

The bell becomes at least 200 mm. wide. There are 8 marginal sense-clubs, each with a
terminal entodermal concretion-mass and without an exumbrella sensory pit. No ocelli(?) In
other respects the bell resembles that of Cephea octostyla having a central dome which bears
about 10 wart-like projections. There are 6 to 8 round-edged, marginal lappets in each
octant. The 8 mouth-arms are short, massive, laterally compressed, and curved outwards.

654 MEDUS.2 OF THE WORLD.

Each mouth-arm bifurcates at its outer end and also gives rise to numerous very short, lateral
branches upon its lower side. These side branches themselves branch somewhat dichoto-
mously, giving a complex system of mouth-bearing ramuli upon the lower side of the mouth-
arm. ‘There are two sorts of appendages between the mouths: small, thin, tubular appendages
with prominent nettling-warts and larger, spindle-shaped appendages. ‘These are, however,
very small in comparison with
the size of the branches of the
arms themselves.

In young medusz there are 4
separate, subgenital cavities with
4 small, external ostia in the inter-
radial sides of the arm-disk. In
older individuals we find a very
variable condition, the different
quadrants of the same medusa
being unlike; but it seems that
a completely separated, subgenital
porticus, such as that found in
Crambessa or Cotylorhiza, is never
formed in Cephea caerulescens.
The canal-system of the umbrella
consists of 8 radial-canals in the
radi of the 8 marginal sense-
organs and 24 intermediate canals
which give rise to numerous side
branches, forming a network of
canals which place all 32 vessels
in communication one with an-
other. There is no distinctly
differentiated, annular ring-canal.
The muscular system of the sub-
umbrella resembles Cotylorhiza in
the form of the radial-muscle
strands. The marginal ring-mus-
cles are, however, very poorly
developed.

The general color appears to
be blue. A narrow zigzag band
of fiery red, around the outer side
of the arm-disk, lies above the
subgenital ostia and bends down-
ward (outward) at each interra-
: dius toward the subgenital ostium.

= Clusters of small, brown, oval
Fic, 406.—* Perirhiza nematophora’’= Cephea cephea, after Kishinouye, in spots are found near the side of
Journal College of Science Tokyo. : :
: : = each subgenital ostium.

This medusa is found in the Malay Archipelago and Maldive Islands, Indian Ocean.
The reddish dots found in the typical C. octostyla around the sides of the arm-disk have, in
this variety, fused into a solid band of color.

AN
NN
i)

SS
SN
3
SSE
SS =

Cephea cephea.

Medusa cephea, Forsxat, 1775, Descript. Anim. Itin. Orient., p. 108, No. 22, Icon., tab. 30 (Non. Tabl. 29).
Medusa octostyla, Linnt, (Gmelin), 1788, Systema Natura, Ed. 13, Pars. 6, p. 3157-

Cephea rhizostomoidea, Pixon ex Lesueur, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 361, No. 100..
Polyrhiza cephea+ Diplopilus couthouyi, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp- 156, 158.

Cephea forskalea+ C. conifera, Haecxen, 1880, Syst. der Medusen, PP- 574, 576, taf: 36, fign. 3-6.

(?) Cephea fusca, Péron ev Lesurur, 1809, loc. cit., p. 361, No. 99.

(?) Casstopea fusca, Dusimurr, 1835, Musée du Jardin des Plantes, No. 111.

Perirhiza nematophora, Kiswinouye, 1902, Journal College Sci. Tokyo, vol. 17, Art. 7, p. 14, plate 2, figs. 11-13.

RHIZOSTOM-2—CEPHEA, 655

Forskal gives a good, clear drawing of this medusa, which bears so close a resemblance
to the figures of “‘ Perirhiza nematophora” of Kishinouye that I am convinced the two are
identical. The medusa is distinguished by the very deep rhopalar clefts in the bell-margin,
its long tapering mouth-arm-flaments, and brown color. Gmelin erred in calling this M.
octostyla, when he quoted from Forskal, for the latter’s Medusa octostyla is very different.

Bell 100 to 140 mm. in diameter. A large dome at apex of exumbrella, nearly as wide
as bell-radius and covered completely with about 30 large, conical, pointed warts, many of
which are bent near their pointed ends. The dome is surrounded by a wide, shallow ring-
furrow, which separates it from the nearly equally wide, flexible marginal zone of the bell.
The 8 sense-organs are set within deep niches in the bell-margin, as is well shown in Forskal’s
figure. There are 80 to go marginal lappets; in each octant 8 or g large, oval, velar between
2 very small, pointed, ocular lappets; the velar lappets are united by a web, so that the bell-
margin appears to be nearly entire. The small ocular lappets are deeply set inward centrip-
etal to the margin.

On the subumbrella a radiating inner zone of folded ridges contains the radial muscles,
and near the bell-margin is an unbroken zone of circular muscles. Arm-disk octagonal,
nearly as wide as bell-radius. The 4 subgenital ostia are very small, compressed clefts. There
is a unitary, cruciform, subgenital cavity. The arm-disk has no canal-system of its own, and
there are no mouths upon its central parts. The 8 laterally compressed, stout, adradial
mouth-arms are somewhat shorter than the bell-radius. Their upper halves are nearly
coalesced where they arise from the arm-disk, but below they fork and each main branch
branches profusely and curves upward. The numerous, frilled mouths are found on the lower,
ventral sides of these mouth-arms and their branches. There are more than 100 long, tapering,
hollow flaments with pointed ends. The largest of these arise from the arm-disk at the points
of origin of the 8 mouth-arms, and they are as long as the diameter of the umbrella and
hollow. Forskal figures 16 such filaments all apparently arising from the arm-disk and
numerous smaller ones arising from between the mouth-frills on the arms, very much as does
Kishinouye 127 years later.

The nearly circular, central stomach gives rise to 8 ocular and about 40 to 48 interocular
radial-canals. The ocular canals are not wider than the others, but they extend straight out
to the rhopalia, giving off numerous side branches into the network-zone of the bell; whereas
the interocular canals lose themselves in this wide network of anas‘omosing vessels which form
a broad zone extending from near the outer edge of the stomach-cavity to the bell-margin.
There is no differentiated ring-canal. The network gives off many blindly-ending branches
which extend downward into the radiating muscular ridges of the subumbrella.

The margins of the velar lobes are brown, but Kishinouye finds that other parts are
colorless, although Forskal’s medusa displayed some reddish-brown on its bell. Forskal
describes this medusa from the Red Sea, and Kishinouye from Misaki, Japan, where it is
found in winter. Péron and Lesueur’s C. fusca, from Malabar and northwesrern Australia, is
probably the same medusa; as is also Diplopilus couthouyt Agassiz, 1862 (Cont. Nat. Hist.
U.S., vol. 4, p. 158), from Hawaii. The medusa appears to be widely distributed over the
Indo-Pacific region. Haeckel’s Cephea conifera from Samoa may be another name for the
same medusa, but its color is not stated and 1:s marginal lappets appear to be indistinct, and
the bell-margin to be practically entire, as in C. caerulea. The decided resemblance, 1n other
respects, between Haeckel’s C. conifera and Forskal’s medusa will appear in the following
description.

“Cephea cephea var. conifera” Haeckel.

Cephea conifera, Harcket, 1880, Syst. der Medusen, p. 576, taf. 36, fign. 3-6.—Hamann, 1881, Jena. Zeit. fiir Naturw., Bd. 15,
p- 246 (anatomy of mouth-arms).

This is probably identical with Cephea cephea.

Bell 100 to 120 mm. wide, 30 to 40 mm. high. A thick-walled, flatly rounded, central
dome upon the exumbrella bears 20 to 30 large and numerous small protuberances and is
separated from the marginal zone of the exumbrella by a deep annular furrow. These solid,
wart-like protuberances of the central dome are scattered irregularly over its entire surface,
as in C. cerulea, not arranged in 2 rows, as in C. dumokuroa. 8 rhopalia are set within deep
niches. 80 indistinctly developed, marginal lappets. In each octant 8 wide, flat, velar lappets,

656 MEDUSA OF THE. WORLD.

flanked by 2 very small, rhopalar lappets. The lappets are so poorly developed that the bell-
margin is practically entire and without notches, in this respect resembling C. cerulea. The
octangular arm-disk is about as wide as the bell-radius, and the 4 subgenital ostia on its
interradial sides are very short, narrow clefts.

The 8 mouth-arms arise in 4 pairs from the perradial angles of the central part of the
arm-disk, but separate widely, one from another, so as to project from the 8 adradial corners
of the sides of the disk. The 8 mouth-arms are each about as long as the bell-radius and
each one bifurcates near its outer end. Numerous short branches arise from the ventral
sides of the mouth-arms and these bear the mouths. A single long, stout filament arises from
each of the 4 perradial corners of the ventral side of the arm-disk at the points of bifurcation
of the 4 primary mouth-arm canals. In this respect the medusa differs from C. ce@rulea,
wherein there are 4 filaments in each perradius of the arm-disk. There are also more than
100 long, slender hlaments between the mouths. These filaments are longer than the bell-
diameter. In the closely allied C. cwrulea the mouth-arm filaments are very short and incon-
spicuous. ‘The radial-muscles of the subumbrella are well-developed and form radiating,

lamella-like ndges as in C,carulea.

Vy £0 ‘There are numerous, fine, anasto-
\ . . .
5 2 = em! mosing radial-canals and a wide
nates ay mye eae
> RN ie fiat TERA on network of vessels near the mar-
; PRCA 6} tI as

Fx Bayh G

s
ae eA TIES gin. Color (?)

Found at the Caroline and
Samoan Islands, tropical Pacific.
This description is presented to
show that there are no appreciable
differences between this medusa

and Forskal’s Medusa cephea.

Cephea cephea var. dumokuroa Agassiz
and Mayer.

Cephea dumokuroa, AGassiz AND Mayer, 1899,
Bull. Museum Comp. Zool. at Harvard
College, vol. 32, p. 172, plates 11, 12,

Q Sy =
saath ar EN figs. 36-39.
Mates tah: Netrostoma dumokuroa, Maas, 1903, Scypho-
‘ he 2 medusen der Siboga Expedition, Monog.
ee II, p- 38.

Bell 300 mm. wide, flat, and
disk-shaped with sides vertical
near the margin. A large promi-
nent domeat center of exumbrella.
The apex of this dome is smooth and without the wart-like protuberances seen in C. caerulea
and C. contfera; instead of which the protuberances of C. dumokuroa are arranged in two ver-
ticels, confined to the sides of the dome. The upper row of protuberances consists of about
8 large, solid, wart-like, bluntly-pointed projections, and below them is a zone of about 12
smaller warts not more than half as large as those of the upper row. There is a wide, shallow
furrow around the dome. 8 rhopalia are deeply sunken within marginal niches. Each sense-
organ contains a terminal mass of white, entodermal concretions. There is no ocellus and
no exumbrella sensory pit.

The marginal lappets are so shallow that one can barely distinguish them, but there are 8
scarcely perceptible, velar lappets in each octant, as in C. cerulea and C. conifera. The arm-
disk is about as wide as the bell-radius and there are 4 very small, round, subgenital ostia, with
a unitary subgenital porticus. 8 short, bifurcated mouth-arms, each about as long as bell-
radius; their free outer ends curve upward and the mouths are confined to the ventral sides of
the arms, the frilled mouths being placed upon short branches which arise from the lower side
of each arm. There are neither filaments nor club-shaped appendages, and in this respect the
medusa differs from C. cwrulea and C. conifera. The central stomach is a wide cruciform space
above the subgenital porticus. 32 radial-canals arise from its margin and diverge into the sub-

Fic. 407.—“ Cephea conifera,” after Haeckel, in Das Syst. der Medusen.

RHIZOSTOM-8—CEPHEA, 657
umbrella. 8 of these canals lead to the rhopalia and 3 are in each inter-rhopalar octant, instead
of 7 as in C. cearulea. There is a wide zone of anastomosing vessels near the margin. The
central stomach also sends 8 canals downward into the arm-disk and this in turn to the 8
mouth-arms.

General color of medusa blue, as in C. cwrulea. The bare apex of the central dome is
streaked longitudinally with blue, and a deep blue entodermal band of colorextends around
the sides of the arm-disk above the subgenital ostia. This blue band is especially wide in
the adradiit above the bases of the 8 mouthe -arms. The unbranched portions of the radial-
canals adjacent to the stomach and the canals of the arm-disk and arms are blue. The
broad network of anastomosing vessels near the bell-margin is dull coffee-colored and the
frilled mouths are of a deeper hue of the same color.

A large swarm of these medusz was found upon the surface off Vanua Mbalavu Island,

Fiji Islands, on November 25, 1897.

Cephea cephea var. ccerulea Vanhoffen.

Cephea cerulea, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped. Valdivia, Bd. 3, Lfg. 1, p. 45, taf. 4, fign. 13, 14.

Bell 57 mm. wide. A dome-like apex 14 mm. wide at the center of the exumbrella is
surrounded by an annular furrow g mm. wide. The dome itself bears 6 large and about 30
small, rounded, wart-like protuberances, the largest of which are 2.4 to 5 mm. wide; numerous
fine punctations between the warts. There are 8 rhopalia. No marginal lappets, but 8 radial
thickenings of the gelatinous substance at the margin in each octant. The 4 subgenital ostia
are very small. The dichotomous mouth-arms are seach 16 mm. long and bear frilled mouths
on their ventral sides. There are 4 long filaments at the point of bifurcation of each of the
4 primary arm-canals, 16 in all; of these, the second and third are 2 to 3 times longer and
much stouter than the first and fourth. Numerous very small filaments are among the mouths.
The central stomach gives rise to 64 radial-canals, which break up into a wide, anastomosing
network zone near the bell-margin. There are 8 rhopalar radial-canals and 7 inter-rhopalar
canals in each octant thus differing from C. dumokuroa, wherein there are only 3 radial-canals
in each inter-rhopalar sector. There is a broad, marginal annulus of ring-muscles in the
subumbrella, and centripetal to this is a wide zone of radial-muscles extending from the
supports of the arm-disk to the zone of ring-muscles.

The disk, arms, and filaments are blue and the frilled-mouths are chocolate-brown.

Found at Dar es Salaam, east coast of Africa (see fig. 408).

Cephea cephea var. setouchiana.

Microstylus setouchtanus, Kisuinouye, 1902, Journal College Sci. Tokyo, vol. 17, Art. 7, p. 11, plates 1, 2, figs. 8-10.
Netrostoma setouchtanus, BRowNe, 1905, Fauna and Geog. Maldive and Laccadive Archipelagoes; vol. 2, p. 967.

Disk 100 to 200 mm. wide with a prominent central dome nearly as wide as bell-radius
and covered completely by 50 or more solid, pointed, wart-like projections of various sizes.
This central dome is surrounded by a wide, annular furrow which separates it from the thin,
flexible, outer zone of the exumbrella. 8 thopalia. 6 to 8 flatly rounded, velar lappets flanked
by 2 smaller, rounded, rhopalar lappets in each octant. 50 to 60 lappets, in all, upon the bell-
margin. The 8 mouth-arms curve outwardly and upwardly in their lower halves. They are
a little longer than the bell-radius and each one is forked, each of the main forks being nearly
as long as the upper, undivided part of the arm. The forks give mse in turn to numerous
pinnate branchlets. There are numerous small, short appendages among the frilled mouths,
and those on the arm-disk at the ends of the perradial oral suture are longer, triangular in
cross-section, and prickly in appearance. The 4 subgenital ostia are circular and much
narrower than the spaces between them. The subgenital cavity is unitary and 4-lobed, as
in Cotylorhiza.

The central stomach gives rise to 8 large thopalar and 24 inter-rhopalar radial-canals,
all of which extend straight Soueward to the bell- margin. All of the canals give off side branches
which form a wide, anastomosing network of esccla! the meshes of ech are mainly polyg-
onal near the center, but rectangular near margin of disk. 8 canals arise from the stomach
at depressed areas near the sides of the perradial septa of the subgenital cavity. These canals

65S MEDUSA OF THE WORLD.

extend downward into the 8 mouth-arms and each give off a horizontal branch which
extends into the center of the arm-disk where they unite in a single short duct.

Fach wart of the central dome is pigmented with lines of numerous, minute, brown dots
converging toward the pointed apex of the wart. There are also brown dots on the sides of
the arm-disk. The oral frills are brown and the gonads pinkish.

Found in August and September in the Inland Sea, and at Misaki and Senzaki, Japan.

A shrimp is commensal with the medusa, hence its popular Japanese name “Yebi-
kuragé” (shrimp medusa).

Cephea typhlodendrium.

Netrostoma typhlodendrium, Scuurtze, L. S., 1898, Denkschr. Med. Nat. Ges. Jena., Bd. 8, p. 457, taf. 34, fign. 10-12a.—Maas,
1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 38.

Bell flatly rounded, 110 mm. wide. The center of the exumbrella is occupied by a low
dome completely covered with about 80 rounded warts of various sizes. There is no ring-
furrow around the dome. The outer parts of the exumbrella are smooth. 8 marginal sense-
organs. 80 marginal lappets composed of 64 rounded or cleft, velar lappets and 16 narrow
sharp-pointed ocular lappets. The 8 mouth-arms arise from a thick arm-disk. The mouth-
arms branch dichotomously and are laterally compressed. Small, spindle-shaped, sharp-
pointed filaments are found only on the arm-disk. There are 4 very small, round, subgenital

Fic. 408.—Cephea cerulea, after Vanhoffen in Valdivia Expedition.
Fic. 409.—Cephea setouchiana, Kishinouye, in Journal College of Science, Tokyo.
ostia and a unitary, subgenital porticus. The stomach gives rise to 32 radial-canals: 8 ocular,
24 interocular. The intérocular canals give rise, distally, to blindly-ending side branches,
peripherally to anastomosing branches; but the 8 ocular canals give off only the peripheral,
anastomosing vessels. ‘These ocular canals are wider than the interocular and extend straight
through the marginal network to the 8 rhopalia, whereas the interocular vessels become lost
in the peripheral network. There is no definite ring-canal.
Found at Amboina, Molluccas, Malay Archipelago.
This may be a well-defined species, for it appears to be distinguished from other members
of the genus by its small, sharp-pointed, ocular lappets.

Genus COTYLORHIZA L. Agassiz, 1862.

Cotylorhiza, AGassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 158.—Harcket, 1880, Syst. der Medusen, p. 609.—Caus, 1883,
Untersuch. Organisation und Entwick. Medusen, p. 60, Leipzig.—VANuGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3,
PP 27, 40.—Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, pp. 32, 80, 89.

: The type species is the well-known Cotylorhiza tuberculata (Medusa tuberculata Macri)
of the Mediterranean.

GENERIC CHARACTERS.

Rhizostomata dichotoma with 8 simple, bifurcated mouth-arms, the terminal branches
of which branch pinnately. The 4 subgenital ostia are simple and funnel-shaped, and there

RHIZOSTOMA2—COTYLORHIZA, 659

is a single subgenital porticus. The appendages upon the mouth-arms are mounted upon
pedunculated filaments. There are 8 marginal sense- -organs and numerous radial-canals
which anastomose laterally without any dennite ring-c ral in the adult. The sense-clubs
have no ocelli and no exumbrella sensory pit. There is a unitary peripheral zone of circular
muscles and an inner zone of radial-muscles in the subumbrella. The exumbrella is smooth
and without an aboral “‘sucker-like” depression, but with a prominent central dome without
wart-shaped elevations upon it.

This genus is sharply separated from Cassiopea, with which it has often been confused,
by its single, unitary, subgenital porticus, its relatively simple bifurcated mouth-arms, and By
having constantly 8 igteadl of an indefinite number of marginal sense-organs. Also there is no
abort ‘sucker” upon the exumbrella, such as is commonly seen in Cassiopea.

Cotylorhiza tuberculata L. Agassiz.
Plate 73, fig. 2

Medusa tuberculata, Macri, G., 1778, Osservazioni Int. Polmone Marino, p. 20.—Linné, (Gmelin), 1788, Systema Natura,
Ed. 13, Pars. 6, p. 3155-

Medusa tuber, Macrt, S., 1825, Atti Reale Acad. Sci. Napoli, vol. 2, Parte 2, p. 74, tav. 4, figs. 1, 2.

Cephea polychroma, PEron rr Lesurur, 180g, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 361—Lamarck, 1816, Hist. Anim.
sans Vert., tome 2, p. 516.

Cassiopea borbonica, Dette Cuiajr, 1823, Memorie sulla storia e notomia degli animali senza vertebre, Napoli, Mem. 3, p- 75,
tav. 3, 4, figs. 1-6; 1841, Animali senza vert., Napoli, tome 7, tav. 140-141.—Kowateysky, 1873, Mem. Imp. Soc. Friends
of Nat. Hist., Moscow, vol. 10, part 2, p. 3, plate 2 (Russian).—Du Pressis, 1881, Bull. Soc. Vaud. Sci. Nat., vol. 17,
No. 86, p. 633, plate 31 (development of egg-scyphostoma).—Dr Merrykowsky, 1882, Archiv. Zool. Exper. et Générale,
tome 10, p.577,planche 29B, figs. 14-20 (development of spermatozoa).—Craus, 1883, Arbeit. Zool. Inst. Wien, Bd. 5, p. 169,
2 taf., fign. 1-11 (development of the ephyra); 1883, Organisation und Entwick. der Medusen, pp. 43, 53, taf. 15, fign.
106-111 (ephyra); 1890, Arbeit. Zool. Inst. Wien, Bd. 9, p. 85, taf. 4-6; 1890, Verhandl. Zool. Buin: Gesell. Wien, Bd. 40,
pp- 54-55 (development of the scyphostoma larva); 1892, Arbeit. Zool. Inst. Wien., Bd. 10, p. 1 (scyphostoma).—Grarrrr,
1884, Arbeit. Zool. Inst. Wien, Bd. 5, p. 343 (seasonal distribution).—Ketrer, 1884, Recueil Zool. Suisse, tome 1, p. 403
(breeding habits of the medusa, and nature of the “yellow cells”).—Vanuorren, 1888, Bibliotheca Zoologica, Bd. 1, Heft 3,
pp: 27, 40 (in the Red Sea).—Hessr, 1895, Zeit. fiir wissen. Zool., Bd. 60, p. 441, taf. 21, fign. 17-19; taf. 22, fign. 27, 28
(nervous system of the subumbrella).—Biceow, R. P., 1900, Mem. Boston Soc. Nat. Hist., vol. 5, No. 6, p. 209.—Hein,
1902, Zool. Anzeiger, Bd. 25, p. 637; 1902, Zeit. fiir wissen. Zool., Bd. 73, p. 302, taf. 20, 21 (development of the gastrula).—
Betxre, 1903, Allgemeine Anat. und Physiol. Nervensystem, pp. 410, 414, 448, etc., fig. 83, 84, 88, 91, etc. (physiology of
the nervous system and of pulsation).—Maas, 1904, Résult. Camp. Sci. Prince de Monaco, fasc. 28, p. 58, planche 2,
figs. 16-19; plate 6, fig. 47; 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 32, taf. 8, fign. 69, 70.—Bouvier,
1907, Bull. Instit. Océanograph, Prince de Monoco, No. 93.

(?) Casstopea corolliflora+ C. canariensis, Tirestus, 1829, Nova Acta. phys. med. N. C., tome 15, pp. 265, 285, tab. 73.

Cephea tuberculata, Escuscuoxtz, 1829, Syst. der Acal., p. 56.

Cephea wageneri, Witt, 1884, Hore Tergestina, p. 58.—Buscu, 1851, Anat. Entwickl. wirbell. Seeth., p. 30, taf. 2, 3.—FRrantzivs,
1852, Zeit. fiir wissen. Zool., Bd. 4, p. 118, taf. 8, fign. 1-4 (siphostoma larva).

Cassiopea borbonica, GeGENBAUR, 1854, Generationswechsel Medusen, p. 2, taf. 2, fign. 32-35.

Cotylorhiza tuberculata, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 158 —Hamann, 1881, Jena. Zeit. fiir Naturwissen,
Bd. 15, p- 254, taf. 9, fign. 8, 9; taf. 10, fign. 19-23, 34, 353 taf. 11, fign. 24-28 (structure of the mouth-arms).

Cotylorhiza tuberculata, and C. ambulacrata (?) Harcket, 1880, Syst. der Medusen, pp. 610, 611.

Cotylorhiza borbonica, Gorter, 1885, Zool. Anzeiger, Bd. 8, p. 554; 1887, Abhandl. Entwick. der Thiere, Heft. 4, Leipzig, 79 pp.;
1893, Zeit. fiir wissen. Zool., Bd. 55, p. 645, taf. 28, 29 (embryology).

Bell usually not more than 150 to 170 mm. in diameter, though according to Will it may
become 300 mm. wide. The dimensions of a large medusa found at Naples on December 28,
1907, were as follows: Bell 168 mm. wide, arm-disk 88 mm. wide, each mouth-arm 46 mm.
long and 54 mm. thick (dorso-ventrally). The gelatinous substance is very rigid. The exum-
brella surface is smooth and without wart-like projections.

The center of the exumbrella is occupied by a smooth, elevated dome, somewhat flatter
than a hemisphere and about as wide as radius of disk. Around the outer edge of this dome
is a depressed region forming a gutter-like ring, somewhat lower than the parts of the bell
nearer the margin. There are 8 marginal sense-organs, which lack an exumbrella sensory
pit and have no ocelli. The sense-club has a very large, bag-like ventral bulb and contains
a terminal mass of orange-colored entodermal concretions.

The marginal sense-organs are flanked by 16 short, blunt, oval rhopalarlappets. There
are typically 10 velar lappets in each octant, the middle 6 of which are sometimes, but not
always, cleft. The outer edges of these lappets are subrectangular with bluntly-rounded
angles. The primary clefts between the lappets are fully twice as long as the secondary clefts
of the 6 middle lappets; but all the clefts are bridged over by a web of subumbrella tissue,
so that they are not complete cuts, but mere grooves upon the exumbrella. ‘The 16 velar

660 MEDUS.2 OF THE WORLD.

lappets adjacent to the 16 rhopalar lappets are fully twice as wide as the others and are rarely
cleft. The actual number of marginal lappets is quite variable, but we may say that there are
typically 96 primary lappets, of which 48 are typically cleft. Counting these clefts we would
then have 18 lappets per octant or 144 1n all. The bell-margin usually bends at right angles
to the general surface of the exumbrella; subumbrella surface convex.

Arm-disk octangular with re-entrant angles and sharply set off from subumbrella. It
is thick and about as wide as radius of bell. Thus in a medusa 168 mm. in diameter the arm-
disk was 88 mm. wide. 4 simple, small, oval, subgenital ostia on interradial sides of arm-disk
are not much wider than the width of a marginal lappet. The 8 adradial mouth-arms are
bluntly simitar-shaped in general outline, are laterally compressed, and only about half as

Fic. 410.—Cotylorhiza tuberculata, from life, by the author, at Naples Zoological Station, December, 1907.

A, oral view with all but one of the mouth-arms removed. The muscular layer is also removed
over the area on to left side of the figure in order to show the canal-system. B, section
through medusa showing central stomach and unitary subgenital space below it. C, cross-
section through subgenital space (sparsely dotted) and stomach (with thickly placed linear
dots); showing the 8 ostia of the mouth-arm canals. D, mouth-arm cut off from arm-disk
and viewed from cut end. E, club-like appendages among the frilled mouths. F, sense-
organ seen from exumbrella side.

long as bell-radius. They are somewhat thicker (downward) than wide and arise from the
arm-disk at 45° apart. Thus in a medusa 168 mm. in diameter the mouth-arms were each
46 mm. long and 54 mm. thick. Each mouth-arm bifurcates near its base and each of the two
main branches gives rise to about 10 to 14 side branches, which in turn give off 30 to 40 smaller
branches, and these again to 100 to 150 smaller branches, which branch still further dendritic-
ally. The farther out the branches the more dendritic and the less dichotomous is their mode
of branching.

A large number of short, club-shaped appendages between the frilled mouths terminate
in bluntly conical to flatly expanded, disk-like ends covered with small, wart-like tubercles.

RHIZOSTOM Aa

LORHIZA. 661

Besides these short appendages, less numerous but somewhat larger ones terminate in a
flattened ball-like to disk-like end set in a socket. At the point of bifurcation of each of the
8 main mouth-arms is a flament which is circular in cross-section and nearly half as long as
the mouth-arm itself. This filament tapers gradually from base to tip, but usually terminates
in a swollen end. Centripetal to these 8 main filaments are 3 to 5 other, somewhat shorter
filaments arising between the frilled mouths of each mouth-arm. Near and at the center of the
arm-disk there are numerous slender filaments, about one-third as long as the main filament,
which terminate in expanded disk-like conical ends. :

A unitary, cruciform, subgenital space opens outwardly at the 4 subgenital ostia. Thus
it is possible to pass a probe into any one of the subgenital ostia and out through the one
on the opposite side of the arm- -disk without penetrating any tissues of the medusa; the
subgenital space being actually outer world (C, fig. 410). The complexly folded, genital
ane 1s developed upon the sides and upper floor of this subgenital space, and thus
upon the lower floor of the central stomach.

The central stomach is large, occupying the spacious cavity of the central dome of the
exumbrella (B, fig. 401). 11 to 13 radial-canals per octant (88 to 124 in all) arise from the
margin of this central stomach and extend outward to the bell- -margin. There is no dis-
tinct ring-canal, but instead there are numerous, lateral anastomoses sbefreen the radiating
canals, forming a complex network of vessels under the floor of the subumbrella. The main
canals of the 8 mouth-arms empty by 8 adradial openings into the central stomach. Each
of these mouth-arm canals bifurcates, and the two main branches give rise to numerous,
lateral diverticula which lead to the gutters of the frilled mouths.

There is a well-developed unitary, marginal ring of circular muscles and an inner zone
of radial-muscles in the subumbrella. Bethe, 1903, has shown that when the medusa pulsates
these inner-lying radial-muscles contract before the ring-muscles, though the latter lie nearer
to the sense-organs, from which the contraction-impulse arises. This more rapid response
of the radial-muscles is due to the fact that thezr latent interval between the reception of the
stimulus and their response is less than in the case of the circular muscles. Bethe also finds
that the normal pulsation consists of 80 to 100 contractions at fairly regular intervals with
periods of total rest between them. These are then followed by a pause Sohich lasts as long
as 3 to 20 of the normal pulsations. For further details of the character of the pulsation, the
reader should refer to Rhizostoma pulmo.

The bell of this medusa is rich olive, tending to orange, or to brownish-yellow, being
especially dark and brownish on dome-like apex of the exumbrella. The rich yellow color
is found on both exumbrella and subumbrella, and is due to the presence of numerous yellow
to brown plant cells (Zoochlorelle) which float in the canal-system and infest the entoderm
of the medusa. Claus, 1883, finds these cells in the 8 lobed ephyra when only 1.5 to 2 mm.
wide.

The arm-disk and mouth-arms are usually pale milky-white tinged with delicate creamy-
yellow. The free outer edges of the mouths are tinged with purple « varying to blue or violet.
The terminal portions of dhe appendages, which arise between the frilled “mouths, are deep
blue tending to purple, or violet.

This medusa is found in the Mediterranean, but is quite capricious in its appearance,
being at times very rare. According to Graeffe it is not seen every year in the Adriatic at
Teveste, but adult medusz are usually: seen from July to September, w hile small ones are found
in July and August. At Naples adult medusz are commoner from August to October than in
winter, when they become very rare, being only occasionally found in mid-winter. Keller is
of the opinion that this medusa is a deep-water species which only occasionally comes to the
surface when sexually mature, and that the young remain near the bottom of the sea. Van-
hoffen, 1888, records the capture of a young individual of this medusa at Assab in the Red
Sea on June 10, 1885. The medusa must have been introduced into the Red Sea through the
Suez Canal. It has also been found in the Atlantic, near the Canary Islands. (See C. ambu-
lacrata Haeckel. )

The development of this medusa has been studied by Busch, Frantzius, Gegenbaur,
Kowalevsky, Claus, Goette, du Plessis, Hein, and others, and has furnished some of the
evidence for a controversy between Claus and Goette concerning the development of the gastro-
vascular cavity of the scyphostoma.

662 MEDUS.© OF THE WORLD.

The young larve are set free from the mouths of the mother medusa as planula or young
eastrule. Segmentation is total and nearly equal. The gastrula is formed by invagination
as in the case of Aurellia. The free-swimming, planula is pyriform to oval, flattened laterally,
and ciliated externally. The entoderm of the planula is entirely encased by the ectoderm
through the closure of the blastopore, and thus the planula is a two-layered sac which attaches
itself to the bottom by means of its broad anterior end, and then loses its cilia.

An invagination of the ectoderm takes place at the posterior (now uppermost) end of the
planula. According to Goette the entoderm is also evaginated at the same time in such manner
that two backwardly projecting pouches remain in the plane of the wide lateral diameter,
while these pouches are absent on the flat sides of the larva. The ectodermal invagination
forms the mouth and cesophagus; while the entodermal evaginations form the first pair of
lateral stomach-pouches. An opening is soon formed where the inyaginated ectoderm has
fused with the entoderm, and thus the throat-tube is placed in communication with the central
stomach. The second pair of gastric pouches now arise go° apart from the first and, according
to Goette,are produced by evagination entirely from the ectoderm of the lower end of the
throat-tube,

Hyde, 1894 (Zeit. fiir wissen. Zool., Bd. 58, p. 521), finds, however, that in the case

Aurellia only the upper floor of the ‘second pair of stomach-pouches is formed from
= ectoderm of the throat-tube, their lower (aboral) floor being of entodermal origin and
derived from the wall of the primitive stomach. Hyde’s research appears to be very care-
fully prepared, and it is probable that the second pair of stomach-pouches in Cotylorhiza is of
mixed (ectodermal and entodermal) origin as in Aurellra. The apparent analogy between the
ectodermal cesophagus of the young scyphostoma and that of the Anthozoa 1s very interesting,
for it may imply a close generic relationship between the Anthozoa and the Scyphomedusz.

In this connection we must, however, give due weight to the work of the Claus-Hadzi
school (see Genus Chrysaora) who find that the 4 primary stomach-pouches and the lining of
the throat are wholly of entodermal origin, and that therefore the scyphostoma resembles the
hydropolyps more closely than the Anthozoa.

The scyphostoma develops 16 tentacles and then gives rise to buds which grow out
from the sides of the body. The wider end of the pyritorm bud is adjacent to the parent
scyphostoma, and the mouth is at this broad end. The bud is set free and swims, rotating
through the water with its narrow posterior end directed forward. Soon, however, the bud
attaches itself to the bottom by means of its narrow aboral end and then develops into a new
scyphostoma.

This asexual development of lateral buds by the scyphostoma of Cotylorhiza seems to
be a normal process and is described by Goette, 1887, p. 24, and Claus, 1892. Claus, 1892,
reared Cotylorhiza in an aquarium and found that eggs laid at Trieste in September developed
into scyphostomz with 16 tentacles and then began to produce lateral buds in the following
July. They strobilated in August. The strobilization is monodiscus, the scyphostoma giy-
ing rise to one ephyra. The 8 marginal sense-organs are apparently developed out of the
bases of the 8 perradial and interradial tentacles, while the 8 adradial tentacles degenerate
and are absorbed. A similar process takes place in Casstopea xamachana, according to R. P.
Bigelow, 1900. The gelatinous substance is secreted by the entoderm.

Claus, 1883, has studied the young ephyrae of Cotylorhiza tuberculata. When only 1.75
mm. wide the ephyra has a simple 4-cornered mouth similar to that of the single-mouthed
Scyphomedusaw. There are 8 long, slender, cleft lobes in the radii of the 8 marginal sense-
organs. ‘The central stomach gives rise to 16 blindly ending radiating diverticula, 8 in the

radii of the marginal sense-organs and 8 adradial in position. ‘There is no ring- canal, These

canals are lined by unicellular yellow-brown alge (Zoochlorelle). There are 4 gastric cirri, one
in each interradius. The 4 lips are simple and cruciform and devoid of a marginal fringe of
tentacles. When about 2.25 mm. in diameter the oral fringe of tentacles begins to develop
around the edges of the still cruciform mouth. When 2.5 to 3 mm. wide the ring-canal develops
by fusions between the adjacent edges of the 16 radiating canals, and 8 adradial velar lappets
begin to grow out from the deep notches between the 8 primitive ephyra-lobes.

When 3 mm. wide each quadrant of the cruciform mouth is bifurcated twice, giving 16
terminal forks to the entire mouth. The central mouth, however, still remains open. At this

RHIZOSTOM

COTYLORHIZA, POLYRHIZA, 663

stage there are still only 24 lappets, 16 rhopalar and 8 velar. When 4.5 to 5 mm. in diameter
the ephyra has 8 (4 pairs) of cleft mouth-arms and the ring-canal has become unrecognizable,
owing to the fusion into a network of numerous, lateral vessels which arise from the 16 radial-
canals. It is evident that at first the ephyra is like that of the single-mouthed Scyphomedusx
and that only later it acquires the characters of the multi- eaoethed Rhizostome. This is
true of all known ephyre of the Rhizostome, and it furnishes the strongest argument for the
theory that the Rhizostome have been derived from the more simply organized, single-
mouthed Scyphomeduse.

For further details of the development of Cotylorhiza the reader should consult the papers
of Claus, Goette, Hein, and Kowaleysky.

Bouvier, 1907, finds Trachurus to be commensal with this medusa.

Haeckel’s Cotylorhiza ambulacrata described from a preserved specimen from Lessona,
Canary Islands, Atlantic Ocean, appears to me to be identical with C. tuberculata, being well
within the limits of variation of the typical medusz found at Naples.

Genus POLYRHIZA L. Agassiz, 1862.

Polyrhiza, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 156.—Haecket, 1880, Syst. der Medusen, p. 576.—VANHOFFEN,
1888, Bibliotheca Zoologica, Heft. 3, p. 40.—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, pp. 32, 81.

GENERIC CHARACTERS.

Rhizostomata dichotoma with mouth-arms bifurcated two or more times. Exumbrella
without a dome but with a central concavity and with radiating furrows. Numerous simple
filaments between the mouths. 8 rhopalia. Numerous pidialec anals and a wide marginal
network of vessels.

Homopneusts frondosa Lesson and Orythta incolor Quoy and Gaimard are believed by
Haeckel to belong to the genus Polyrhiza, but the descriptions and figures of these meduse
are so vague, fanciful, and evidently inaccurate that we can not consider them. One should
consult Lesson, R. P., 1829, Voyage de la Coquille, Mollusques, plate 12; and Quoy et
Gaimard, 1833, Voyage d |’Astrolabe, Zoophytes, tome 4, p. 297 (not plate 25, figs. 6 to 10;
these are mollusca).

Polyrhiza vesiculosa L. Agassiz.

Cephea vesiculosa, EHRENBERG, 1835, Abhandl. Berlin Acad., p. 260.
Polyrhiza vesiculosa, AGassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 156.—Hamann, O., 1881, Jena. Zeit. fiir Naturw., Bd.15,
Pp. 247, taf. 9, fig. 7 (anatomy of mouth-arms).

Bell 50 to 60 mm. wide, flat, with a pit at center of exumbrella. 32 dichotomous, radiating
furrows are separated by a deep annular furrow from the equally wide marginal zone of ie
exumbrella. 8 rhopalia. 80 lappets. In each octant are 8 rectangular aon and 2 small
thopalar lappets. The mouth-arms branch dichotomously 4 to 6 times. ‘There is a large
cluster of long, simple filaments at the center of the arm-disk. 32 radial-canals, 8 large
thopalar and 24 narrower canals, all connected one with another by a wide-meshed network
of anastomosing vessels. There is no definite ring-canal.

Bell rose-red, knobs of mouth-tentacles brownish-black. Filaments transparent. Found
in the Red Sea, at Tur and Suez.

RHIZOSTOMATA TRIPTERA Vanhéffen, sensu Maas, 1903.

Rhizostomata triptera+ R. trigona, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 41, 44.
Rhizostomata triptera, MAAS, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 46.

Rhizostome in which the lower parts of the 8 separate mouth-arms are 3-winged or
Y-shaped in cross-section, being expanded in a yentral and 2 dorsal lamella. The 3 lamellz
narrow outwardly and meet in a point at the lower end of the arm. The frilled mouths are
borne upon the 3 lamella, especially along their free edges. The mouth-arms do not bear
scapulets. The ring-muscles of the subumbrella are powerful and the radial-muscles weak.

There is no sharp line of demarkation between the Rhizostomata dichotoma with mouth-
arms V-like in cross-section and the Rhizostomata triptera wherein the arms are Y-shaped in

664 MEDUS4 OF THE WORLD.

cross-section. The two groups may, however, be maintained apart more for convenience of,
classification than because of any non-intergrading distinction between them.

Indeed the chief distinction between them is that in the Rhizostomata dichotoma the
radial muscles are powerfully and the circular muscles weakly developed whereas in the
Rhizostomata triptera the reverse is the case, the circular muscles being the more powerful.

Another distinction is that in the Rhizostomata dichotoma the axial duct of each mouth-

arm simply bifurcates sending a

branch to each arm of the V-shaped
lower part of the mouth-arm. In the
Rhizostomata triptera, however, each
s axial duct gives off 3 lateral branches
which extend downward along the
lines of the 3 rows of frilled mouths
and usually rejoin the axial duct at

the lower end of the arm. This may
be made clearer by saying that in the
Rhizostomata triptera the arm is Y-
shaped in cross-section, and the axial
duct passes down through the middle
of the Y while its 3 lateral branches
extend down near the 3 ends of the Y.
In the Rhizostomata dichotoma, however, a single duct extends down in the angle of the V and
sends off 2 branches into the arms of the V (see text-figures 404 and 411).

The genera are very closely related, being distinguished by the presence or absence of
appendages upon the mouth-arms and by the arrangement of the canal-system. A description
of the genera follows

Fic. 411.—Diagrammatic representation of the shape and position of
the mouth-arms in the Rhizostomata tri ptera.

Catostylus L. Acassiz, 1862. Neither clubs, filaments, nor other appendages upon the mouth-arms. The network of
canals on the inner side of the ring-canal ends blindly without connecting with the stomach.

Lychnorhiza Harcker, 1880. Similar to Catostylus but with filaments, and no clubs upon the mouth-arms.

Crambione Maas, 1903. Similar to Catostylus but with clubs and filaments upon the mouth-arms.

Mastigias L. Acassiz. Each mouth-arm terminates in a naked club-shaped extremity. Numerous clubs or filaments
among the mouths. The network of canals arising from the inner side of the ring-canal connects with the stomach.

Pseudorhiza yon L¥NDEN¥FELD, 1882. Similar to Mastigias, but without lateral clubs or filaments among the mouths.
A terminal club present. The canals which arise from the inner side of the ring-canal between the radial-canals
end blindly without connecting with the stomach.

Phyllorhiza L. Acassiz, 1862. Mouth-arms with lateral filaments, but without clubs, as in Lychnorhiza. Canal system
as in Mastigias.

Versura Haecket, 1880. Mouth-arms with clubs and filaments asin Crambione. 4 perradial canals arise directly from
the stomach, but the 4 interradial canals result from the fusion of a network of vessels which arise from each inter-
radial side of the stomach. No ring-canal, but a marginal network of vessels. An outer and an inner zone of ring-
muscles with an annular separation between them.

Lobonema, gen. nov. Marginal lappets of the bell elongated so as to resemble tentacles. Mouth-arm membranes per-
forated. Exumbrella covered with papilla.

Genus CATOSTYLUS L. rome 1862.

Catostylus+ Toxocl ytus+ Rhacopilus, Aassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 152, 153.

Crambessa, Haecxen, 1869, Zeit. fiir wissen. Zool., Bd. 19, Pp. 509.—VON feted ELD, 1888, Zeit. fiir wissen. Zool., Bd. 47, p
231.—Browne, 1905, Report Pearl Oyster Runes Gulf of Manaar, p. 519.

Toxocl ytus + Crambessa, Harcxe, 1880, Syst. der Medusen, pp. 585, 619.

Crambessa+ Loborhiza, VANHGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 28, 41, 44; 1902, Wissen. Ergeb. Valdivia
Exped., Bd. 3, Lfg. 1, p. 52.

Toxoclytus + Crambessa+ Loborhiza, Maas, 1903, Scyphomedusen Siboga Exped., Monog. 11, pp. 47, 61, 80, 81.

The type species is Catostylus mosaicus of Australia, first described as Cephea mosaica
by Quoy and Gaimard, 1824. Agassiz designates this as the type of the genus.

GENERIC CHARACTERS.

Rhizostomata triptera the mouth-arms of which bear neither clubs, filaments, nor other
appendages. 16 radial-canals, 8 rhopalar, and 8 adradial. The rhopalar canals extend to
the bell-margin, but the adradial canals end in the ring-canal. On both its inner and outer
sides the ring-canal gives off anastomosing vessels which may join with the radial-canals,
but which do not connect directly with the central stomach.

RHIZOSTOM Al

ATOSTYLUS 665

Among characters of minor importance, the marginal zone of circular muscles in the
subumbrella is only partially interrupted in the 8 principal radii.
pit with radiating furrows above each sense-club.

Vanhoffen, 1902, shows that Haeckel’s Crambessa and Toxoclytus are identical and
must be merged. The only possible distinction appears to be that in Toxoclytus there are 4
separate genital sacs, whereas in Crambessa there is a unitary, cruciform genital cavity.

Catostylus is very closely allied to Lychnorhiza and C ‘rambione, being distinguished solely
by having ‘neither filaments nor clubs upon the mouth-arms.

Haeckel’s Crambessa is equivalent to the genera Catostylus, Toxoclytus, and Rhacopilus
of L. Agassiz, 1862. The name Catostylus ees precedence oyer all of the others, and Haeckel
should have used this name instead of inv enting a new one.

The lower ends of the mouth-arms of C. orsini and C
mouths and the extremity is blunt, triangular, and naked.

Meduse of Catostylus are often found in brackish or muddy harbors. They are often
infested with commensal plant- -cells which may give them a peculiar opaque yellow- brown
coloration as in C, mosarcus in certain parts of Australia; for yon Lendenfeld finds that in the
harbor of Melbourne the medusa is not infested with plant-cells and is deep cobalt-blue in
color, whereas in the harbor of Sydney it is infested and is opaque, light yellow-brown. I have
also seen a swarm of these cobalt-blue meduse in Brisbane Harbor, Queensland.

It is possible, as Vanhoffen surmises, that “Cephea” dubreuillii Reynaud (1830, Lesson’s
Centurie Zoologique, p. 75, planche 23) may be a Catostylus, but the description and plate
are so unsatisfactory that I think knowledge will be advanced by dropping the species. It

There is an exumbrella

. stuhlmanni are devoid of frilled

comes from Pondichery, Indian Ocean.
Catostylus is abundant in the Indo-Pacific region, but only 2 or 3 species are known
from the Atlantic coasts of Africa and southern Europe, and not one has been found in North

Synopsis of the Forms of Catostylus.

C. cruciatus.

C. palmipes.

C. tagi.

C. pictonum

C. tagi (?).

C. mosaicus.

Diameter of bell
in mm.

Shape of bell, and
character of ex-
umbrella surface.

Number of lappets
in each octant of
bell-margin.

Length of mouth-
arms in terms of
bell-radius (r)

Length of 3-wing-
ed, pointed, lower
end of each arm
in terms of length
of simple, cylin-
drical, upper part
of arm.

Color.

Where found.

120 to 150

Hemispherical, with
deep radiating fur-
rows.

Six; 4 large triangu-
lar velar, 2 very
small ocular.

1 to 0.5

Bell yellowish-white.
Gonads and ring-
canal rose-red, or
with bluish-white
bell, deep blue lap-
pets, and red
mouth-frills.

Coast of Brazil,
harbor of Rio de
Janeiro.

64

Hemispherical, with
fine granulations.

Eight; 6 large square!

velar, and 2 oval
ocular.

ft

Northern coast of
Australia to Am-
boina.

§00

Flatter than a hemi-
sphere with dendri-
tically branching
furrows.

Ten; 4 pairs of large,
triangular velar,
and 2 small, point-
ed ocular.

3 to4

Opalescent yellow or

bluish-white. Some-
|

times brown.
Ridges of exum-
brella purple-
brown. Gonads
yellowish.

In harbors from
Portugal to Sene- |
gambia, Africa.

400

With regularly rec-
tangular elevations
bordered by fur-
rows.

Ten; 4 pairs of large
triangular velar,
and 2 small, pointed
ocular.

3 to 4?

Yellowish or opales-
cent greenish-
white, rarely
reddish-yellow.
Gonads greenish or
yellow.

| Brittany, Atlantic

coast of France,
August.

359

Nearly hemispheri-
cal, covered with
coarse granulations.

Variable, about 16
oval, long, all sim-
ilar each to each.

I to 0.5

Bell and arms yel-
lowish-white.
Opaque. Some-
times cobalt-blue.

East coast of Aus-
tralia, Brisbane to
Melbourne. In
large swarms in
harbors.

666

MEDUS OF THE WORLD.

Synopsis of the Forms of Catostylus—Continued.

Diameter of
bell in mm.

Shape of bell
and character
of surface of
exumbrella.

|

| Number of lap]
pets in each
octant of bell- |
margin.

Length of
mouth-arms
in terms of
bell-radius(r).

Length of 3-
winged,
pointed, lower
end of each
arm in terms
of length of
simple cylin-
drical upper
part of arm.

Color.

Where found.

C.stuhlmanni C. orsini. C. stiphrop- | C. viridescens.| C. ornatellus. | C. tripterus. | C. purpurus.
terus.

200 65 100 80 ? 50 115
Hemispherical, Flatly round- | Flat, with Hemispherical,) Flatly round- | Hemispherical.| Flatly round-
surface gran-| ed, smooth. smooth sur- with smooth | ed, granular ed, smooth

ular with face. surface. surface with surface.

sharp pointed
projections
on lappets.

Fourteen; 12
long, rounded|
velar, 2 short,
small ocular.

1too.g5r

Bell yellowish-
brown, milky:
yellow with
purple-brown
blotches and
streaks. Arms
colorless.
Mouths
brownish-
purple.

4 miles above
mouth of
Quilimane
River, East
Africa, Feb-
Tuary to
March.

Eighteen; 16
long, sharp-
pointed velar,
and 2 smaller
ocular.

Assab, Red
Sea.

Seven; 5 large} ? lost.
cleft velar,
and 2 slender,
sharp-point-
ed ocular.
—r r
5 5
Exumbrella | Bell sea-green,
with 4 per- | Arms color-
radial areas | less. Mouths
of brown dark-violet.
spots.
Ternate,
Malay Arch-| gami River,
ipelago. East Africa,

in November.)

lines over
lappets.

Ten; 4 pairs
of large,
bluntly point-
ed velar, and
2 small, sharp
pointed ocu-
lar.

0.66 r.

?

Mouth of Pen-| Near Puna
Island, Guay-

aquil, Equa-
dor.

Six; 4 wide
quadratic

velar, and 2
narrow, long,

projecting
ocular.

0.5

Fernando Po
Island, coast

of Guinea,

West Africa.

4 cleft, and 2
simple velar,
and 2 rho-
palar lappets
in each oc-
tant.

0.75 r.

5107

Uniform dark
brownish-
purple.

Manila Bay,
Philippine
Islands.
Common.

American waters. There are a number of local races, as is the case with Cassiopea, which
also thrives in harbors.

Catostylus mosaicus L. Agassiz.

Cephea mosaica, Quoy et Gaimarp, 1824, Voyage de I’Uranie, Zoologie, p. 569, planche 85, fig. 3.
Rhizostoma mosaica, Huxey, 1849, Philosoph. Trans. Roy. Soc. London, pp. 422, 432, plate 38, figs. 26, 27; plate 39, figs. 28-34.
Catostylus mosaicus+ C. wilkesti, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 152.
Crambessa mosaica, Harcker, 1880, Syst. der Medusen, p. 622.—von Lenpenrexp, 1883, Annals and Mag. Nat. Hist., ser. 5,
vol. 12, p. 259 (nematocysts in the gelatinous substance); 1884, Proc. Linnean Soc. New South Wales, vol. 9, pp. 428, 926;
1888, Zeit. fiir wissen. Zool., Bd. 47, p.231, taf. 19, fign. 10, 13; taf. 21, fign. 21,23; taf. 23, fign. 44-46, 51, 58, 59; taf.24,
fign. 63-65; taf. 25, fign. 66-78; taf. 26, fign. 82, 83, 93, 96, 97; taf. 27, fign. 108, 109, 111-113, 115-119 (detailed descrip-
tion).—Acassiz, A., AND Mayer, 1898, Bull. Museum Comp. Zool. at Harvard College, yol. 32, p. 16, plates 2, 3, 5 figs —
Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 47.

Bell fully 250 to 350 mm. wide, somewhat flatter than a hemisphere when expanded.
Exumbrella covered with coarse granulations. 8 rhopalia without an ocellus, but with an
exumbrella sensory pit having radiating furrows in its floor. There are about 128 long, oval,
marginal lappets, about 16 in each octant, all similar each to each and with smooth exumbrella
surfaces. Arm-disk somewhat wider than bell-radius. There is a unitary subgenital cavity.

RHIZOSTOMA—CATOSTYLUS. 667
A gelatinous papilla is found upon the subumbrella on the outer side of the opening of each
subgenital ostium. The 8 mouth-arms are about 1.5 times as long as bell-radius. The laterally
compressed, simple, upper part of each arm is only about one-sixth as long as the 3-winged,
tapering lower part. The 3 expanded membranous lamella of the lower parts of the arms are
120° apart, and their free, outer edges branch profusely and bear the frilled mouths. The
mouth-arms taper to a pointed end below. No clubs, filaments, or other appendages. The
mouths are bordered by small knobbed tentacles, which wave incessantly. 16 radial-canals
leave the cruciform central stomach and are connected by a ring-canal which gives off, both
on its outer and inner sides, an anastomosing network of vessels which fuse with the radial-
canals, but do not extend inward to the margin of the stomach.

This medusa is normally cobalt-blue, butt in Port Jackson, New South Wales, Australia,
it is infested with plant cells (Zodxanthelle), which give it a uniform creamy or brownish-
yellow color except along the upper edges of the wing-like folds of the mouth-arms, where
the deep blue color appears. In Brisbane Harbor and at Melbourne, Australia, the medusa
is not commonly infested with plant cells and is deep cobalt in color.

C. mosaicus occurs in vast swarms in the harbors and estuaries of the Australian coast
from Brisbane to Melbourne during the Australian summer and autumn.

It swims by an incessant series of pulsations of its bell-rim and tends to oppose the current.
It is the most abundant medusa along the Australian coast, and is often cast up on the beaches
in long wind-rows during storms.

A small fish, Trichinurus declivis, is often seen living commensally with the medusa.

Catostylus cruciatus.

Rhizostoma cruciata, Lesson, 1829, Voyage de la Coquille, Zooph., p. 121, planche 11, fig. 1.
Rhacopilus cruciatus, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 153.

(2) Rhacopilus cyanolobatus, AGassiz, L., Ibid., p. 152. After mss. of Couthouy.

Crambessa cruciata, HArckeEL, 1880, Syst. der Medusen, p. 620.

Bell hemispherical, 120 to 150 mm. in diameter. 8 marginal sense-organs and 48 lappets.
4 large triangular velar and 2 very small ocular lappets in each octant. There are about 32
deep, radial furrows in the exumbrella. Arm-disk as wide as bell-radius. The 8 mouth-arms
are 1.5 times as long as bell-radius. The simple, cylindrical, upper part of each arm 1s short,
while the lower part is about 4 times as long and tapers to a point. These lower parts of the
mouth-arms are 3-winged 1 In cross-section aad thickly covered with frilled mouths.

The medusa is yellowish-white with rose-red gonads and ring-canal, according to Lesson;
but according to Agassiz’s note from Couthouy’s manuscript, the bell is bluish-white with
deep-blue marginal lappets and with carmine mouths upon the arms. It should be borne in
mind that Crambessa mosaica of Australia is sometimes yellowish-white while other individuals
are deep cobalt-blue. This yellowish-white 1s caused by unicellular, commensal plant cells
which sometimes infest the medusa.

Found at Santa Catharina Island and at Rio de Janeiro, Brazil.

Catostylus palmipes.

Crambessa palmipes, HarcKer, 1880, Syst. der Medusen, p. 620.—Lunet, 1883, Annals and Mag. Nat. Hist., ser. 5, vol.
p- 268.—Scuuttz, L. S., 1898, Denkschr. Med. Nat. Gesell., Jena, Bd. 8, p. 453, taf. 33, fig. 1; taf. 34, fig. 11.

Bell 64 mm. wide, hemispherical. Exumbrella covered with fine granulations. 64
marginal lappets. In each octant 6 quadratic, truncated velar lappets, flanked by 2 oval
ocular lappets only half as large as the velar ones. Mouth-arms thick, somewhat shorter than
the bell-radius. The cylindrical, upper shaft of the arm is only about one-sixth as long as the
pointed, 3-winged, lower part. These short, upper shafts of the 8 arms are bound one to
another along their sides by 8 membranes, leaving only the pyramidal lower parts of the arms
free. Neither clubs nor filaments. Subgenital ostia 3 to 4 times as wide as the columns between
them. Color (?) Found from northern Australia to Amboina, Malay Archipelago. Most
fully described by Schultze.

Lunel, 1893, observes that this medusa is often found commensal with a fish Carnex
melampygus.

66S MEDUS2 OF THE WORLD.

\ variety of this medusa, in which the webs spanning between the basal parts of the mouth-
arms and the arm-disk are not so well developed as in the typical C. palmipes, was found by the

Fisheries Bureau steamer Albatross at a depth of 150 feet in Manila Harbor, Philippine
i I cae on January 13, 1908. Bell 56 mm. wide, flatter and more conical than a hemisphere.
Exumbrella finely granular without furrows. 64 lappets. 2 small, oval ocular and 6 indis-
tinct, rectangular, sometimes cleft, velar lappets in each octant. Arm-disk 35 mm. wide where
it arises from subumbrella, and 27 mm. wide at level of origin of 8 mouth-arms. 4 subgenital
ostia slightly wider than perradial columns. A unitary cruciform subgenital cavity. Free parts
of upper arms 5.5 mm. long, lower 3-winged parts of arms 30 mm. long, 19 mm. wide, with-
out appendages. Mouths extend to blunt tips of mouth-arms without naked areas. Canal-
system as in C. purpurus. General color in formalin dull ocher-violet, gelatinous substance
milky.

Catostylus tagi.

Crambessa tagi, Haecken, 1869, Zeitschrift fiir wissen. Zool., Bd. 19, p. 509, taf. 38, 39, 8 fign; 1880, Syst. der Medusen, p. 621.—

GrenacHER und No xt, 1876, Abhandl. Senckenberg Naturf. Gesell., Frankfurt, Bd. 10, p. 123, taf. 1-7, 17 fign—Greer,

R., 1881, Zool. Anzeiger, Bd. 4, p. 564.

(?) Crambessa pictonum, Harcket, loc. cit., p. 621.

Bell hemispherical, 500 mm. wide. Exumbrella covered with dendritically branched
ridges which extend upward from the outer, pointed ends of the lappets toward the apex of
the bell. 8 rhopalia, with an ocellus on the aboral side and an exumbrella sensory pit, the
floor of the pit covered with radiating, branching furrows. 80 marginal lappets; in each
octant 4 pairs of oval, pointed, velar between 2 small, pointed, ocular lappets, not half as wide
or as long as the velar lappets. Arm-disk somewhat wider than bell-radius. The 4 subgenital
ostia are ager than the columns between them, and there is a unitary subgenital porticus.

The 8 mouth-arms are as long as the bell-diameter. The simple, laterally compressed,
upper part of each arm is less than one-third as long as the 3-winged, lower part. These lower
parts bear 2 lateral, outwardly projecting wings which are about 60° apart, and a ventral
(inner) wing which is 150° from the lateral wings. ‘The wings taper to a point at the lower
end of the aoe -arm. The free edges of the 3 Sel ARES: leaf-like expansions of the arms
are complexly folded and bear numerous mouths which are bordered by a row of small ten-
tacles. There are neither clubs, filaments, nor other appendages upon the mouth-arms. The
circular muscles of the marginal zone of the subumbrella are interrupted in the 8 principal
radi.

The cruciform central stomach gives rise to 16 radial-canals, 8 of which extend to the
sense-organs and 8 are intermediate and adradial in position. All are connected by a ring-
canal. Centripetal to the ring-canal the 16 radial-canals give off an anastomosing network
of vessels which fuse with the ring-canal, and on its outer side the ring-canal gives off a net-
work which extends into the lappets and fuses with the outer ends of the 16 radial-canals.
Each of the 8 principal mouth-arm-canals gives off 3 side branches which extend down the
3 membranous leaves of the arm and send branches off to the mouths. These 3 branches
then fuse again with the central canal at lower end of mouth-arm.

The Poiied is opalescent yellowish or milky bluish-white, sometimes brown. The
dendritic ridges of the exumbrella are reddish or brownish- purple. Gonads yellowish.

This arena is found in brackish waters near the mouths of rivers from Senegambia,
Africa, to France. It is well described and figured by Grenacher and Noll.

“Crambessa pictonum” of Haeckel, from the mouth of the Loire and in le Croisic harbor,
France, is closely related to, if not identical with, C. tag:. It is distinguished, according to
Haeckel, by the peculiar rectangular elevations separated by furrows upon the exumbrella.
These rectangles are not quite as wide as the largest velar lappets and are all of the same size.
They are arranged in a 4-sided, cruciform system upon the exumbrella and are separated
one from another by deep, parallel furrows. The velar lappets are said to be somewhat
wider than they are long, and the mouth-arms are shorter than in C. tagr. Haeckel found this
medusa in August on the southern coast of Brittany. He states that when the medusa’s bell
is 30 mm. wide, the mouth-arms have only a single row of mouth-frills on their ventral sides,
as in his genus Haplorhiza, and the lateral, leaf-like wings develop later.

RHIZOSTOMA#®—CATOSTYLUS. 669

Catostylus stuhlmanni.

Crambessa stuhlmanni, Cuun, 1896, Mittheil. Naturhistor. Museum, Hamburg, Bd. 13, Pp. 10, taf. 1, 2 fign.

Bell hemispherical, 80 to 200 mm. wide. 8 marginal sense-organs, set within deep clefts
in the bell-margin. 112 marginal lappets. The ocular lappets are short, pointed and small,
but the 12 velar lappets in each octant project farther outward and have rounded margins.
They are separated one from another by long, deep grooves extending up the sides of the ex-
umbrella. Each of these lappets is provided with a median longitudinal row of sharp-pointed
projections on the exumbrella side. The 8 mouth-arms are bluntly pointed and are shorter than
the bell-diameter. The subgenital ostia are one-third to one-fourth as wide as the columns
between them. A unitary, subgenital porticus.

The bell is yellowish-brown or milky-yellow, besprinkled with purple-brown blotches,
which are most numerous near the margin. The marginal lappets have rusty-brown, longi-
tudinal median streaks. The arms are colorless and the mouths are usually spotted with
brownish-purple.

8 specimens found 4 miles above the mouth of Quilimane River, East Africa, in February
and March. Described in detail by Chun, 1896.

This species resembles C. orsini in that the mouths are not developed upon the lower,
pointed, knob-shaped ends of the mouth-arms. The outer zone of circular muscles of the
subumbrella is interrupted near the margin in the 8 principal radu, but centripetal to this
they are unbroken and form a complete annulus.

Catostylus orsini.

Mastigtas orsint, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 34, 44, taf. 4, fign. 2-4; 1902, Wissen. Ergeb.
deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, pp. 48, 49-—Maas, 1903, Scyphomedusen der Siboga
Expedition, Monog. 11, pp. 63-66.

Umbrella 65 mm. wide, with smooth, exumbrella surface flatly rounded with incurved
margin. 8 marginal sense-organs. 144 small, sharp-pointed, marginal lappets. 16 velar
lappets between 2 somewhat smaller, ocular lappets in each octant. A radial furrow extends
up the side of the exumbrella in the line of the cleft between each adjacent pair of lappets.
The 8 sense-organs are set within deep niches and there is an exumbrella sensory pit with
radiating furrows above each sense-club. The subumbrella displays a deep annular furrow
on the inner side of which lies the ring-canal. Centrifugal to this furrow is a zone of power-
ful circular muscles, which are not interrupted in the 8 principal radii.

The arm-disk is nearly as wide as the radius of the bell and the 4 arm-disk pillars are
wider than the ostia of the subgenital porticus. The simple upper part of each of the 8 mouth-
arms is very short and only one-third as long as the 3-winged lower part of the arm. It is
also thin and ribbon-like, and in this respect is in marked contrast to the large, 3-cornered
lower part of the arm. The upper part of the arm bears no dorsal mouths, but only a single
row of frilled mouths along its ventral side. The large, 3-sided, lower part of the arm is elongate,
prismatic, with a short gelatinous, pyramidal, bluntly pointed, 3-cornered knob at its end.
Altogether the entire mouth-arm is about as long as the bell-radius. The lower part of the arm
bears frilled mouths, but neither filaments nor other appendages. The lower end of the arm
isnaked and devoid of mouths, as in Catostylus stuhlmannt, and forms a blunt, triangular
knob which on the outer side is nearly half as long as the upper part of the arm itself, but
only one-third of this length on the two radial sides.

8 canals extend down the middle of the 8 mouth-arms and send ramifying branches to
the frilled mouths. These 8 arm-canals enter the small, central stomach, from which arise
16 straight radial-canals, 8 ocular and 8 adradial, connected one with another by a wide,
circular vessel. An anastomosing network of vessels arises on the inner side of the ring-canal
between the radial-canals, although this network does not fuse with the radial-canals them-
selves, but arises solely from the ring-canal. The radial-canals are about twice as wide as
the ring-canal. The unitary, subgenital porticus is very small. Color (?) Found at Assab,

Red Sea, in June.

670 MEDUS OF THE WORLD.

Catostylus stiphropterus.
Crambessa stiphroptera, Scuurtze, L. S., 1897, Abhandl. Senckenberg, Naturforsch. Gesell. Frankfurt a. M., Bd. 24, Heft 2,

p- 159, taf. 15, fign. 4, 5, 5a-

Bell flatly rounded, 100 mm. wide, with a smooth, exumbrella surface which lacks the
protuberances found in C. mosaicus. There are 8 marginal sense-organs. The rhopalar,
marginal lappets are slender and sharp-pointed, and in each octant there are at least 5 larger,
cleft, velar lappets about 10 mm. long and 6 mm. wide at their bases. The arm-disk is about
38 mm. in diameter, somewhat less in width than the bell-radius. The 4 arm-disk-columns
are nearly as wide as the 4 narrow, subgenital ostia. Lower arm 5 times as long as upper, the
total length of both not quite equal to that of the bell-radius. Upper arm wholly free, differing
in this respect in the number of its marginal lappets and in its narrow subgenital ostia from
C. palmipes

The exumbrella displays 4 perradial areas of indistinct, round, brown spots which do
not extend to the bell-margin.

Found at Ternate, Malay Archipelago.

Catostylus viridescens.

Crambessa viridescens, Cuun, 1896, Mittheil. Naturhist. Museum, Hamburg, Jahrg. 13, p. 12, taf. 1, fig. 2.

Bell 80 mm. wide, hemispherical. Marginal lappets (?) Arm-disk wider than the bell-
radius. Subgenital ostia wider than the spaces between them. 8 short mouth-arms, not
longer than bell-radius. Upper arm about one-fifth as long as the lower part. Bell sea-green,
mouth-arms colorless. Frilled mouths dark-violet. Two specimens found at the mouth of
the Pangani River, East Africa, late in November. The marginal lappets were lost in both
specimens. Characterized chiefly by its sea-green color.

Catostylus ornatellus.

Loborhiza ornatella, VaNuOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 28, 41, taf. 2, fign. 3-6

Disk flatly rounded, the gelatinous substance thick. Size (?) The exumbrella is finely
and evenly Soran but these granules fuse into rows upon the marginal lappets. 8 mar-
ginal sense-organs and 80 marginal lappets. In each octant there are 4 pairs of bluntly pointed,
nearly rounded, velar lappets, Read 2 very small, sharp-pointed, lancet-shaped ocular lappets.
The velar lappets adjacent to the ocular lappets project farther outward and are sharper-
pointed than the remaining velar lappets. There is a powerfully developed zone of ring-
muscles in the subumbrella, but these are relatively indistinct and somewhat interrupted
in the radii of the 8 ocular radial-canals.

The arm-disk is supported by 4 thick arm-pillars, which flare outward at their sub-
umbrella bases so as to recall a Maltese cross when viewed looking toward the subumbrella
surface. The 4 perradial columns of the arm-disk are about as wide as the 4 genital ostia,
but they appear wider thaa the genital ostia, for their flaring bases curve around in 8 hook-
like lateral projections so as to partially close the openings oF the genital ostia. The opening
of each genital ostium is still further blocked by a triangular pointed flap of the arm-disk
which projects over the middle of the ostium, so that each ostium appears as if constricted
into 2 side-openings (see figures by Vanhoffen, 1888, taf. 2). The arm-disk is octagonal,
and there is a unitary subgenital porticus with 4 folded, U-shaped gonads.

The 4 pairs of eaonthe -arms are very thick, but only about two-thirds as long as bell-
radius. The simple upper half of each arm is short, but the lower half gives rise to 2 dorsal,
wall-like lamella which bear the mouths on their free outer edges. The ventral side of Gath
mouth-arm also gives rise to a similar lamella; and thus the lower parts of the mouth-arms
are 3-rayed in cross-section. The 2 dorsal lamella are set off one from another at an angle
of about 60°, while the ventral lamella is at an angle of 150° from the 2 dorsal lamella. The
outer edges of these 3 wing-like lamella fold in and out and give rise to short, lateral branches,
along the edges of which the numerous mouths are placed. There are neither filaments nor
other appendages among the mouths. The 3 wings of the mouth-arms end in a blunt point at
the lower extremity of the mouth-arms:

RHIZOSTOM2®—CATOSTYLUS. 671

The 8 ocular radial-canals are joined one to another by a thick, irregularly anastomosing,
network of canals. The ring-canal is not clearly defined. Other radial-canals (?) Color (?)
Found at Puna Island, near Guayaquil, coast of Equador, South America.

Catostylus tripterus.

Toxoclytus tripterus, Hance, 1880, Syst. der Medusen, p. 586.
Crambessa triptera, VANHOFFEN, 1902, Wissen. Ergeb. deutschen Tiefsee Exped. Valdivia., Bd. 3, Lfg. 1, Pp. 52.

Bell 50 mm. wide, hemispherical. 8 rhopalia, 48 lappets. In each octant 4 wide, nearly
quadratic, velar lappets between 2 smaller, but longer, conspicuously projecting, ocular
lappets. 8 arms, somewhat longer than bell-radius, are grouped 1 in 4 pairs, and each consists
of a stout, long, nearly cy hadtical upper part of the arm which ts twice as long as the 3-cornered
pyramidal, lower part of the arm. There are 3 wide, leaf-like projections on this lower part
of the arm, and the mouths on their edges are only slightly folded. There are no appendages
between the mouths. 4 horseshoe-shaped gonads.

Found on the west coast of tropical Africa at Fernando Po Island, on the coast of Guinea.
Color (?)

Catostylus turgescens.

Toxoclytus turgescens, ScHuttzeE, L. S., 1898, Denkschr. Med. Nat. Ges. Jena., Bd. 8, p. 455, taf. 34, figs. 13, 14.

This is described by Schultze from a single specimen which appears to be quite abnormal—
so much so that I have but little faith in its value.

Bell flatly rounded, g0 mm. wide. Exumbrella smooth. g marginal sense-organs.
Marginal lappets narrow and sharp-pointed. Number (?) Arrangement (?) 6 subgenital
ostia aides than the arm-shafts which separate them. Arm-disk flat. The 6 upper arms are
thick, arrowhead-shaped and curve outward; they are about 19 mm. long and 17 mm. wide.
Lower arm sharply pointed, only about 12 mm. long. ‘There are thin. Salaments upon the
arm-disk between the mouths, but no other appendages. The canal-system consists of a fine
anastomosing network on the inner and outer sides of the ring-canal. This network does
not reach ‘he margin. ‘There are also unbranched radial- canals and blindly-ending centrip-
etal canals. Amboina, Moluccas. Color (? )

Catostylus purpurus, sp. nov.

This form is closely related to Catostylus stiphropterus, from Ternate, but differs in the
number and arrangement of its marginal lappets and in its deep, uniform dark brownish-
purple coloration.

Disk flatter than a hemisphere, 88 to 115 mm. wide, 26 to 35 mm. high. Exumbrella
smooth. 8 rhopalia flanked by short, narrow, bluntly rounded lappets. A furrowed exumbrella
sensory pit above each rhopalium Rhopalar lappets somewhat narrower than the velar.
In each octant there are 4 cleft and 2 simple velar lappets arranged in a definite manner, see
a and p, fig. 412. In the middle of each octant there are a pair of cleft velar lappets, and
these are femeal on their outer sides by 2 simple velar lappets, which are in turn bordered
by 2 cleft velar lappets. Thus the lappets of each octant are arranged in sequence as fol-
lows: (1) a small, simple, rhopalar lappet adjacent to the sensory- Be iise (2) a cleft velar
lappet; (3) a simple velar lappet; (4 and 5) 2 cleft velar lappets; (6) a simple velar lappet;
(7) a cleft velar lappet; (8) a small rhopalar lappet. Thus the bell-margin displays 96 nearly
equally spaced notches, there being 16 rhopalar and 80 velar terminal lappets.

The arm-disk is about as mide as the bell-radius at its origin from the subumbrella, but
at the level of the origins of the 8 mouth-arms it 1s somewhat less than three-eighths as wide
as the bell-radius. There are 4 long, narrow, genital ostia nearly as wide as the 4 perradial
columns of the arm-disk. Each ostium is constricted by a thick, wide, median gelatinous
projection from the arm-disk. A long, finger-shaped papilla arises from the subumbrella
surface in the median line on the outer side of and close to the opening of each genital
ostium, and this is in some specimens flanked by a pair of cocks-comb shaped, gelatinous
projections from the floor of the subumbrella as is shown in c in text-figure 412. The arm-
disk is notched in each perradius. The unitary subgenital cavity is wide and cruciform.

672 MEDUS OF THE WORLD.

There are 8 separate mouth-arms, each three-eighths as long as the bell-diameter. The
lower, 3-winged, expanded part of each arm is about 5 times as long as the simple, flattened,
upper part of the arm. Each mouth-arm is bluntly pointed and its frilled mouths lack fila-
ments or other appendages. The mouth-frills extend to the extreme tip of the arm and there
is no naked, terminal portion.

A zone of powerfully developed, unbroken, circular, subumbrella muscles extends from
the outer edge of the arm-disk to the bell-margin. The gelatinous substance of the bell is
very tough and of a leathery consistency.

"16 radial-canals leave the central stomach: 8 rhopalar and 8 adradial. These are con-
nected by a ring-canal on the outer side of which there is a fine-meshed and on the inner side
a coarse-meshed network of anastomosing vessels.

The medusa is dull, uniform dark brownish-purple, resembling old leather soaked in
water. It is abundant in Manila Bay, Philippine Islands, where it occurs over the bottom
in shallow water.

Seven specimens found in Manila Bay on December g, 1997, are in the collection made
by the U. S. Fisheries Bureau steamer Albatross, and a larger one on March 11, 1908. This
largest specimen serves as the type of the species in the National Museum at Washington. Its
dimensions in mm. are as follows: Bell 115 wide, evenly rounded, 35 high; arm-disk 75 wide
where it arises from the subumbrella, 52 wide at level of origin of mouth-arms; mouth-arms
58_long, upper arm 7 long, lower arm 51 long and 30 wide.

Fic. 412.—Catostylus purpurus, sp.nov. Drawn by the author, from specimens obtained by the U.S. Bureau of
Fisheries steamer Albatross in Manila Harbor.

A, oral view, half natural size. Only two of the mouth-arms are shown; § others are cut off close to their points
of origin, and one is shown cut across in its expanded 3-winged part. B, side view. C, genital ostium
showing subumbrella papilla flanked by a pair of cocks-comb-shaped subumbrella projections. D,
exumbrella view of rhopalium showing furrowed sensory pit.

Genus LYCHNORHIZA Haeckel, 1880.

Lychnorhiza+ Cramborhiza, HArcKe1, 1880, Syst. der Medusen, pp. 587, 633.—VANHGFFEN, 1888, Bibliotheca Zoologica, Bd. 1,
Heft. 3, pp. 28, 41.
Lychnorhiza, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, pp. 48, 80; 1906, Revue Suisse de Zool., tome 14, p. 102.

The type species is Lychnorhiza lucerna Haeckel, from the coast of Brazil, Rio de Janeiro
to Pernambuco.

GENERIC CHARACTERS.

Rhizostomata triptera with filaments, but without clubs, upon the 3-winged mouth-arms.
No axial terminal club at the end of each arm, and no club-shaped appendages between the
mouths. The stomach gives rise to 16 radial-canals: 8 rhopalar and 8 adradial. The rhopalar
canals extend to the bell-margin, but the adradial ones end in the ring-canal. Blindly ending
centripetal vessels arise from the inner side of the ring-canal and may anastomose to some
extent. On its outer side the ring-canal gives off a network of anastomosing vessels which
extend into the lappets.

RHIZOSTOMA8—LYCHNORHIZA. 673

Among characters of minor importance the circular muscles of the subumbrella are
entire and not broken in the 8 principal radii. The sense-clubs have each a sensory pit with
radiating furrows over its floor. The subgenital ostia are wider than the columns between
them.

This genus is so closely allied to Crambrone that the two might readily be merged. It
may, Hower er, be distinguished by having no club-like appendages between the fouthe frills.

Lychnorhiza lucerna Haeckel.

Lychnorhiza lucerna, HAecKEL, i880, Syst. der Medusen, p. 587, taf. 34, 8 fign.—Hamann, 1881, Jena. Zeit. fiir Naturw., Bd. 15,

p- 249 (anatomy of mouth-arms). i
Cramborhiza flagellata (young medusa), Harcket, Ibid., p. 646.

Lychnorhiza flagellata, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft 3, Pp: 29, 42, taf. 2, fig. 7; taf. 3, fign. 1-3.

The bell is flatter than a hemisphere, 120 to 150 mm. in diameter, and its exumbrella
surface is besprinkled with fine granules and minute, sharp-pointed projections. There are
8 marginal sense-organs and 48 marginal lappets. The 16 lappets flanking the 8 marginal
sense-organs are small, sharply pointed, and triangular, while the 4 velar lappets of each
octant are 3 times as broad and 3 to 4 times as long as those flanking the sense-organs. These
velar lappets are more nearly oval in outline and not quite so sharply pointed as are the ocular.
The 8 stout, adradial mouth-arms arise from a large, gelatinous base which projects from
the center of subumbrella. These 8 mouth-arms are laterally compressed, separate one from
another, and about as long as the bell-diameter. The outer sides of the upper halves of the
mouth-arms are smooth and bear no mouths, but below this the arm is developed into a ventral
median and 2 large lateral wing-like membranes, all 3 of which meet at a point below. The
edges of these 3 membranes are much folded and are lined by numerous mouths which are
surrounded by minute, clubbed tentacles. In addition, the edges adjacent to the mouths
bear numerous long filaments, 120 to 160 upon each mioutbe -arm. Near their bases these
filaments are wine. but they expand beyond into a ribbon-like shape, each of the narrow
edges of the ribbon being lined by a row of small, club-like nematocyst-organs. Each mouth-
arm bears a pair of very long filaments and 15 to 20 somewhat shorter Berens: the remainder
being still shorter. The longest filaments exceed the length of the mouth-arms themselves.

There is a well-developed, unbroken zone of circular muscles in the outer part of the
subumbrella. The central stomach is cross-shaped, the arms of the cross being in the diameters
of the principal radii, while the 4 subgenital pits lie in intermediate positions. 16 radial-
canals extend out from the central stomach: 4 in the principal radii, 4 in the secondary, and
8 in the tertiary (adradial) radii. The 8 principal radial-canals extend to the sense-organs,
but the 8 adradial ones end in the ring-canal, which is at some distance inward from the
margin. The ring-canal gives off 32 blindly- -ending, centripetal vessels, 2 between each pair
of radial-canals. On its outer side the circular vessel gives rise to about 160 radiating vessels
(20 in each octant) which extend into the lappets and are connected by numerous, anas‘omosing
vessels forming a marginal network. Each of the 8 mouth-arms sends a canal into the central
stomach. This main canal of each mouth-arm gives rise to 2 side branches, each of which
extends down a lateral wing of the mouth-arm under the mouths, while the central canal
extends down the center of the lower side of the mouth-arm. The gonads are much-folded
membranes lining the inner walls of the 4 subgenital pits. They fill the greater part of the
stomach cavity and according to Haeckel, they project outward through the subgenital pits.
This is, however, unknown in any other rhizostomous medusa and is, I believe, merely a result
of shrinkage, etc., in the preservative fluid.

Haeckel studied a single specimen of this medusa from Rio de Janeiro, Brazil. He
gives a detailed description ae aes by figures.

I am inclined to believe that “LZ. flagellata” is only the young of L. lucerna. In order
the more readily to aid future students in settling this question, we present a detailed description
of “L. flagellata:?’ The disk is 80 mm. wide, about 30 mm. high, and evenly rounded. It is
very tough with thick gelatinous walls. The exumbrella is covered with fine granules, which
become larger near the marginal lappets and set themselves in elongate lines over the lappets.
There are 8 marginal sense-organs and 48 marginal lappets. In each octant 4 large, bluntly
triangular, velar lappets between 2 very small, sharp-pointed, lancet-shaped, ocular lappets

674 MEDUSA OF THE WORLD.

which are hardly half as long and one-fifth as wide as the velar lappets. The marginal sense-
organs are similar to those of Catostylus. The ring-muscles of the subumbrella are very
powerfully developed, but are partially interrupted over the 8 ocular radial-canals. The arm-
disk is about as wide as the radius of the umbrella. It is 8-sided, the narrow sides being in
the radii of the arm-pillars; and the wide sides, which are 3 times as wide as the others, are
in the radii of the 4 subgenital ostia. The subgenital ostia are thus 3 times as wide as the arm-
pillars between them.

8 thick mouth-arms arise from the arm-disk, and these are about as long as the bell-
radius. The lower part of the arm is somewhat longer than the upper. There are 2 well-
developed, thick, dorsal mouth-lamella or “wings” which project from the lower arm and
these fuse with the ventral mouth-lamella at the pointed end of the arm. The ventral side of
the mouth-arm is complexly folded and gives rise to lateral lappets. The filaments, which
arise at the ends of these lappets between the mouths, are shorter than in the mature L. /ucerna.
The 4 sides of the genital organs are bent at right angles, thus forming a cross of 4 right-
angled membranes mehich are very much folded. The gastrogenital cavity and subgenital
porticus are small and much reduced.

16 radial-canals (8 ocular and 8 interocular) emerge from the cruciform, central stomach.
These 16 radial-canals are put into connection one with another by a wide ring-canal which
is about half-way between the center and the margin. Peripheral to this ring-canal there is
a network of vessels, although the 8 ocular canals run through and fuse with this network.
32 blindly-ending, centripetal vessels extend inward from the ring-canal toward the center of
the disk, but they end blindly before reaching the edges of the stomach. There are 2 of these
blindly-ending diverticula between each successive pair of radial-canals, and in some cases
they fuse one Sith another. Color (?)

Found at Pernambuco and at Contigeriba on the coast of Brazil. The most complete
description is that of Vanhéffen, 1888, from which the above has been mainly derived.

I am inclined to believe that this medusa will prove to be only a young stage of Lych-
norhiza lucerna Haeckel. The marginal lappets, mouth-arms, and canal-system are similar
in both. According to Haeckel the bell is flatter and thinner in L. lucerna than in L. flagellata;
also in the mature L. lucerna the gonads protrude through the subgenital ostia, but this may
be due to defects in preservation or to the general breaking up of these organs which commonly
occurs in medusz when the genital products are set free. Vanhoffen did not compare his
specimen of L. flagellata with Haeckel’s type in the Berlin Museum. The only distinguishing
features according to the accounts of Haeckel and Vanhoffen are as follows:

Lychnorhiza lucerna: Lychnorhiza flagellata:
Disk flat, gelatinous substance thin. Disk nearly hemispherical, gelatinous substance
Umbrella 120 to 150 mm. wide. thick.

Umbrella 80 mm. wide ?*

Mouth-arms are twice as long as the radius of the Mouth-arms are only a little longer than the ra-

umbrella. dius of the umbrella.

Mouth-arm filaments longer than the mouth-arms. Mouth-arm filaments very short (broken off ?).
| 4 separate, subgenital cavities. Gonads protrusive. Subgenital porticus present. Gonads do not pro-

| trude through the subgenital ostia.

*Vanhdffen does not state the size of his specimen.

The presence of a unitary subgenital porticus hele be flagellata” is its only really distinctive
character, but this is often highly variable in development in different specimens ‘of the same
medusa. See Maas, 1903, Scyphomedusen Siboga Exped., p. 36.

Lychnorhiza bartschi, sp. nov.

Named in honor of Dr. Paul Bartsch to whose care and skill the excellent preservation of
the meduse upon the Philippine expedition of the Albatross is due.

Bell 74 mm. wide, flatter than a hemisphere and with smooth exumbrella surface. Gelat-
inous substance thick but not very rigid, 8 rhopalia, each with an ocellus and an exumbrella

RHIZOSTOMA8—LYCHNORHIZA. 675

sensory pit with dendritic furrows over its floor. 96 (812) lappets, 10 bluntly pointed velar
lappets between 2 somewhat smaller ocular lappets in each octant. Arm-disk 48 mm. wide
where it arises from the subumbrella, but only 40 mm. wide at the level of origin of the 8 mouth-
arms. The 4 subgenital ostia (s go) are crescent-shaped and each is covered above by a
gelatinous fap. They are only half as wide as the perradial columns between them. Each
perradial column exhibits a niche, 1, figs. 413 and 414, on its outer side which bears a super-
ficial resemblance to the subgenital ostia. The subgenital cavity is unitary. » Bly

The 8 mouth-arms are laterally compressed and 36 mm. long, the lower 3-winged parts of
the arms being 24 long and 23 wide. Numerous simple, laterally flattened, tapering filaments
arise from between the frilled mouths on all sides of the mouth-arms and from the arm-disk.
The filaments upon the arm-disk are about 30 mm. long, but those from the outer parts of the
mouth-arms are shorter.

The central stomach is cruciform and about 40 mm. wide. 16 simple ‘radial-canals, 8
thopalar and 8 adradial. These are all put into intercommunication with a wide ring-canal
which is at some distance inward from the margin. The adradial canals terminate in this
ring-canal, but the rhopalar canals extend onward to the sense-organs. On its inner side the

413. 413. 414.

Fic. 413.—Lychnorhiza bartschi. Drawn by the author. a, Oral view. 8, sense-organ seen from exumbrella side.
Fic. 414.—Lychnorhiza bartschi, sp. nov. Natural size, drawn by the author. sgo, subgenital ostium. 1, niche in perradial
column of arm-disk.

ring-canal gives rise to 16 blindly ending networks of vessels which do not connect either with
the stomach or with the radial-canals. On its outer side a fine-meshed network of vessels arises
from the ring-canal and fuses with the rhopalar vessels. Around the margin at the bases of
the lappets there is a marginal ring-canal of fine caliber. There is a unitary uninterrupted
system of ring-muscles in the marginal zone of the subumbrella, but there are no radial
muscles.

The gelatinous substance is translucent and milky in formalin, and the gonads, mouth-
frills, and canal-system are milky-yellow. I am told by Dr. Bartsch that these colors in the
living animal were nearly as they appear in the formalin specimen.

A single specimen was found by the U. S. Fisheries Bureau steamer, Albatross, at Jolo
Anchorage, Philippine Islands, on February 13, 1908 (text-figs. 413 and 414).

676 MEDUS OF THE WORLD.

Genus CRAMBIONE Maas, 1903.

Crambione, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, pp. 48,81; 1906, Révue Suisse de Zool., tome 14, p. 103.
The type species is Crambione mastigophora, Maas, from the Malay Archipelago.
GENERIC CHARACTERS.

Rhizostomata triptera in which each mouth-arm is 3-winged and the wings bear secondary
branches. All 3 of these wing-like expansions and their branches bear mouths, among which
there are clubs and filaments.” No terminal club at the end of each arm. With a unitary, sub-
genital porticus and with 4 slit-like, subgenital ostia.

The canal-system consists of 8 vessels which extend outward to the bell-margin in the
perradii and interradii, and 8 adradial canals which end in the ring-canal at some distance
inward from the bell-margin. On the outer side of the ring-canal is a network of vessels, and
on the inner side the ring-canal gives off a network between each 2 radial-canals, which does
not connect with the radial-canals themselves. The circular muscles are unitary, being
unbroken by radial strands. The marginal sense-organs have a pair of eye-spots and a
sensory pit with large radial furrows.

This genus is closely related to Catostylus, but is distinguished by having clubs and fla-
ments upon its mouth-arms, these being absent in Catostylus. It is also very closely allied to
Lychnorhiza, but has both cae and filaments upon its mouth-arms, whereas Lychnorhiza

has filaments only.
Crambione mastigophora Maas.

Crambione mastigophora, Maas, O., 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 49, taf. 6, fign. 47-53; taf. 8,

figs. 71-74; taf. 11, fign. 100, 104; taf. 12, fig. 113; 1906, Revue Suisse de Zool., tome 14, p. 103.

The bell may become 400 mm. wide and is highly arched and rounded. The gelatinous
substance of the center is thick, while the margin is sharply set off from the center and is
thin-edged. The exumbrella is smooth. There are 8 marginal sense-organs. These sense-
clubs have each a bulbular swelling on the subumbrella side and 2 lateral ocelli near the outer
end of the club. There is also an entodermal lithocyst mass. There is a large, heart-shaped
sensory pit on the exumbrella side above each sense-organ, and prominent radiating furrows
spread out from the center of the pit-cavity just aboye the base of the sense-club. 2 small,
pointed, lanceolate, ocular lappets flank each of the 8 sense-organs, and in each octant are
also 8 to 10 velar lappets which are elongate, with rounded outer edges and deep clefts between
them, and which increase in number with age.

The arm-disk is very wide and 8-sided. The 4 interradial, subgenital ostia are narrow,
elongate, and slit-like, but not as long as the arm pillars between them. 4 perradial, slit-like
depressions or fosse in the arm-disk are somewhat higher than the subgenital ostia to which
they bear a close superficial resemblance; they are not to be confused with subgenital ostia,
however, for they are mere depressions in the surface of the arm-disk. The 8 adradial mouth-
arms alternate in position with the subgenital ostia and the perradial fossz. In the young
medusa they are grouped in 4 pairs, but in the adult they arise at equal intervals from the sides
of the arm-disk. Basal parts of mouth-arms massive, nearly circular in cross- section; in
their lower halves each gives rise to 3 projecting, lateral expansions or “‘wings” which meet at
the lower end of each arm, giving a pyramidal general outline to the outer half of each mouth-
arm. ‘There are numerous mouths along the lower inner lamella of each mouth-arm and
along the edges of the 2 lateral wings, as in Catostylus. But unlike Catostylus many small,
club-shaped and some long, tapering, filamentous appendages arise from both the lower and
upper sides of the mouth-arms between the mouths.

The central stomach is cruciform, the axes of the cross being in the perradii. 4 perradial,
4 interradial, and 8 adradial canals arise from the stomach. The perradial and interradial
canals extend to the bell-margin, but the 8 adradial vessels end in the ring-canal, which lies
some distance inward from the margin of the bell. On its outer side the ring-canal gives
off a network of vessels which anastomose with the perradial and interradial canals. Centrip-
etal to the ring-canal and arising from it, between the 16 radial-canals, are 16 open networks
of vessels. The ring-canal and the 16 radial-canals are of uniform and moderate width.
The peripheral network of vessels is of finer caliber and the 16 networks on the inner side
of the ring-canal are of wider caliber than the outer network, but not as wide as the radiating
vessels. Thea inner networks do not fuse with the radial- eral,

RHIZOSTOM®—CRAMBIONE, MASTIGIAS, 677

The 4 interradial gonads form a cross following the lines of the cruciform stomach of the
medusa but interior to the border. The adjacent gonads lie so close one to another that the
genital cross is extremely narrow and elongate. ier is a unitary, subgenital porticus or
chamber which serves as a brood-sac for "he planula larve. There is a well- -developed,
peripheral ring-muscle in the subumbrella, and this is not Leglees by radial muscle-strands,
such as are found in Mastigzas.

The gelatinous solbsianree | is translucent and milky. The frilled mouths are whitish and
the clubs reddish. The gonads are flesh-colored, pinkish, or reddish.

This medusa is fond at Amboina and at other places among the islands of the Malay

Archipelago. It is described and figured in detail by Maas, 1903.

Crambione cookii, sp. nov.

Plate 74, fig. 2

Bell hemispherical, 110 mm. wide, gelatinous substance tough. Exumbrella smooth in
the flexible zone above the margin, but the inflexible central part of the dome is reticulated
by a network of deep furrows trending more or less radially outward from the apex. 8 rho-
palia. 88 large, pointed, marginal lappets, equal in size each to each. Mouth-arms 1.5 times
as long as bell-radius. The lower two-thirds of each arm is 3-winged and the outer edges of
these lamella are complexly folded and bear the mouths. 4 slender, uniform filaments, as
long as the bell-radius, arise from the arm-disk. There are about 2 to 6 globular, gelatinous
appendages on the outer sides of each mouth-arm. These are about 6 mm. long.

The gelatinous substance of the bell is opaque, horny, milky-yellow. The marginal
ring-muscles of the subumbrella are brown and the valleys of the exumbrella furrows are of
a lighter shade of the same color. Mouth-arms and vesicles translucent milky-blue. Mouth-
frills brown.

Found by me on the surface along Great Barrier Reef, off Cooktown, Queensland, Australia,
May 4, 1896, during Dr. Alexander Agassiz’s exploration of the reefs. Named in honor of
the distinguished navigator, Captain James Cook, whose yoyage first made the Queensland
coast known to the world, and whose ship, the Endeavour, met with misfortune in June, 1770,
near the place wherein this medusa was found.

Genus MASTIGIAS L. Agassiz, 1862.

Mastigias, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 152.—Cxaus, 1883, Organisation und Entwick. der Medusen,
p- 61.—von LenpENFELD, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 300.—Kisuinovure, 1895, Zoological
Mag. Tokyo, vol. 7, No. 78.—Cuun, 1896, Mittheil, Nat. Mus. in Hamburg, Jahrg. 13, p. 13—Maas, 1909, Abhandl.
Akad. Wissen., Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 46.

Mastigias + Eucrambessa, HarcKer, 1880, Syst. der Medusen, pp. 622, 624.

Mastigias+ Desmostoma, VANHGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp- 33, 35, 44, 45-—Maas, 1903, Scyphome-
dusen der Siboga Exped., Monog. 11, pp. 62, 66, 81.

Mastygias, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Valdivia, Bd. 3, Lfg. 1, p. 46.

The type species is the widely distributed MM. papua of the Indo-Pacific region. It was
first described as Cephea papua by Lesson, 18209.

GENERIC CHARACTERS.

Rhizostomata triptera with 3-winged mouth-arms which terminate in a naked, club-
shaped extremity. There are also smaller clubs and filaments between the frilled mouths.
The mouths are developed not only along the edges of the 3 leaf-like wings of the lower parts
of the mouth-arms, but also over parts of their flat, expanded sides. The central stomach
gives rise to 8 rhopalar canals and numerous, interocular radial-canals all of which anastomose
and finally connect with the ring-canal. The rhopalar canals extend straight to the sense-
clubs, but jthe inter-rhopalar canals end in the ring-canal. On its outer side the ring-canal
gives off a network of vessels which extend into the lappet-zone and fuse with the outer ends
of the rhopalar canals. The ring-muscles of the subumbrella are interrupted in the 8 rhopalar
radii. A unitary subgenital porticus. No furrows in the exumbrella sensory pits.

The genus Desmostoma of Vanhéffen, 1888, conforms in all respects to Agassiz’s Mastigias
except that there are clusters of filaments upon its arm-disk at the bases of the mouth-arms,
whereas the other species of Mastigias lack filaments and have only small clubs on the sides

678 MEDUS® OF THE WORLD
of the mouth-arms in addition to the terminal club, I have merged it with Mastigias for the
genera among Rhizostomata triptera are already too numerous, and are distinguished upon
differences of “such slight importance that the distinctions threaten to confuse rather than to
clarify the system of aesecaGem

Mastigias is closely allied to Pseudorhiza, but may be distinguished by 1 its numerous,
complete, interocular radial-canals; whereas the converging vessels on the inner side of the
ring-canal between the 16 radial-canals in Pseudorhiza end blindly without reaching the
stomach.

The canal-system of von Lendenfeld’s Phyllorhiza punctata is similar to that of Mastigras,
but the mouth-arms bear numerous very long filaments without any definite terminal club.

The following synopsis of the forms of Mastigias may be of service. r is the length of
the radius of the exumbrella.

Tabular Synopsis of the Forms of Mastigtas.

M. papua. | M. papua var. | M. papua var. | M. ocellata. M. pantherina.| M. gracile.

| siderea. siboge.
| Number of 8X8. Rounded. | 88. Rounded. | 9X8. Rectangu- | 6X8 or 12X8 16 X8. Rectan-| 5X8 or 1oX8.
| velar lappets. lar. | truncated. Rec- | gular.
| tangular.
Length of r (rear, r r 2r —r
mouth-arms.
Length of ter- | 2r r 2r r 4to6r One-sixth r.
| minal clubs.
| Color. Bell blue, greenish.) Bell blue, greenish,| Bell yellow (?) Bell reddish- Bell brown ?
olive, or brown; | olive or brown; | with orange (?)) brownwithring-| with darker
with white, with white, spots. Canals like spots of white} margin.
brown, or yellow-) brown, or yellow-| and clubs violet | and brown. White spots
ish spots. ish spots. to rose-colored. ringed with
black.
Where found | Fiji Islands, Zanzibar, East Malay Archi- Indian Ocean, Samoa. Red Sea. Dis-
and remarks.| Japan, Malay Africa. pelago. Hongkong, tinguished by
Archipelago, Cocos Islands. long filaments
Indian Ocean. on arm-disk.

Mastigias papua L. Agassiz.

Cephea papua, Lesson, 1829, Voyage de la Coquille, Zooph., p. 122, planche 11, figs. 2, 3.

Cephea papuensis, Grirritu, 1832, Cuvier’s Animal Kingdom, plate 3, fig. 3-

Mastigias papua, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 152.—Haercket, 1880, Syst. der Medusen, p. 623.—Maas,
1903, Scyphomedusen der Siboga Expedition, Monog. 11, pp. 66, 69, taf. 7, fign. 60, 62, 63; taf. 12, fig. 111.

Pseudorhiza thocambaui, AGassiz, A., and Mayer, 1899, Bull. Museum. Comp. Zool. at Harvard College, vol. 32, p. 173, plate
13, figs. 40-44 (young stages).

Mastygias papua, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Valdivia, Bd. 3, Lfg. 1, p. 47, taf. 4, fign.

17-19.
Mastigias physophora, Kisuinovye, 1895, Zoolog. Magazine, Tokyo, vol. 7, No. 78, 3 pp., plate 13, figs. 1-13.—Scuuttze, L. S.,
1898, Denkschr. Med. Nat. Gesell. Jena, Bd. 8, p. 443.
Mastigias physophora, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedition, Dampfer Valdivia, Bd. 3, Lfg. 1, p. 49-
Mastigias papua var. physophora, Maas, 1909, Abhandl. Akad. Wissen. Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 46.

Bell 30 to 80 mm. wide, usually hemispherical, sometimes flatter and sometimes fuller
than a hemisphere. Gelatinous substance frm. Exumbrella surface with very fine granu-
lations. 8 rhopalia, each with a pigmented mass of crystalline concretions and a shallow,
exumbrella, sensory pit without furrows. 80 marginal lappets. In each octant 2 small,
pointed, ocular and 8 larger, rounded velar lappets with deep furrows between them extending
a short distance up the sides of the exumbrella. Arm-disk somewhat wider than bell-radius.
The 4 subgenital ostia are about twice as wide as the columns between them and are each
somewhat constricted in the middle. Subgenital porticus unitary. 8 mouth-arms, each about
as long as bell-radius. The simple upper part 1.5 times as long as the 3-winged lower portion
ofthe arm. The frilled mouths are developed not only along the edges of the 3 wings, but also
for some distance inward along the sides of each leaf. Each mouth-arm usually terminates
at its lower end in a club-like filament which may be as long as the diameter of the bell, but
is sometimes reduced in size or even wholly absent. This club is triangular in cross-section
and contains an axial canal. A large number of small, club-shaped vesicles arise from between

RHIZOSTOM#—MASTIGIAS. 679

the mouths on the outer sides of the mouth-arms. The central canal of each mouth-arm gives
off 3 side branches which lead to the 3 rows of frilled mouths of the winged, lower part of the
arm. All 3 of these canals fuse again with the axial-canal at the base of the terminal club and
extend onward as the axial-canal of the club.

The central stomach is cruciform and gives off 8 straight radial-canals which extend to
the sense-organs. ‘These canals are all connected by a wide ring-canal in a zone at a consider-
able distance inward from the margin. About 7 to 9 anastomosing radial-canals arise from
the stomach in each octant between the rhopalar canals and fuse with the ring-canal. On its
outer side the ring-canal gives off a fine-meshed network of anastomosing vessels which extend
into the lappets and fuse with the outer ends of the 8 rhopalar canals. “The circular muscles
of the marginal zone of the subumbrella are widely interrupted in the 8 principal (rhopalar)
radi. The gonads are 4 folded walls forming cruciform sides of the subgenital porticus.

Color quite variable. Bell and mouth-arms usually greenish-blue, or olive-green to
olive-brown, and there are a number of yellow, white, or occasionally brown, blue, or green
oval spots over the exumbrella, especially near the margin. ‘The frills of the mouths may be
olive, greenish-blue, yellowish-green, or brown. ‘The 8 rhopalar radial-canals are darker.

This medusa is widely distributed over the Malay Archipelago, Indian Ocean, and
China Sea to Japan, and outward over the Pacific to the Fiji Islands.

Agassiz and Mayer found an ephyra of the medusa in Suva Harbor, Fiji Islands, in Jan-
uary, 1898. It was 5 mm. in diameter and quite flat and disk-shaped. There were 8 marginal
sense-organs. The central mass of dark-brown entodermal pigment granules of the sense-organ
was developed, but the peripheral shell of transparent granules had not yet made its appear-
ance. There were 24 marginal lappets, the 16 ocular lappets being about twice as long as the 8
velar lappets. There were 16 radial-pouches from the stomach, of which 8 went to the sense-
organs and 8 to the velar lappets. The sub-
genital porticus was already unitary and the
brachial disk was suspended from the floor of
the subumbrella by means of 4 gelatinous pillars,
exactly as in the adult. The ephyra possessed
only a simple, central mouth opening, having 4
cruciform lips. The margins of the lips were
lined with a row of short, slender tentacles
with knob-like ends exactly like those that sur-
round the mouths on the mouth-arms of the
adult medusa. No trace of the genital organs

Fic. 415.—Mastigias papua, after Vanhofien, could be detected, but the gastric cirri were

ao ites ped ik represented by 12 short filaments (3 in each
quadrant). The color of the ephyra was very similar to that of the adult. The ring-canal had
not yet begun to develop. ; ;

Mastigias papua swims very rapidly by an incessant contraction and expansion of the
bell-rim. Being an abundant and variable form, it has given rise to many nearly related
varieties, such as M. papua var. siboga of the Malay Archipelago and M. stderea of the east
coast of Africa. M. physophora Kishinouye is another variety found abundantly off the
coasts of Shima and Sagami, Japan, during summer and autumn. Its bell is at least 100 mm.
in diameter and is light-brown with numerous, round, dark-brown spots near the margin.

Schultze finds this medusa at Amboina, Moluccas, in January and February, and it is
evidently only a large, dark-colored variety of M. papua. Kishinouye gives an excellent
series of drawings of this medusa.

Mastigias papua var. siderea.

Mastigias siderea, Cuun, 1896, Jahrb. Wissen. Anstalten Hamburg, Jahrg. 13, p- 13, taf. 1, fig. 3.

Mastigias, sp., (young medusa ?), Scuuttzr, 1898, Abhandl. Senckenberg, Gesell., Bd. 24, Heft. 2, p. 161, taf. 15, fig. 2.
Mastygias siderea, VANHOFFEN, 1902, Wissen. Ergeb. deutsch. Tiefsee Exped., Dampfer Valdivia, Bd. 3, Lfg. 1, p. 49-
(?) Mastigias siderea, Scnuttze, L. S., 1898, Abhandl. Senckenberg. Gesell. Naturf., Bd. 24, p. 161, taf. 15, fig. 2.

(?) Eucrambessa miilleri, Hancxer, 1880, Syst. der Medusen, p. 624.

(?) Mastygias miilleri, VANHGFFEN, 1902, Wissen. Ergeb. Valdivia Exped., Bd. 3, Lfg. 1., p. 49-

Bell flatly rounded, 70 mm. wide. 8 marginal sense-organs and 80 marginal lappets.
Ocular lappets narrow, but the 8 intermediate lappets in each octant are semicircular in outline,

680 MEDUSZ OF THE WORLD.

Arm-disk wider than bell-radius, and the subgenital ostia are twice as wide as the radial
supports between them. 8 wide, ocular radial-canals and 7 anastomosing, radial vessels in
each octant. Mouth-arms are twice as long as bell-radius, the simple upper part of the arm
being somewhat shorter than the lower, 3-winged part. Each arm terminates in a single club
as long as the bell-radius.

Bell light yellowish-brown with round white spots, which are largest over the ring-canal
and smaller near the margin, where they are arranged in 3 or 4 radiating rows between each
successive pair of marginal sense-organs. 8 blackish streaks along the 8 ocular radial-canals
on the subumbrella, and also white specks in each octant of the subumbrella between the
stomach-pouches and the circular furrow. Arms brown with small white spots. Filaments
yellowish.

Found along the Zanzibar coast, East Africa, in August and September, and in the western
parts of the Indian Ocean.

Chun, 1896, gives a detailed description of the adult and the young of this species.

Haeckel’s Eucrambessa miiller: from Madagascar is probably identical with this species,
but is so imperfectly described that we will never be able to determine it with certainty.

Mastigias papua var. siboge Maas.

Mastigias papua var. siboge, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 66, taf. 6, fign. 54-57; taf. 7,

fign. 58, 59, 61, 64; taf. 8, fign. 75-77; taf. 9, fign. 84, 85; taf. 12, fig. 110.

Bell massive, rounded, and when mature 120 mm. in diameter. There are about 9
rectangular velar lappets with rounded angles in each octant between sense-organs. The
4 interradial ostia of the subgenital porticus are 3 times as wide as the columns between them
There are 7 to 10 anastomosing radial-canals between each successive pair of rhopalar canals.
Mouth-arms as long as the bell-radius. There are numerous small, rounded clubs upon
each of the mouth-arms and also a terminal appendage, which is triangular in cross-section
and nearly as long as the bell-diameter.

The ground color is yellowish (?) with orange (?) spots. There are no ring-shaped
spots upon the exumbrella. There are 8 violet radial bands upon the rhopalar canals. The
terminal appendages of the mouth-arms are sprinkled over with violet spots. The canal-
system is rose-colored and the gonads are orange.

This variety is found in the Malay Archipelago, and is described in detail by Maas, 1903.
It is distinguished from the typical M. papua by the absence of “eye spots” upon the exum-
brella, by its yellow or orange color, and by its nearly rectangular velar lappets.

Mastigias ocellata Haeckel.

Medusa ocellata, Mopeer, 1791, Nova. Acta. Phys. Med., N. C., tome 8, Append., p. 27.

Cephea ocellata, PEron et Lesueur, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 361.

Cephea ocellata+ Hidroticus rufus, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 156, 158.

Mastigias ocellata, Harckxer, 1880, Syst. der Medusen, p. 623.—VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp-
33) 44, taf. 5, fign. 3-6; 1902, Wissen. Ergeb. Valdivia Exped., Bd. 3, Lfg. 1, p. 49—Maas, 1902, Scyphomedusen Siboga
Exped., Monog. 11, p. 63.

This medusa is distinguished from M. papua by the peculiar “eye spots” on the exum-
brella. These may be described as white circles with a brown center and brown rim. There
are also other simple brown spots on the exumbrella. The mouth-arms are shorter than in
M. papua, being shorter than the bell-radius, and the terminal clubs are not longer than the
bell-radius. Velar lappets more numerous than M. papua, there being about 12 rounded
velar lappets between 2 narrow, pointed, prominently projecting, ocular lappets in each octant.
There are 15 to 20 anastomosing radial-canals in each octant between the rhopalar canals,
instead of about 7 to 9 as in M. papua.

General color reddish, with numerous white, brown-rimmed, and centered “eye spots”
on the exumbrella. Tips of terminal club blue.

[he medusa becomes about 50 to 60 mm. wide and is found in the eastern parts of the
Indian Ocean and in the China Sea. Straits of Sunda, Hongkong in October, Cocos, and
Philippine Islands. The Albatross found small meduse in March and April, and a mature one
in January in the Philippines, in 1908.

RHIZOSTOMA—MASTIGIAS. 681

Mastigias pantherina Haeckel.

Mastigias pantherina, HarcKe1, 1880, Syst. der Medusen, p. 624.—Vanunorren, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3,
P- 445 1902, Wissen. Ergeb. Valdivia Exped., Bd. 3, Lfg. 1, p. 49. :
Mastigias papua, Maas, 1903, Scyphomedusen der Siboga Exped., Monog. 11, p. 63.

This is known only from a preserved specimen, briefly described by Haeckel. It appears
to be related to, if not identical with, M. ocellata, having the same peculiar “‘eye spots” on the
exumbrella. The velar lappets are said to be truncated aa rectangular and to be more numer-
ous than in M. ocellata, there being 16 in each octant instead of about 12, as in M. ocellata.
Mouth-arms are much longer than in M. ocellata, being nearly as long as bell-diameter. The
simple upper part of the arm is hardly half as long as the 3-winged lower part, whereas in
M. ocellata and M. papua the upper part is longer than the lowest part of the arm. Terminal
club very much longer than in other forms of Mastigias, being 2 or 3 times as long as the
bell-diameter.

Bell dark-brown with white spots ringed with black. Bell-margin black.

Found at Samoa, tropical Pacific.

Mastigias gracile.

Desmostoma gracile, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft 3, pp. 35, 45, taf. 4, fign. 5-7.

Disk flatly rounded or hat-shaped, 35 mm. wide, thin at margin, but very thick at apex.
Exumbrella besprinkled with irregularly placed clusters of small warts. 8 marginal sense-
organs, the marginal lappets irregularly arranged, there being 5 or 10 rectangular velar lappets
in various octants; thus some marginal sense-organs may be close together, while others are
far apart. There is a wide, well- -developed zone of ring-muscles, Eanhnedl however, to the
peripheral parts of the subumbrella, its inner edge being beyond the periphery of the arm-
pllars. The subgenital ostia are twice as eades as pillars of the arm-disk. These ostia
appear double, thus giving the false appearance of 8 instead of 4 genital pits. This is due to
the fact that each of the 4 interradial gonads is separated into 2 lateral rays by means of a cen-
tral gelatinous flap which divides the subgenital ostium into 2 side-openings.

The 8 mouth-arms are hardly as long as the bell-radius and consist of a simple, short,
thick, upper part of the arm and an expanded 3-winged, lower part, which is 3 to 4 times as
long as the upper. The lower part is thickly beset with frilled mouths, there being short,
isolated, gelatinous knobs strewn between the mouths and a short, rounded terminal knob
at the free end of each arm, about one-sixth as long as the arm itself. There is also a large
cluster of about 8 to 20 linear filaments upon the arm-disk at the bases of the 8 mouth-arms.
These are about 1.5 times as long as the diameter of the bell.

Central stomach large and cruciform, 8 radial-canals arise from it and extend outward
to the marginal sense-organs; of these the 4 interradial canals are long and the 4 perradial
ones short. Between these 8 main canals are numerous, slender canals which arise from the
periphery of the stomach and anastomose with themselves and with the main radial-canals.
All of these canals are set into communication one with another by means of the circular
canal near the bell-margin. On its outer side the ring-canal gives off a network of slender
vessels which anastomose over the lappets. The subgenital porticus is very small. Color ( ?)

Vanhoffen describes 3 examples of this medusa fan Assab, on the Red Sea. They were
found in September. He gives it the generic name Desmatonn defining the genus as being
similar to Mastrgias, but with a large cluster of filaments upon the arm-disk between the
mouth-arms. The distinction appears to me to be too slight for generic, although important
for specific, differentiation.

Mastigias (?) rosea Vanhoffen.

Rhizostoma rosea, REYNAUD, 1830, in Lesson’s “Centurie zoologique,” p. 97, planche 34.

Toxoclytus roseus, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 153.—Harcxet, 1880, Syst. der Medusen, p. 586.—
LENDENFELD, VON, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 288.

Mastigias roseus, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 45.

Disk flat and hat-shaped, short, deep radial furrows on the exumbrella surface between
the lappets; 56 to 64 (?) small, elongate, marginal lappets, all of the same size and shape.
8 separate mouth-arms, hardly as long as the bell-radius; upper part of each arm cylindrical,

682 MEDUSZ OF THE WORLD,

lower half pyramidal and 3-winged, one wing centripetal, the other two centrifugal; outer
surfaces of all 3 wings covered with deep furrows which are complexly folded and contain
the frilled mouths. A club-shaped, gelatinous appendage arises from the free lower end of
each arm and numerous other appendages spring from the sides of the arms between the
furrows. No scapulets. 4 horseshoe-shaped gonads.

The bell-margin and frills of the mouth-arms are deep rose color, while the gonads are
paler. This species is found in the tropical Atlantic.

Size (?) Exact locality (?)) A more accurate and modern figure of this form is greatly
to be desired for we can not now be certain even of the generic position of this medusa.

Genus PSEUDORHIZA von Lendenfeld, 1882.

Pseudorhiza, VON LENDENFELD, 1882, Zool. Anzeiger, Bd. 5, p. 380; 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 293;
1887, Descriptive Catalogue Australian Museum, Sydney, part 1, p. 23.—Maas, 1903, Scyphomedusen der Siboga Expedi-
tion, Monog. 11, pp. 53, 71, 80.

Monorhiza, Haacke, 1887, Jena Zeit. fiir Naturw., Bd. 20, p. 614.

The type species is Pseudorhiza aurosa yon Lendenfeld, of Victoria and South Australia.

GENERIC CHARACTERS.

Rhizostomata triptera with eight 3-leaved mouth-arms, one or all of which terminates
ina ais large club. No other clubs or filaments among the mouths. 8 rhopalia. 16 radial-
canals, 8 rhopalar and 8 adradial. The rhopalar canals extend to the bell-margin, but the
adradial ones only to the ring-canal which connects with all of the 16 radial-canals. On the
outer side the ring-canal gives off an anastomosing network, and on its inner side a number
of blindly-ending, centripetal vessels which may anastomose. The wide, circular muscle
of the subumbrella is only partially interrupted in the radii of the radial-canals. The sense-
club bears an ocellus and there is a shallow, exumbrella sensory pit without radiating furrows.
The central mouth persists at the center of the arm-disk.

Haacke’s genus Monorhiza is similar to Pseudorhiza, but in Monorhiza only one mouth-
arm bears a terminal club, whereas all 8 bear each a terminal club in Pseudorhiza. The
distinction may be deemed to be of specific rather than of generic value.

Pseudorhiza is so closely allied to Mastigias that we might readily merge the two genera
into one, designating it by the older name Mastrgtas. In Mastigi1as, however, one finds small
clubs or filaments arising from the sides of the mouth-arms between the mouths, and these are
wholly absent in Pseudorhiza; also in Mastrgias the inter-rhopalar canals which arise from
the inner side of the ring-canal connect with the stomach, whereas in Pseudorhiza they end
blindly. These blindly-ending, centripetal canals on the inner side of the ring-canal in Pseu-
dorhiza may or may not anastomose. They appear not to anastomose in Haacke’s P. haeckelt1,
but in von Lendenfeld’s P. aurosa they are said to form a network; yet von Lendenfeld believes
these medusz to be identical, and they are certainly closely related.

The terminal club in all Rhrzostomata is merely the naked extension of the axial shaft
of the mouth-arm. It is triangular in cross-section as is the arm itself, and the axial duct of
the arm extends onward into it. Thus it is not homologous with the club-like appendages and
filaments which arise between the mouth-arms.

Pseudorhiza aurosa von Lendenfeld.
Pseudorhiza aurosa, VON LenpDENFELD, 1882, Zool. Anzeiger, Bd. 5, p. 380; 1884, Proc. Linnean Soc. New South Wales, vol. 9,

p- 293, plate 3, 1 fig.; Ibid., p. 426.—1887, Descriptive Catalogue Australian Museum, Sydney, Part 1, p. 23; 1888,

Zeit. fiir wissen Zool., Bd. 47, p. 218, taf. 18, fig. 1; taf. 19, fign. 3-7; taf. 20, fign. 14-16; taf. 23, fign. 39, 42, 43, 49-52,

57; taf. 24, fign. 60, 62; taf. 26, fign. 84, 87-92, 94, 95; taf. 27, fign. 98-107, 114 (detailed description).

Umbrella 400 mm. wide, flatly rounded, about 130 mm. high. Exumbrella rough and
reticulate. 8 marginal sense-organs and in each octant there are 2 long, narrow, pointed,
ocular lappets and 6 velar lappets. Each of the velar lobes consists of 3 secondary lappets.
lhe arm-disk is about as wide as the radius of the umbrella and gives rise to 8 mouth-arms
which are about as long as the diameter of the umbrella. There is a central mouth on the
(lower) subumbrella side of the arm-disk and 4 pairs (8) of deep gutters extend out from this

RHIZOSTOM2—PSEUDORHIZA. 683

mouth along the lower side of the 8 mouth-arms. These arms are 3-leaved and the free edges
of these leaf-like expansions branch profusely and complexly. The 8 club-shaped axial appen-
dages which arise from the lower ends of the 8 arms are each about as long as the diameter of
the disk.

The 4 subgenital ostia are somewhat wider than the supports between them. There
is a single subgenital cavity. The cruciform, central stomach gives rise to 16 radiating
canals, 8 to the sense-organs and 8 to the intermediate positions. These 16 canals are put
into connection one with "omnes by a ring-canal. On the outer side of this ring-canal is an
anastomozing network of vessels, and extending inward from the ring-canal are 160 blindly-
ending, centripetal vessels, 10 Reevecn each pair of adjacent radial- Sonal

Umbrella colorless, the valleys of the reticulate elevations of the exumbrella violet. The
entoderm of gastrovascular cavity brown. Upper parts of mouth-arm grooves rose-colored.
Arms colorless and transparent. Mouth-frills along the margins of the grooves and distal
ends of the long axial mouth-arm clubs rich violet.

Found at Port Philip, Victoria, and at Adelaide, South Australia.

Described in detail by von Lendenfeld, in Zeit. wissen. Zool. It differs from Haacke’s
“Monorhiza” in that there are 8 moderately long, mouth-arm filaments, instead of only one
very long filament, and the centripetal canals anastomose into a network on the inner side
of the ring-canal instead of remaining separate, as in Haacke’s medusa.

Pseudorhiza haeckelii Haacke.

Pseudorhiza haeckelisz, Haacke, 1884, Biol. Centralbl., Bd. 4, p. 291.
Monorhiza haeckelii, Haacke, 1887, Jena. Zeit. fiir Naturwissen, Bd. 20, p. 614, taf. 37, fign. 1-9.

Disk hemispherical to hat-shaped, 200 to 250 mm. wide and 50 to 100 mm. high.
Exumbrella roughened with polygonal, wart-like reticulations. 4 elongate, wart-like pro-
tuberances upon the subumbrella in the 4 interradit beyond the 4 subgenital ostia. 8 marginal
sense-organs flanked by 16 short, narrow, sharp- pointed lappets. The 48 velar lappets are
wide, ehore and rounded. There are thus 8 marginal sense-organs and 64 lappets. The
central mouth opening is 4-cornered, and the central disk gives rise to 4 pairs of laterally
compressed 3-leaved mouth-arms. Each leaf of these mouth-arms gives rise to many flat,
fern-like expansions. A single filament, 300 mm. long, arises from the lower end of one
of the mouth-arms; it is spindle-shaped and 3-cornered in cross-section. The 4 subgeni-
tal ostia are wider than the gelatinous columns between them. The central stomach is
Maltese-cross-shaped and gives rise to 16 canals, 8 extending outward in the radu of the
marginal sense-organs, and 8 being adradial. These 8 ocular canals extend outward to the
rhopalia, but the adradial canals end in the ring-canal near the middle zone of the subumbrella.
This ring-canal gives rise in each octant to about 18 narrow, unbranched, non-anastomosing,
centripetal canals which end blindly. On the outer side of the ring-canal is a network of
anastomosing vessels which fuse with the 8 rhopalar radial-canals. The circular muscles of
the subumbrella are well- -developed over the peripheral half of the under side of the bell and
are only partially interrupted in the 8 principal radu.

The gonads form 4 U -shaped walls of the subgenital porticus and are much folded. The
gastric filaments are so inconspicuous that Haacke failed to find them, although according to
von Lendenfeld they are present. Like Chrysaora, Pseudorhiza haeckelit 1s hermaphroditic,
for in addition to the central gonads there are sporadic spermaries situated in root- like ento-
dermal filaments in the “‘gutters” or food crevices of the mouth-arms.

The furrowed network of the exumbrella is violet-brown and the entoderm of the gutters
of the mouth-arms dark-red. The large filament glistens in metallic copper- -color, and in
young specimens it is blue-violet at the base, with a dark-colored, blue and red, spiral, ento-
eceal band of color extending throughout its central cavity. The peripheral canal-system
of the subumbrella is brownish-red.

Found in the Gulf of St. Vincent, South Australia, and described in detail by Haacke,
1887. Von Lendenfeld regards this medusa as being identical with, or only a variety of, his
Pseudorhiza aurosa.

684 MEDUS.E OF THE WORLD.

Genus PHYLLORHIZA L. Agassiz, 1862.
Phyllorhiza, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 158.—Harcxex, 1880, Syst. der Medusen, p. 588.—von Len-

penrecp, 1884, Proc. Linnean Soc. New South Wales, vol. 9, p. 296.—Vanunirren, 1888, Bibliotheca Zoologica, Bd. 1,

Heft. 3, p- 41.

Phyllorhiza chinensis L. Agassiz and P. trifolium Haeckel are too imperfectly described
to be recognizable, and P. punctata von Lendenfeld, from Australia, is the type species and
the only adequately determined form.

Phyllorhiza chinensis is probably Cephea cephea, and P. trifolium is, according to Haeckel,
described from a preserved and mutilated specimen. It has 96 marginal lappets. Exumbrella
finely granular. Arms with 3 semicircular, pinnately-branched, separated lappets or wings,
with 24 long and numerous, short filaments, the largest being equal to the bell-diameter. Bell
75 mm. wide. Japanese Sea.

GENERIC CHARACTERS.

Similar to the closely allied Lychnorhiza but the centripetal vessels which arise from the
inner side of the ring-canal join with the central stomach, as in Mastigtas, instead of ending
blindly as in Lye hnorhiza. Also the ring-muscle of the subumbrella is interrupted in the 8
principal radii. The canal-system resembles that of Mastigias, but the mouth-arms have
no terminal clubs.

Phyllorhiza punctata von Lendenfeld.

Phyllorhiza punctata, Von Lenvenrevp, 1884, Zool. Anzeiger, Bd. 7, p. 429 (development).—1884, Proc. Linnean Soc. New

South Wales, vol. 9, p. 296, plate 4, 1 fig.; p. 307, plate 5, figs. 1-4; 1888, Zeit. fiir wissen. Zool., Bd. 47, p. 223, taf. 18,

fig. 2; taf. 19, fign. 8, 9, 11, 12; taf. 21, fign. 17-20, 22; taf. 22, fign. 27-35, taf. 23, fign. 40, 53-55; taf. 26, fign. 79,

80, 86; taf. 27, fign. 110, 117 (detailed description).—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 60.

Umbrella somewhat flatter than a hemisphere, 500 mm. wide. Exumbrella 50 mm.
thick, with a finely granular surface. 8 marginal sense-organs. In each octant there are
2 sickle-shaped ocular lappets, 4 simple lappets near the ocular lappets, and 4 double lappets
near the middle of each octant. There are thus 14 lappets in each octant, considering each
double-lappet as two. The large, double lappets are partially fused by a basal web, and the
ocular lappets are only half as wide as the others and are sharp-pointed, while the others are
rounded. Radial furrows extend centripetally over the exumbrella between the lappets. The
arm-disk is thick and a little wider than the bell-radius, octagonal in shape and has a canal-
system of its own. It is thickly beset with filaments on its ventral side, to which young embryos
in the gastrula stage adhere. The 4 subgenital ostia are oval and more than twice as wide as
the pillars between them. The eight 3-leaved mouth-arms are two-thirds as long as diameter
of umbrella. Their 3 mouth-bearing edges branch pinnately, but the pinnz are only rudi-
mentary. The lower parts of the mouth-arms bear numerous, tapering, bluntly-ending fila-
ments, some of which are two-thirds as long as the mouth-arms themselves. The circular
muscles of the subumbrella are interrupted in the 8 principal radii.

8 radial-canals, 4 perradial and 4 interradial, arise from the cruciform, central stomach
and extend to the 8 marginal sense-clubs. A wide ring-canal, at some distance inward from
the margin, connects all 8 radial-canals. On its outer side the ring-canal gives rise to a fine-
meshed network of vessels (which fuse also with the radial-canals) extending into the lappet
zone. On its inner side the ring-canal also gives off a similar network of vessels which connects
with the central stomach and with the 4 interradial, but not with the 4 perradial, canals.

The arm-disk, main stems, branches of the mouth-arms, and gelatinous substance of
the umbrella are colorless. In the gelatinous substance of the umbrella, close to the surface,
are groups of unicellular, yellow, plant cells which give the whole surface a brown color. Also
in the gelatinous substance, close to the surface of the exumbrella, there are cloud-like masses
of minute, highly refractive bodies which give the medusa a spotted appearance, the spots being
whitish. The frilled mouths are brown and the filaments colorless.

This medusa is found in Port Jackson, New South Wales, Australia, and is described
in elaborate detail by von Lendenfeld (Zeit. fiir wissen. Zool.). He finds that during develop-
ment the marginal sense-organs decrease from 24 to 16, and finally to 8.

When the medusa is 15 mm. wide there are 24 marginal sense-organs and 48 marginal
lappets. The 2 ocular lappets of each octant are sharp-pointed, narrow, and elongate, while

RHIZOSTOM®—PHYLLORHIZA, VERSURA, 685

the 4 intermediate marginal lappets are broad and bluntly rounded. The 2 intermediate
sense-organs of each octant lie in the clefts of the 2 lappets on both sides of the central fissure
of each octant.

When the meduse are about 30 mm. wide the 16 intermediate sense-organs disappear
and a new set of 8 sense-organs develops in the middle cleft of each octant. The medusa
now has 64 lappets and 16 marginal sense-organs. The 2 lappets on both sides of the 8 velar
sense-organs are now double and finally divide completely. When 50 mm. wide the meduse
lose their § intermediate sense-organs, and there are then 8 radial and interradial rhopalia and
80 marginal lappets. von Lendenfeld’s observations of this remarkable process of development
await confirmation.

Genus VERSURA Haeckel, 1880.

Crossostoma, used for Mollusca by Morris and Lycert, 1850.
Crossostoma, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 155.—Maas, 1903, Scyphomedusen der Siboga Exped., Monog.
II, pp. 54, 81.
Versura+ Crossostoma, Harcket, 1880, Syst. der Medusen, pp. 606, 607.
The older species are inadequately described and we may designate /’. palmata
Haeckel, from the Malay Archipelago, as the type of the genus.

GENERIC CHARACTERS.

Rhizostomata triptera with clubs and filaments upon the mouth-arms. The 4 perradial
canals arise directly from the stomach, but the 4 interradial canals result from the fusion of a
number of anastomosing vessels which arise from the interradial sides of the stomach. There
is no definite ring-canal, but merely a marginal network of vessels. There are no radial-
muscles in the subumbrella, but the ring-muscles are well-developed.

Among characters of minor importance, the subgenital ostia are wide openings, wider
than the columns between them, and the sense-organs have a simple, exumbrella pit without
radiating furrows. At the center of the arm-disk is a prominent, raised cluster of frilled
mouths having filaments between them.

The older species are so imperfectly described that it will be hopeless to attempt to
determine them and they had best be omitted from further consideration. For example:
““Crossostoma corolliflora”’ Haeckel is probably a Cotylorhiza; “C. dubreuillir’”’ may be a Cato-
stylus, and “C. frondifera’”’ may be a Casstopea (see Haeckel, Syst. der Medusen, pp. 608,
609).

Versura palmata Haeckel.

Versura palmata, Harcxet, 1880, Syst. der Medusen, p. 606, taf. 40, fign. 9-12.—Goetre, 1886, Sitzungsber. Akad. Wissen.
Berlin, Jahrg. 1886, p. 836.—Vanuorren, 1888, Bibliotheca, Bd. 1, Heft. 3, p. 42.—Hamann, 1881, Jena. Zeit. fur Natur-
wissen., Bd. 15, p. 253.

Bell flat, shield-shaped, 60 mm. in diameter, 20 mm. high. 8 rhopalia set within very
shallow niches in the bell-margin. Ocular lappets very small and sharply pointed; the velar
lappets vary greatly in number, ranging from about 4 double ones to 12 in each octant. They
are barely ‘discernible, being separated by very short, narrow clefts. Arm-disk about two-
thirds as wide as bell- panes the 4 subgenital ostia are twice as wide as the perradial columns
between them. There is a unitary, narrow, cruciform subgenital cavity. The 8 mouth-arms
are somewhat shorter than the bell-radius. The simple upper axial shaft of each arm is not
quite half as long as the 3-winged lower part, which is Y-shaped in cross-section and nearly
as wide as long. The free edges of the Y are complexly branched and folded and bear the
frilled mouths. Each lamella of the mouth-arm displays 6 to 7 very deep clefts. There are
numerous, small, club-shaped vesicles scattered among the mouths and a larger terminal club
at the lower end of each arm.

The central stomach is Maltese-cross-shaped. Canal-system of bell (?) There is a
marginal zone of circular muscles which are only partially interrupted in the 8 principal radii.
Centripetal to this zone is another muscular zone which is interrupted in the 8 principal
radii, the fibers of which are bowed outward toward the areas of interruption. Color (?)

Haeckel records this medusa from the Malay Archipelago, and Goette studied specimens
from Zanzibar, Singapore, and Nagasaki, Japan. It is distinguished only by its indistinct
velar lappets and the terminal club of its mouth-arms.

686 MEDUS.X OF THE WORLD.

Versura vesicata.
Versura vesicata, HarcKeL, 1880, Syst. der Medusen, p. 645.

This is very briefly mentioned by Haeckel. It is closely related to, if not identical with,
I’, palmata, but has twice as many velar lappets. The ocular clefts in the margin are deep
and the subgenital ostia are only half as wide as the pillars between them. The vesicular
club at the end of each arm is larger than in /’. palmata, being one-fourth as long as the bell-
radius. Northwestern coast (?) of Australia. Size, etc. (? )

Versura pinnata Haeckel.
Versura pinnata, Hace, 1880, Syst. der Medusen, p. 607.

This Cocos Island medusa is very briefly described by Haeckel. Bell flat, 80 mm. wide.
8 deeply-cleft rhopalar niches. 144 lappets. In each octant 16 indistinct quadratic velar,
between 12 small, pointed, ocular lappets. Subgenital ostia as wide as the columns between
them. Mouth-arms somewhat longer than bell-radius and twice as long as wide. It may be
identical with /’. palmata, being described only from a preserved and presumably contracted
specimen.
Versura anadyomene.

Crossostoma anadyomene, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 56, taf. 7, fign. 65-68.
Crossostoma, sp., Gorrre, 1886, Sitzungsber. Akad. Wissen., Berlin, Jahrg. 1886, p. 837.

Bell about 200 mm. in diameter and quite flat. Exumbrella covered with a network
of anastomosing furrows, leaving polygonal elevations between them. This network is wide
at the center and finer-meshed at the margin, and the general trend of the furrows is mainly
outward from the center. Gelatinous substance of bell very thin, being only a few millimeters
thick even at center. The canal-system can be seen by looking through the bell from the
exumbrella side. There are 8 marginal sense-organs, 4 perradial and 4 interradial. These
are small and probably lack ocelli, and on the exumbrella side above each sense-organ there
is a small, simple sensory pit without radiating furrows. The 16 ocular lappets which flank
the 8 marginal sense-organs are small and lanceolate. In each octant between sense-organs
are 8 large, semicircular velar lappets, which alternate somewhat irregularly with about 8
small, narrow, tongue-shaped lappets. The arm-disk is rectangular to cruciform, with long
interradial slit-like, subgenital ostia and 4 narrow, perradial pillars between them. The
subgenital sinus is a flat, narrow, cruciform space and the genital cross is very narrow. The
medusa is very delicately formed, the lower parts of the mouth-arms being thin and leaf-like.

Each of the 8 mouth-arms is about as long as the bell-radius and very strongly compressed
laterally. The upper part of the arm has the form of a knife blade, the sharp edge being
inwards (axial) and the thick, rounded side being abaxial. Two rhomboidal, expanded,
leaf-like wings arise from the abaxial side of each arm. The lower sides of the two lateral
wings and the inner (axial) part of each arm gives rise to numerous, flat, membranous side
branches which expand outwardly. These side branches bear the frilled mouths. Small
club-shaped vesicles arise from between the mouths of the two lateral, abaxial wings of each
mouth-arm, while the ventral side of each mouth-arm gives rise solely to tapering filaments
between the mouths. The rows of frilled mouths on the inner (axial) sides of the mouth-arms
extend to the center of the arm-disk, where they form a projecting rosette.

A main canal arises from each of the 4 perradial corners of the stomach and sends branches
into the mouth-arms. It is remarkable that each of the lateral, abaxial wings of the lower
arms contain 2 separate axial-canals, each of which sends off side branches to the mouths.
These side branches do not anastomose and thus there is a double canal-system in each of
the lateral wings of the lower arms.

The central stomach is cruciform, the arms of the cross being perradial. 4 perradial
canals extend uninterruptedly from the 4 angles of the central stomach to the 4 perradial
sense-organs. The 4 canals to the interradial sense-organs do not arise directly from the
stomach, but from 4 areas of anastomosing vessels which form a network on the interradial
sides of the central stomach. These network-like areas of vessels arise in numerous canals
from the entire interradial sides of the stomach. They send out a few branches which join

RHIZOSTOM-E—VERSURA, 687
the 4 perradial canals. There is no true ring-canal, although all 8 of the radial-canals are
placed in communication one with another by a marginal zone of anastomosing vessels which
extend into the lappets. There are no radial-muscles in the subumbrella, but there are 2
separate concentric ring-muscles, which are further divided into 8 sectors, the lines of separa-
tion being in the radii of the 8 radial-canals. The outer zone of ring-muscles lies close to the
bases of the marginal lappets. The inner zone is not a true circle but is widest in the 4 interradii
and narrowest in the 4 perradial lines. The gonads and canal-system appear to be reddish.

This medusa is described by Maas from a single specimen found in the Malay Archi-
pelago by the S:boga expedition. It may be identical with the Versura briefly described by
Goette, 1886 (Sitzungsber. Akad. Wissen. Berlin, Jahrg. 1886, p. 837), from the east coast
of Africa. Maas gives a detailed description of the medusa. The species is distinguished
by its prominent velar lappets, its very thin, delicately formed bell and mouth-arms, and the
furrowed surface of its exumbrella.

Versura maasi, sp. nov.

Named in honor of Prof. Dr. Otto Maas in recognition of his notable researches upon
medusz.

Bell 90 mm. wide, flatter than a hemisphere and evenly rounded. Exumbrella finely
granular, without furrows. Gelatinous substance fairly thick but not very rigid. 8 rhopalia,
each with a pigment mass and an exumbrella pit with smooth floor. 112 marginal lappets.

Sy

WS
SS)

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246) UE
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eae (2

Fic. 416.—Versura maasii. Drawn by the author, from a specimen obtained by the U.S.
Fisheries Bureau steamer 4/batross in the Philippine Islands.

A, oral view showing 6 of the mouth-arms cut off close to the arm-disk. In the lower sector the
circular muscles are removed exposing the canal-system. B, magnified view of one of
the clubs from between mouths of the mouth-arms. C, mouth-arm seen from the
outer (abaxial) side. D, exumbrella view of one of the marginal sense-organs.

The 16 rhopalar lappets are only slightly narrower than the velar lappets. There are usually
12 velar lappets in each octant. The outer edges of all lappets are bluntly rounded. The
arm-disk is five-ninths as wide as the bell-diameter in the perradius where it arises from the
subumbrella, but is only one-fourth as wide as the bell-diameter at the level of the origin of the
8 mouth-arms. The 4 interradial subgenital ostia are twice as wide as the perradial arm-disk
columns. There is a wide unitary subgenital porticus. The 8 mouth-arms are each one-
third as wide as the bell-diameter. The 3-winged lower part of each arm is somewhat more
than twice as long as the unbranched proximal shaft of the arm. The 2 lateral, outer wings
of each arm are deeply cleft (fig. 416, c). There are a very few, small, club-like appendages
(fig. B) among the mouths of the mouth-arms, but the center of the mouth-arm disk bears
a great number of clubs. These clubs are laterally flattened, the largest being only 10 mm.
long and besprinkled with nematocyst-bearing warts which are especially numerous upon

68S MEDUS2 OF THE WORLD.

their outer ends. The mouth-arms are strongly compressed laterally, being only 3 mm. in cir-
cumferential and 11 mm. in radial width at their points of origin from the mouth-arm disk.
A single duct extends into each mouth-arm, but this soon sends off a pair of side branches to
the lateral wings, and a pair of secondary branches arises from these and extends down the
main shaft of the mouth-arm (fig. 416, c).

There are 8 wide rhopalar canals of which the 4 perradial ones arise directly from the
stomach, but the 4 interradial arise from the confluence of a pair of forks. 7 to 9 narrow
vessels arise from each inter-rhopalar octant of the stomach and extend outward toward the
bell-margin, giving off numerous, anastomosing side branches which form a network connect-
ing all the canals. There is no definite ring-canal.

Near the bell-margin there 1s a sharp, angular bend in the subumbrella surface so that
the outer annulus of the subumbrella extends downward vertically and at nght angles with
the inner zone of the subumbrella. There is a wide annulus of circular muscles in the sub-
umbrella. These muscle-fibers are unbroken but somewhat thinned in the 8 rhopalar radii,
and the muscular-zone is wider in the interradi than in the perradii, but does not extend to
the edges of the arm-disk.

In formalin the rhopalar radial-canals and the proximal parts of all other canals adjacent
to the stomach are bluish-purple. The mouth frills are brownish to brownish-purple. The
bell is milky and the muscles and gonads dull brownish-yellow.

A single perfect specimen was obtained by the U. S. Fisheries Bureau steamer Albatross
on April 8, 1908, along the shore at Mantocao Island, west coast of Bohol, Philippine Islands.

=
\

Fic. 417.—Lobonema smithiit. Drawn by the author, from a preserved specimen.
View of subumbrella. Muscular system shown on right and vascular system on left.

Genus LOBONEMA, gen. nov.

The type species and only known form is Lobonema smithit from Manila Bay, Philippine
Islands.

GENERIC CHARACTERS,

Rhizostomata triptera in which the marginal lappets are greatly extended, tapering to
pointed ends. Mouth-arms with numerous filaments. Mouth-arm membranes perforated
by window-like openings. 8 rhopalia, 16 radial-canals, and a ring-canal which gives off
anastomosing vessels on both its inner and outer sides. The inner network does not connect

RHIZOSTOMA2—LOBONEMA. 689

with the stomach. The subumbrella exhibits a well-developed system of entire ring muscles.
There are numerous prominent, tapering, nematocyst-bearing papilla upon the exumbrella.
All 16 of the radial-canals extend to the bell- -margin. There is a sensory pit on the exumbrella
side above each rhopalium and the floor of this pit is furrowed.

Lobonema smithii, gen. et sp. nov.

This species is named in honor of Dr. Hugh M. Smith, Deputy United States Fish Com-
missioner, who found it in Manila Bay, Philipine Islands. The Albatross found a perfect
specimen of this medusa, and a quadrant of its disk and all of its mouth-arms were preserved.
There were also two other imperfect specimens, so that all three taken together afford data for
a partial description of the medusa.

Bell flatter than a hemisphere, 236 mm. across from each sense-club to the one 180° from it.
Gelatinous substance thick, tough, and rigid. Exumbrella regularly besprinkled with erect,
gelatinous papillz which are largest and most abundant at the center of the exumbrella but dis-
appear near the margin and are not seen over the lappets. Near the center of the exumbrella
these papillz are about 6 to 10 mm. apart and each is about 35 to 40 mm. long and 3 to 5 mm.
wide at the base; they are conical, usually more or less curved, and taper to pointed ends.
Their surfaces are thickly covered with nematocysts, which give a bristling appearance to the
disk of the medusa. 8 rhopalia which lack ocelli in specimens preserved in formalin or alcohol.
On the exumbrella side above each sense-club there is a shallow, heart-shaped, sensory pit
with dendritic ridges over its floor. The rhopalia are flanked by very small, oval, ocular lappets
only 3 mm. long and 2.5 mm. wide. There are 32 (4x8) velar lappets which are most extra-
ordinary, each being a to 100 mm. long and tapering gradually from base to tip. They are

modified so as to a ecuble superficially tentacles of semzostomous Scyphomedusz and trail
downward from bell-margin, waving flexibly to and fro as do veritable tentacles; I can find no
muscles in these lappets, however: and do not believe that they can contract and elongate.
There are deep clefts in the exumbrella surface between the lappets, but these clefis are bridged
over by a thin subumbrella membrane spanning between the lappets. The 8 inter- rhopalar
grooves are 35 mm., the 8 rhopalar 16 mm. and the 16 intermediate clefts 31 mm. long. The
8 rhopalar clefts are A shaped and the exumbrella sensory pit is at the middle of the crotch
of the A with the divided groove on either side of it (see text-figure 418, c). The grooves
between the velar lappets are simple, undivided, linear clefts.

16 radial-canals, 8 thopalar and 8 inter-rhopalar, leave the central stomach and all extend
to the bell- -margin. There is a fairly distinct ring- -canal about 30 mm. inward from the sense-
clubs, and this ring-canal gives rise on both its inner and outer sides to an anastomosing net-
work of vessels eas connect with the 16 radial-canals, but not directly with the Wore
This network of vessels extends downward throughout the length of the tapering lappets
trending mainly longitudinally but with frequent anastomoses.

The muscular system forms an annulus about 68 mm. wide in the subumbrella from the
margin of the arm-disk to the zone of the thopalia. The circular muscles are powerfully
developed, and are only thinned but not broken in the rhopalar radii. There are no radial
muscles and no muscles in the lappets.

The arm-disk is 100 mm. wide but as it was cut off, I can make no statements in ref-
erence to the size or form of the subgenital ostia or of the gonads.

The 8 mouth-arms are separate, 150 mm. long, and each is 3-winged below. The upper
shaft of each arm is 60 mm. and the 3-winged lower part go mm. long. It is remarkable that
each of the 3 lateral membranes is perforated by 3 windows or openings (see diagram A,
text-figure 418). The axial duct of the arm extends down the center and gives off side branches
in the tissue between the windows to the mouths. These side branches are joined one to
another by longitudinal canals near the frilled mouths (see text-fgure 418, B).

There are numerous appendages upon the mouth-arms arising between the mouths.
Those near the lower pointed ends of the mouth-arms are large, spindle- -shaped, more or less
triangular in cross-section and taper to pointed ends. Those arising higher up are more
slender, and above these there are mere thread-like filaments. The appendages are usually
70 to 100 mm. long, and the large ones contain an axial duct. The general color of the medusa
in formalin is milky-gray. The mouths and gonads being darker than other parts.

690 MEDUS® OF THE WORLD.

The U. S. Fisheries Bureau steamer Albatross found this medusa in Manila Bay at the
ship’s anchorage on April 25, 1908, and again at station D, 5222, between Marinduque and
Luzon, 9 miles off San Andreas Island, on the surface on April 24, 1908.

Dr. Hugh M. Smith tells me that this medusa inflicts a very severe sting upon persons who
may venture to handle it, and he believes it to have been the species which stung nine bathers

Fic. 418.—Lobonema smithii. Drawn by the author, from a preserved specimen.
A, diagrammatic illustration of one of the mouth-arms, to show the window-like openings in
side walls of arms. B, side view of a mouth-arm, showing canal-system (dotted).
C, rhopalium and one of the marginal lobes.

in Manila Bay whose cases were reported upon by Edward H. H. Old, Asst. Surgeon, U. S.
Navy. One of these cases proved fatal, and they all occurred during the summer months of
1906-07. ‘The skin where the sting occurs becomes red and vesiculated and ‘“‘weeps” as does an
eczema. Soon general pains develop throughout the body, especially in the lumbar region.

RHIZOSTOMA2—LOBONEMA, THYSANOSTOMA, 691

The mucous membranes give rise to a thin copious secretion. The patient becomes hysterical,
coughs almost incessantly and throws himself about the bed, nauseated, weeping, and with an
anxious congested face. The pulse becomes rapid and some degree of fever usually develops.
The general symptoms develop in from 10 to 15 minutes after the infliction of the sting.
The most efficient remedy was found to be a hypodermic injection of one-sixth to one-
eighth grain of morphine sulphate and an external application of an alkaline solution such
as bicarbonate of soda. A report upon these cases is presented by Dr. Old in the Philippine

Journal of Science, vol. 3, p. 329, 1908.

RHIZOSTOMATA LORIFERA Vanhiffen.

Rhizostomata lorifera, VaNWOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 45.—Maas, 1903, Scyphomedusen der Siboga
Exped., Monog. 11, p. 75.

Leptobrachide, Craus, 1883, Organisation und Entwick. Medusen, Leipzig —von LenpENFELD, 1888, Zeit. fiir wissen, Zool., Bd.
47, P- 211.—Maas, 1906, Revue Suisse de Zool., tome 14, p. 104.

CHARACTERS OF THE GROUP.

Rhizostome with very elongate, narrow, lash-like mouth-arms. The lower parts of the
arms are 3-winged in cross-section, the mouths being developed upon and near the angles.
The upper parts of the mouth-arms are very short and partially
fused to the arm-disk by a series of arches spanning from one
arm to another. The ring-muscles of the subumbrella are pow-
erfully and the radial muscles weakly developed.

The Rhizostomata lorifera are only a subordinate group of
the Rhizostomata triptera from which they have been derived by
the elongation of the mouth-arms and the reduction of the lateral
expansions of the arms. A description of the genera follows:

Thysanostoma L. Acassiz, 1862. Mouth-arms without terminal clubs. 3 rows
of frilled mouths extend down the angles throughout the entire length of
the lower arm.

Lorifera Harcxet, 1880. Similar to Thysanostoma but with a naked knob
at the lower end of each arm.

Leptobrachia Branvt, 1838= Leptobrachta+ Leonura HaeckeL. Mouths confined
to the lower and upper ends of the mouth-arms, so that the mouth-arms are
devoid of mouths in the mid-regions of their lengths.

Genus THYSANOSTOMA L. Agassiz, 1862.

Thysanostoma, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p.153.—Haecket, 1880,
Syst. der Medusen, p. 625.—VanuorreNn, 1888, Bibliotheca Zoologica, Bd. 1, Heft.

Fic. 419.—Diagrammatic representa- 3> P: 45-—KisuinoureE, 1895, Zool. Magazine, Tokyo, vol. 7, p. 133.—Scuuttzr,
tion of the form and position 1898, Denkschr. Med. Nat. Gesell., Jena, Bd. 8, p. 448.—Maas, 1903, Scypho-
of the mouth-arms in the Rhz- medusen der Siboga Exped., Monog. 11, pp. 75, 81; 1906, Revue Suisse de Zool.,
zostomata lorifera. tome 14, p. 105.

The type species is Thysanostoma thysanura from the Indo-Pacific region. It is possible
that Lesson’s inadequately described Rhizostoma brachyura may be the same medusa.

GENERIC CHARACTERS.

Rhizostomata lorifera having mouth-arms bearing 3 rows of frilled mouths from base to
lower end, without a terminal club.

Among characters of minor importance, the 4 interradial, subgenital ostia are wider than
the perradial columns between them. There are 8 rhopalar canals, and a ring-canal which
gives off a network of vessels on both its inner and outer sides. This network connects with
all the radial-canals and also at numerous points with the central stomach. ‘The well-devel-
oped circular muscles are only partially interrupted in the 8 principal radii. There is a small,
shallow, exumbrella pit above each sense-organ, without furrows in the floor of the pit.

692 MEDUS.Z OF THE WORLD.

The only difference between this genus and the closely allied Lortfera is that the frilled
mouths are developed even to the tips of the lower ends of the mouth-arm and there is no
terminal club, whereas Lorifera has a naked terminal club.

Thysanostoma thysanura Haeckel.

(?) Rhizostoma brachyura, Lesson, R. P., 1829, Voyage de la Coquille, Zoophyt., tome 2, p. 153; 1830, Centurie Zoologique,
p- 227, planche 80.

Thysanostoma thysanura, Harcxe, 1880, Syst. der Medusen, p. 625, taf. 39, fign. 1-9.—Scnuttze, L. S., 1898, Denkschr.
Med. Nat. Gesell., Jena., Bd. 8, p. 448, taf. 33, fig. 3; taf. 34, fig. 8—Maas, 1903, Scyphomedusen der Siboga Expedition,
Monog. 11, p- 75, taf. 10, fign. 93, 94; 1906, Revue Suisse de Zool., tome 14, p. 105.—KisHINovyeE, 1910, Journal Col-
lege of Sci., University of Tokyo, vol. 27, art. 9, p. 23.

Thysanostoma denscrispum, Kisninovuyr, 1895, Zoological Magazine, Tokyo, vol. 7, No. 83, p. 133, plate 18, figs. I-13.

The bell is go to 120 mm. wide. The exumbrella exhibits polygonal facets or granular
elevations which are larger at the center than at the margin. Its outline is dome-like and
flatter than a hemisphere, recalling the appearance of a shield. There are 8 marginal sense-

ae
2 8)
g

.

Fic. 420.—Thysanostoma thysanura. Drawn by the author, froma specimen taken in a
seine by the U. S. Fisheries Bureau steamer Albatross at Panabutan Bay, Philip-
pine Islands, Feb. 6, 1908.

A, oral view showing all but one of the mouth-arms cut off. One mouth-arm is cut off
close to its point of origin, but the other 6 are cut off at their widest, Y-shaped
regions. B, side view of basal part of mouth-arm, with section of same. C, sec-
tion near distal end of mouth-arm, somewhat enlarged, showing T-shape of cross-
section of arm the and 4 arm-canals. D and E, rhopalar lappets from subum-
brella and exumbrella sides respectively.

organs which are flanked by 16 small, pointed lappets and between each successive pair of
sense-organs are 6 to 12 bluntly rhomboidal lappets which are very variable in size even in
different parts of the same octant. The sense-clubs have a pigmented mass of lithocysts
and there is a very small, shallow, simple, exumbrella pit.

The 8 mouth-arms are each about 1.5 to 3 times as long as the bell-diameter; upper arm
only about one-twelfth as long as the lower arm. Lower arm delicately formed, 3-winged
and Y-shaped or T-shaped in cross-section throughout its length; in its upper part it tapers

RHIZOSTOMAS—THYSANOSTOMA, LORIFERA, 693

slightly and then continues throughout the greater part of its length of nearly uniform width,
ending in a blunt, distal extremity, covered with frilled mouths. ‘The wings are thin, longi-
fecinal lamella, one being inward (axial) and two flaring outward ( (lateral). The 2 lateral
wings give rise each to phere small, secondary wings near Ane upper, outermost end. In the
upper third of the lower arm the frilled mouths are developed upon the outer sides as well as
along the edges of the 3 wings. In the middle third they are confined to the edges of the 3
wings; and in the lower (diecal) third, they are developed upon the sides and edges as in the
upper third of the arm. Thus in the proximal and the distal thirds of the lowes arms the
mouths recall the condition seen in the mouth-arms of Mastigias and Crambione. There
are neither terminal knobs nor other appendages upon the mouth-arms, but there are
numerous short, slender, filiform, tubular appendages upon the arm-disk.

The arm-disk is quadrangular with rounded angles, with its sides about three-eighths as
long as the bell-diameter. The subgenital ostia are large and gaping, 4 times as wide as the
perradial columns of the disk, and are not narrowed by median flaps. There is a single, large,
but low, subgenital porticus.

There are only ring-muscles in the subumbrella. These are strongest near the margin
and the muscle-mass is widest in the 4 interradiu. The muscles are ‘only partially inter-
rupted in the 8 principal radii. The canal-system of the subumbrella is characterized by the
considerable width of the 8 rhopalar canals, which are somewhat wider than the others. The
ring-canal is at some distance inward from tthe bell-margin and the anastomozing network of
vessels extends on both sides of the ring-canal, fusing with the 8 radial-canals and with the
central stomach. 4 canals arise from the perradial sides of the cruciform stomach and extend
downward through the 4 pillars into the arm-disk. Here each canal divides into 4 branches,
2 horizontal ones leading into the center of the arm-disk and 2 vertical leading down into 2 of
the oral arms. Each arm-canal gives off 3 side branches which together with the axial-canal
extend down the mouth-arms, the side branches giving off branchlets to the 3 rows of mouths.

This medusa is found in the Malay Archipelago, from Amboina, Philippines, and Moluc-
cas to Japan. It is described in detail by Haeckel, L. S. Schultze, and Kishinouye. The
exumbrella is violet or mauve colored at its center, grading into russet at its margin. Sub-
umbrella flesh-colored. Gonads and mouth russet or hazel-brown. Mouth-arms and arm-
disk violet.

Kishinouye’s T hysanostoma denscrispum, from Japan, is, I believe, only the young of
T. thysanura. Lesson’s Rhizostoma brachyura, from New Guinea, has a whitish bell with
rusty-colored margin and yellowish-red mouth-frills, the general color pattern being very
similar to that of the more highly colored Japanese medusa.

The dimensions in mm. of a specimen obtained at Mindanao, Philippine Islands, by
the U. S. Bureau of Fisheries steamer Albatross are as follows: Bell, 100 wide; perradial
diameter of arm-disk, 74; diameter of arm-disk at level of origin of mouth-arms, 48; genital
ostium, 40 wide; mouth-arms, 220 long, 24 wide at widest part, 12 wide at their blunt tips;
8 to 12 velar lappets in each octant; filamentary appendages on the arm-disk, 10 to 15 long;
exumbrella finely granular.

In another large medusa from Mausalay, Mindoro, Philippine Islands, taken by the
Albatross on June 4, 1908, from a depth of 150 feet, the bell is 120 mm. wide and the mouth-
arms 190 long. In a half-grown medusa obtained on the surface at the same time and place
the bell is 59 mm. wide with finely granular exumbrella. Mouth-arms 67 long. Arm-disk
41 mm. wide at its origin from the subumbrella and 33 mm. wide at the level of the origins
of the mouth-arms. ~

Genus LORIFERA Haeckel, 1880.

Himantostoma, Acassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 152.—HarckeL, 1880, Syst. der Medusen, p. 627.—Van-
HOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 45—Maas, 1903, Scyphomedusen der Srboga Exped., Monog. 11.

Pp- 77, 81.
Lorifera, Harcket, 1880, Ibid., p. 628.

The type species is L. lorifera of the Indo-Pacific region. This genus is distinguished from
the closely allied T hysanostoma only by the naked, club-shaped extremities of its mouth-arms.

The name Himantostoma is preoccupied, having been used by Loew, 1853, for Diptera.
We must therefore use Haeckel’s alternative name Lorifera.

694 MEDUS® OF THE WORLD.

GENERIC CHARACTERS.
Rhizostomata lorifera in which the 8 mouth-arms bear rows of three continuous frilled
mouths, but terminate each in a naked knob.

Among characters of minor importance the subgenital ostia are usually wider than the
perradial disk-columns. The circular muscles of the subumbrella are practically entire. The
sense-clubs have each an ocellus and a well-developed, exumbrella pit with radiating furrows.

All the species are from the Indo-Pacific region.

Tabular Description of the Species of Lortfera.

H. lorifera: H. flagellata:

Diameter of bell in mm. 150 to 160. Exumbrella smooth. 200. Exumbrella granular.
Number of velar lappets in each octant.| 6 double. 8 rounded.

Length of mouth-arms in terms of bell- | 3 to 4r 2r

radius (r).

Length of terminal knob in terms of | One-sixtieth; a very small, swollen, oval | Two-thirds; slender and tapering.
length of mouth-arms. bulb.
Color. Bell amethyst. Margin white with dark-| ?

violet spots. Mouths dark-violet.

Where found. Red Sea to Pacific Ocean. Malay Archipelago to Hawaiian Islands.

Lorifera lorifera Haeckel.

Rhizostoma lorifera, EuRENBERG, 1835, Abhandl. Berlin. Acad., p. 260.
Leptobrachia lorifera, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 154.
Himantostoma lorifera, Haecker, 1880, Syst. der Medusen, p. 628, taf. 38, fign. 1-6.
(2?) Himantostoma sueurti, AGassiz, loc. cit. p. 152.
Lorifera arabica, HAtcKEL, loc. cit., p. 628.

Bell 150 to 160 mm. wide with very thin walls and smooth exumbrella surface. 8 rhopalia.
64 marginal lappets. In each octant 6 short, wide, bluntly-rounded, double, velar lappets
between 2 rudimentary, oval, ocular lappets. The 4-sided arm-disk is somewhat wider than
the bell-radius. The 4 subgenital ostia are 3 to 4 times as wide as the perradial columns
between them. There is a narrow, cruciform, subgenital cayity. The 8 long, tapering, whip-
like mouth-arms are twice as long as the bell-diameter. Near the arm-disk they are only 5 to
6 mm. wide and taper outwardly, being only 2 to 3 mm. at their ends where they terminate
in a naked, oval knob 5 to 6 mm. long. A ventral and dorsal row of complexly folded, frilled
mouths is found in the upper half of each mouth-arm, and thus the upper half of the arm is
triangular in cross-section, with a double row of mouth-frills at each of the 3 angles. The
ventral (inner) ridge of mouths disappears about the middle of each arm, the lower halves of
the arms being thus ribbon-like with only the 2 lateral rows of mouth-frills persisting to the
base of the terminal knob. Neither clubs nor filaments between the mouths. There is a wide
unitary zone of circular muscles in the subumbrella and apparently no radiating muscle-fibers.

Stomach cruciform, the 4 perradial oral rays of the cross being 60 mm. long and twice as
wide (30 mm.) in their outer half as they are near the center of the bell. 8 rhopalar radial-
canals arise from the stomach and extend to the marginal sense-organs. These 8 canals are
put into connection one with another by a ring-canal at some distance inward from the margin.
On its outer side the ring-canal gives off a network of vessels which fuse with the rhopalar
canals, and on its inner side there is a wider-meshed network of vessels which fuse with the
radial-canals and with the central stomach. There are traces of 8 narrow, adradial canals in
the network.

The bell is amethyst-color with a white margin and with a dark-violet spot upon each
lappet. The frills of the mouths are dark-violet and the gonads reddish-yellow.

Found at Tur, near Saini, Red Sea, in November.

This medusa may be identical with Agassiz’s “Himantostoma sueurit’”’ from the China
Sea; but in this form there are only 5 velar lappets in each octant instead of 6 as in L. lorifera.

RHIZOSTOM AB

LORIFERA. 695

Lorifera lorifera ‘“‘var.”’ pacifica.

Himantostoma loriferum, var. pacifica Scuuitzs, L. S., 1897, Ablandlung. Senckenberg. Naturf. Gesell., Bd. 24, Heft 2, p. 153,
taf. 15, fign. 1, 1a, 6; 1898, Denkschrift. Med. Nat. Gesell. Jena, Bd. 8, p. 446, taf. 34, fig. 9 (young medusa).

This variety is described by Schultze from Ternate and from Amboina, Malay Archi-
pelago.

Bell flatly rounded, 200 mm. wide and 50 to 60 mm. high. 8 marginal sense-organs.
64 marginal lappets; ach octant has 2 sharp-pointed rhopalar lappets and 6 velar lappets
separated one from another by long, deep furrows; the outer edges of these velar lappets are
rounded and each lappet usually displays a median cleft, as in ii lorifera, or is even further
divided; outer edges of velar lappets evenly rounded. The 4 subgenital ostia are 3 to
4 times as wide as the arm-columns between them. The 8 mouth-arms are 1.5 times as long
as diameter of disk. Thus in a medusa 200 mm. in diameter the arms are each 290 mm. long.
The simple upper arm is only 10 mm. long; below this each arm expands into a 3- winged
appendage, with two outer and one inner wing. This 3-winged part of mouth-arm is very
short and extends below in a very long, cieden 3-sided lash, the angles of which bear frilled
mouths. This lash terminates below in a small, vesicular, naked aot The gastric canal
which enters each lower mouth-arm sends out a longitudinal branch into each of the 2 dorsal
wings. These branches are put into communication with the central canal of the arm by
frequent cross-branches, and the side branches into the mouth-grooves often anastomose.

In the middle of the bell is a dark blue-violet area. Near the bell-margin the color is
brownish or white. The marginal lappets are violet. The smooth, outer side of the upper arm
istransparent. The thick, proximal parts of the lower arms are light-brown in their basal parts,
but throughout the greater portion of their lengths they are violet.

This ‘“‘variety”’ is probably identical with Haeckel’s L. lorifera from the Red Sea. The
slight differences mentioned by Schultze may readily be due to individual variation. For
example, in the medusa from the Malay Archipelago the lower side of the arm-disk along the
lines of the 8 axial rows of mouth-openings is beset with a felt-like mass of long, thin filaments.
These are absent, however, in Haeckel’s medusa from the Red Sea. Schultze has seen Haeckel’s
original specimen in Berlin and finds that it is a male, whereas the medusa from the Malay
Archipelago is a female. It is possible, therefore, that this difference in the condition of the
lower surface of the arm-disk may be sexual.

Lorifera flagellata.

Himantostoma flagellata, HarcKet, 1880, Syst. der Medusen, p. 629.—Maas, 1903, Scyphomedusen der Siboga Expedition,
Monog. II, p. 77, taf. 10, fign. 87-92; taf. 11, fig. 101.

The bell is flatly rounded and may be 200 mm. in width. The gelatinous substance is
thick, tough, and of a porcelain-like whiteness. The exumbrella bears fine granulations,
and near the margin are light-brown punctations, especially numerous over the marginal
lappets, where they are rendered especially conspicuous owing to the white color of the under-
lying gelatinous substance.

There are 8 marginal sense-clubs each with a large swollen end containing a concretion
and a pigment- spot of horseshoe shape. The covering scale over each sense-club 1s wider
than it is long, and there is a well-developed sensory pit upon the exumbrella side. The
bottom of this pit exhibits radiating furrows. The 8 sense-clubs are flanked by 16 short,
pointed, ocular lappets, and between each successive pair of sense-organs are typically 8
large, rounded, velar lappets. Thus there are in all 80 marginal lappets.

The arm-disk is 8-sided and the 4 interradial, subgenital ostia are nearly twice as wide
as the pillars between them. These pillars of the arm-disk are peculiar. Near their points of
origin from the subumbrella each pillar is divided so that 2 arches of gelatinous substance
extend downward to the base of the upper arm. Spanning these arches is a well-dey eloped,
gelatinous membrane which also spans the 4 interradial spaces between the arm-disk-pillars
and overlaps the 4 subgenital ostia. This peculiar arrangement is well described by L. S.
Schultz, 1898, in L. lorifera var. pacifica (see Denkschr. Med. Nat. Gesell., Jena, Bd. 8,
p- 447). The arches and the cross-spanning membrane extend so far down the length of the
upper arm that only a small portion of the latter is free.

696 MEDUS42 OF THE WORLD.

The 8 mouth-arms when contracted are hardly longer than the bell-diameter. The
lower arms are free and taper to their pointed lower ends. They are about 6 times as long as
the upper arms. At the upper end of each of these lower arms there are 2 abaxial wings so
that the arm is here 3-winged in cross-section and the frilled mouths are developed only along
the thin edges of the 3 membranous wings. The main shaft of the lower arm is, however,
triangular in cross-section and the mouths are developed upon the sides as well as upon the
angular edges of the arm. Each lower arm terminates at its pointed lower end in a long,
tapering filament which is about two-thirds as long as the lower arm itself. Numerous, short,
slender filaments arise from between the mouths, especially along the lines of the angular
edges of the lower arm. There are no club-shaped appendages.

Stomach cruciform, the arms of the cross wide and not quite as long as the center, where-
in the arms come together. Thus the stomach ts relatively wider than in other species of
Lorifera. The canal-system of the bell consists of 8 canals in the radii of the sense-organs,
which are put into communication one with another by a network of anastomosing vessels
which arise not only from the 8 radial-canals but from the edges of the stomach. There
is no clearly developed ring-canal and it is difficult to determine the number of canals which
arise from the stomach between each successive pair of rhopalar canals. There is a very wide
zone of circular muscle-fibers in the subumbrella. This muscle is only somewhat thinned but
not actually interrupted in the 8 principal radii. The 4 gonads are horseshoe-shaped and com-
plexly folded. This medusa is found at the Hawaiian Islands and in the Malay Archipelago.

In a small specimen found by the U. S. Fisheries Bureau steamer Albatross at station
D 5226, in the Philippine Islands on May 4, 1908, the bell is 50 mm. wide and mouth-arms
56 mm. long, the slender, tapering filaments at the ends of the arms being 18 mm. and the
mouth-bearing parts of the arms 38 mm. long.

Genus LEPTOBRACHIA Brandt, 1838.

Leptobrachia, Branpt, 1838, Bull. Acad. Sci. St. Pétersbourg, tome 1, p. 191.—AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4,
Pp- 154-—Maas, 1903, Scyphomedusen der Ssboga Exped., Monog. 11, p. 81.

Leptobrachia+ Leonura, HarcKet, 1880, Syst. der Medusen, pp. 630, 631.

Leonura, Harcxet, 1881, Deep-Sea Medusx Challenger Expedition, Zool., vol. 4, p. 133-

Leptobrachia+ Leonura, VANUGFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 45.

GENERIC CHARACTERS.

Rhizostomata lorifera in which the long, linear mouth-arms bear no frilled mouths near
the middle of their lengths; but near their poin‘s of origin from the arm-disk there is a ventral
row of mouths, and below the naked mid-region there are 3 lines of mouths, 1 ventral and 2
dorsal. The mouth-arms terminate below in a naked pointed end, free of mouths.

Among characters of minor importance, the slit-like subgenital ostia are wider than the
columns between them. 16 radial-canals extend to the bell-margin and a well-developed
ring-canal gives off a network of vessels on both its inner and outer sides; these networks
fuse with the radial-canals. A unitary, circular muscle is in the marginal zone of the sub-
umbrella.

Leptobrachia leptopus Brandt.

Rhizostoma leptopus, CHAMisso ev Eysennarpr, 1821, Nova Acta Phys. Med. Nature Curios., tome 10, p. 356, taf. 27, fign. 1,

A and D.

Leptobrachia leptopus, Branvt, 1838, Bulletin Acad. Sci. St. Pétersbourg, tome 1, p. 191.
Leonura terminalis, Harcker, E., 1880, Syst. der Medusen, p. 646; 1881, Deep-sea Medusx Challenger Expedition, Zool., vol. 4,

Pp- 133, plate 32, figs. 1-8.

Leonura leptura, HarcKet, 1880, loc. cit., p. 631.

The following description is derived from Haeckel’s account of his “‘Leonura termi-
nalis” which is only a modern name for Brandt’s Leptobrachia leptopus =Rhizostoma le ptopus
Chamisso and Eysenhardt.

Bell flatter than a hemisphere, 80 mm. Ma Exumbrella covered with regularly arranged,
polygonal elevations bordered by furrows. 8 rhopalia. 80 marginal lappets. All of the
lappets are sharply pointed and are largest at the middle of each octant, the smallest being
adjacent to the rhopalia, the lappets i increasing successively in size and being largest midway

RHIZOSTOMA—LEPTOBRACHIA. 697

between the rhopalia. Converging furrows extend up the sides of the exumbrella from the
clefts between the lappets. Arm-disk 4-sided, rarely as wide as the bell-radius, with subgenital
ostia 3 times as wide as the columns between them. A unitary, cruciform, subgenital cavity.
The 8 slender mouth-arms are about as long as the bell-diameter. The arms are triangular in
cross-section and end below in a triangular, pointed, naked extremity nearly one-fourth as
long as the entire arm itself. 8 rows of frilled mouths radiate outward from ‘a raised rosette
of frills at the center of the arm-disk and extend down the ventral angle of each mouth-arm
for a distance about one-fourth the length of the arm. Below this the arm is naked, triangular,
and devoid of all mouths for about one-fourth of its/length. Below this naked region are 3
double rows of frilled mouths, 1 on the ventral and 2 on the dorsal angles of the arm covering
a length equal to about one-fourth the arm; below this region is the naked terminal club. Thus
from base to lower end we find: (1) a length wherein there are only ventral mouths; (2) a mid-
region devoid of mouths; (3) a part wherein there are 3 double rows of frilled mouths, 1 ventral,
2 dorsal; (4) the pointed, naked, terminal club. There is a unitary marginal zone of ring-
muscles in the subumbrella.

The cruciform, central stomach gives rise to 16 radial-canals, 4 perradial, 4 interradial,
and 8 adradial. All extend to the bell-margin and are connected by the ring-canal at some
distance inward from the margin. A network of anastomosing vessels connects the ring-
canal with the radial-canals on the inner side. On its outer side the ring-canal gives off a
forked canal into each velar lappet and a network of anastomosing vessels which connects

with all of the canals in the lappets.

Found by the Challenger expedition near Juan Fernandez Island, off the Pacific coast of
South America.

Haeckel’s “‘Leonura leptura,’ from near New Zealand, is probably another name for
L. terminalis. It differs only in having rectangular instead of pointed velar lappets and the
mouth-arms are 3 times as long as the bell-diameter, whereas the arms of L. terminalis are
said to be only about as long as the diameter of the bell. The “quadratic” marginal lappets
of L. ‘“‘leptura” are probably due to the loss of their originally pointed ends, an accident which
frequently occurs to Scyphomeduse.

It seems probable that Rhizostoma leptopus of Chamisso and Eysenhardt, 1821, is the
same medusa. It is described from the Radack Islands, tropical Pacific. The mouth-arms

are not quite twice as long as the bell-diameter, thus resembling
Haeckel’s ZL. terminalis.

The bell is light-violet, the margin and frilled mouths
being darker. 32 reddish-violet spots on the exumbrella near

the margin. Gonads yellow.

RHIZOSTOMATA SCAPULATA Vanhoffen.

Rhizostomata scapulata, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p.
42.—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 72.

Stomolophide+ Rhizostomide, CLavs, 1883, Untersuch. tiber Organisation und Entwick.
Medusen.—von LENDENFELD, 1888, Zeit. fiir wissen. Zool., Bd. 47, p. 208.

Rhizostomz in which each of the 8 mouth-arms bears a
pair of wing-shaped outgrowths, called scapulets, or shoulder
ruffles which arise from the dorsal side of each arm near its
Fic. 421—Diagrammatic representa- point of origin from the arm-disk. Frilled mouths are devel-

honvof shape jab mouth-arm-in oped upon the upper and outer sides of these scapulets as well

Rhizostomata scapulata. <n,

as upon the lower parts of the mouth-arms. The circular
muscles of the subumbrella are powerfully and the radial-muscles weakly developed or absent.
A description of the genera follows:

Rhizostoma Cuvier, 1800. 8 free mouth-arms, the lower parts of which are Y-shaped or 3-winged in cross-section,
These 3 wings meet at a point at the lower end of the arm, and a naked, 3-cornered, club-like appendage arises from
this point. There are neither secondary clubs nor filaments upon the mouth-arms.

Rhopilema Harcxet, 1880. Similar to Rhizostoma, but with numerous clubs or filaments upon the mouth-arms.

Eupilema Hacker, 1880. Similar to Rhizostoma but there are neither clubs nor filaments upon the mouth-arms.

Stomolophus L. Acassiz, 1862= Brachiolophus + Stomolophus Harcxet, 1880. 8 laterally coalesced mouth-arms form-
ing an 8-sided throat-tube for the central mouth. Lower ends of mouth-arms are free and branch complexly.

MEDUS OF THE WORLD.

698

Genus RHIZOSTOMA Cuvier, 1799, sens. restr.

Rhizostome, Cuvier, 1800, Bull. des Sci. Soc. Philomathique de Paris, tome 2, p. 69, planche 4.

Rhizostoma, Peron et Lesueur, 1809, Annal du Muséum Hist. Nat. Paris, tome 14, p. 362.—Escuscuo.tz, 1829, (in part),
Syst. der Acal., p. 45.—pe Brarnvirre, 1834, Manuel d’Actinologie, Paris, p. 297.—Lesson, 1843, Hist. Nat. Zooph.
Acalephes, p. 411.—Gecensaur, 1856, Zeit. fiir wissen. Zool., Bd. 8, p. 210.—Acassiz, L., 1862, Cont. Nat. Hist. U. S.,
vol. 4, P- 150.

Pilema, Haecxet, 1880, Syst. der Medusen, p. 591.

Rhizostoma, Craus, 1883, Organisation und Entwick. Medusen, p. 60, Leipzig —Vannorren, 1888, Bibliotheca Zoologica, Bd. 1,
Heft. 3, pp. 31, 43-—Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 80.

The type species is Rhizostoma pulmo of the Mediterranean.

GENERIC CHARACTERS.

Rhizostomata scapulata with 16 scapulets upon the outer sides of the 8 mouth-arms.
The 8 mouth-arms are free, not fused together. Lower arms 3-winged, each terminating in
a single, club-like, gelatinous appendage. There are no other clubs, nor filaments among the
mouths. The 4 subgenital ostia are narrow cross-slits, each constricted by a wart-like papilla
upon the subumbrella. 16 radial canals, 8 rhopalar and 8 adradial, all of which extend to
the bell-margin. A network of anastomosing vessels arises from the outer halves of these 16
radial-canals, and this network gives rise centripetally to 16 blindly-ending areas in the
sectors between the radial-canals. There is no marginal ring-canal, but the most direct and
widest connection between the radial-canals is through the middle of the zone of anastomos-
ing vessels. 16 triangular areas of circular muscle-fibers alternate with the 16 radial-canals.
The marginal sense-clubs lack ocelli. There is an exumbrella pit above the sense-club and -
the bo:tom of this depression is furrowed with radiating ridges.

This genus is distinguished from Eupilema Haeckel by having 8 terminal clubs upon its
mouth-arms whereas these are absent in Eupilema. It is distinguished from Rhopilema by

Synopsis of the varteties of Rhizostoma pulmo.

R. pulmo. R. lutea. R. octopus. R. corona. R. capensis.
| Diameter of bell | 150 to 600 200 to 300 As in R. pulmo. 200 to 400 200 to 300 (?)
| in mm,
Character of ex- | Finely granular. Rough with oval As in R, pulmo. Smooth (?) Smooth or finely
umbrella surface. warts. granular.
| Number of mar- | 80 80 96 to 112 140 to 180 ?
ginal lappets.
| Shape of velar lap-| Semicircular. Short, oval. Short, pointed. Small, truncated. Semicircular.
pets.

Not as long as lower
arm.

Shorter than lower
arm.

Nearly twice as long
as lower arm.

Longer than lower | As in R. pulmo.

arm.

Length of upper
arm.

Somewhat shorter
than upper arm.
Width nearly uni-
form throughout.
Terminal knob
with toothed angle.

Half as long as upper
arm. Pyramidal,
3-cornered with
toothed angles and
bluntly pointed
end. -

Longer than upper
arm with a slender
basal stalk and
swollen club-
shaped outer end.

Shorter than, or
equal to, upper
arm. Widest near
base. Constricted
at base. No basal

| stalk.

Longer than entire
arm. With long
slender basal stalk
and swollen club-
like outer end.

Length and shape
of terminal club.

Color. Bell milky-yellow, | Warts of exum- As in R. pulmo. ? Bell and clubs
occasionally red- brella reddish- bluish. Mouth-
dish. Marginal brown. Frilled frills brownish-red.
lappets blue or vio-)_ mouths yellowish.
let. Mouth-frills | Terminal club
orange-yellow to deep purple-brown.
brownish-red or
violet.

Where found. Mediterranean, Straits of Gibralter. | Atlantic coasts of | Red Sea at Suez and| Cape of Good Hope,

Europe, France to | Tur. South Africa.
Scotland.
Remarks. Described in detail | Closely related to | Imperfectly known. | So imperfectly
by Grenacher and | R. pulmo, but Apparently closely | known that it can

Noll.

distinguished by its
swollen, club-like,
terminal append-

ages.

related to R. pulmo,

not be determined.

PLATE 73.

Fig. 1. Rhizostoma pulmo, 0.75 natural size. Zoological Station, Naples,
Italy, November 29, 1907.

Fig. 2. Cotylorhiza tuberculata, 0.75 natural size. Zoological Station, Naples,
Italy. November 30, 1907.

Drawn from life, by the author.

MAYER
PLATE 73

| Gg BW OUU OS

queen’ Re ae yy DLIIITN )
eee ee ee NS Se bay
. pe Bee EG,

RHIZOSTOMA2—RHIZOSTOMA, 699

having but a single filament (a terminal one) on each mouth-arm, while in Rhopilema there
are, in addition to the terminal knob, many lateral appendages between the frilled mouths.

The varieties of Rhizostoma are known only from the Mediterranean, Red Sea, and At-
lantic coasts of Europe and Africa, and are closely related to R. pulmo of the Mediterranean;
distinguished one from another only by the relative lengths, and the various shapes of the
terminal clubs.

Rhizostoma pulmo Agassiz.
Plate 73, fig. 1.

Potta marina, ALDRovANDI, 1642, Zoophyt. Lib., tome 4, pp. 73, 76. Also; Pulmo marinus, pp. 75, 77+

Medusa pulmo, Macrt, 4778, Atti Real. Acad. Sci. Napoli, vol. 2, p. 45, tav. 1.—Linné, (Gmelin), 1788, Systema Nature, Ed. 13,
Pars. 6, p. 3155- '

Rhizostoma aldrovandi, Péron er Lesueur, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, p. 362, No. 102.—Detre Cuiaje,
1822, Mém. Anim. senza Vert., Regno Napoli, tav. 74, fig. 10; tav. 75, figs. 1-10; Ibid., 1841, vol. 7, tav. 142, 143.

Cephea aldrovandi, pe LaMarck, 1816, Syst. Anim. sans. Vert., tome 2, Pp- 517-

Rhizostoma cuviert, Cuamisso et Eysenuarpt, 1821, Nova Acta Acad. Nat. Cur., Leop. C., tome 10, p. 377, plate 34.—pe
Brainvitie, 1834, Manuel d’Actinologie, p. 297, planche 44, fig. 1.

Rhizostoma pulmo, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 150.—C aus, 1883, Untersuch iiber Organisation und
Entwick. der Medusen, p. 43, taf. 12-15, fign. 86, 105 (development of the ephyra).—Grarrre, 1884, Arbvit. Zool. Inst.
Wien, Bd. 5, p. 343 (seasonal distribution)—Vanuorren, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p- 31.—v_E VeEscov1,
1895, Zool. Results, vol. 1, p. 37-—von Urxkixt, 1go1, Mitth. Zool. Sta. Neapel, Bd. 14, p. 620 (rhythmical pulsation).—
Bere, 1903, Allgemeine Anat. and Physiol. Nervensyst., pp. 87, 90, 108, 410, 426, 432, etc., fign. 33-35, 83, 87, etc. (rhyth-
mical pulsation); 1908, Archiv fiir Ges. Physiologie, Bd. 124, p. 541; Ibid., 1909, Bd. 127, p.219.—Hararrt, 1904, Journal
of Exper. Zool., Baltimore, vol. 1, p. 73, figs. 1-6 (regeneration).

Rhizostoma cuviert, Branpt, A., 1870, Mém. Acad. Sci. St. Pétersbourg, tome 16, No. 6, 29 pp., 1 taf. (detailed description).—
Craus, 1877, Denkschrift. Wien Acad., Bd. 38, p. 47, plate 10, 11; 1881, Zool. Anzeiger, Bd. 4, p. 79 (young ephyra),
1883, Arbeit. Zool. Inst. Wien, Bd. 5, p. 9, taf. 2, fig. 12 (young ephyra); 1884, Arbeit. Zool. Inst.Wien, Bd. 5, p. 169;
taf. 2, fig. 12 (ephyra).—Brancnarp, 1883, Zool. Anzeiger, Bd. 6, p. 67 (the blue coloring matter is not cyanein).—Hessr,
1895, Zeit. fiir wissen. Zool., Bd. 60, p. 411, taf. 20-22 (nervous system and sense-organs).

Pilema pulmo, Harcxet, 1880, Syst. der Medusen, p. 591 (record of literature)—Hamann, 1881, Jena. Zeit. fiir Naturwissen,
Bd. 15, p. 250, taf. 10, fiz. 13 (structure of mouth-arms).

Bell pyriform, somewhat higher than a hemisphere, usually not more than 150 mm. in
diameter, though specimens 600 mm. wide may occasionally be found. Surface of exumbrella
finely granular, being covered with small nettling-warts. 8 marginal sense-organs, each
containing an orange-colored mass of concretions of entodermal origin; no ocellus. Above
each sense-organ on the exumbrella side is a wide triangular pit, the bottom of which exhibits
diverging furrows. Each sense-organ is flanked by a pair of narrow, elongate, lanceolate,
sharp-pointed rhopalar lappets. 8 evenly rounded, velar lappets, all similar each to each in
size and shape in each octant. Thus there are in all 80 marginal lappets (16 rhopalar+64
velar).

The total length of the mouth-arms, including their terminal clubs, is about equal to the
bell-diameter. In its upper part the arm-disk is 4-sided and narrow, but below it widens out,
becomes 8 to 16-sided and gives rise to the 16 (8 pairs) short, simitar-shaped scapulets
having fringed mouths upon their upper, convex sides. The 8 mouth-arms are each 3-winged,
or Y-shaped in cross-section, two of the wings being directed outward, one being centripetal.
These wing-like expansions bear numerous mouths, the lips of which are fringed by a row of
short, flexible, knobbed tentacles.

The terminal knobs are triangular in cross-section, somewhat contracted in the middle
of their lengths and with a simple, central canal. They are usually a little shorter than the
upper arms, but may equal or slightly exceed them in length. The mouth-bearing, lower
arms are somewhat shorter than the free upper arms between the zone of the 16 scapulets
and the winged portion of the arms. Ordinarily the scapulets are hidden away under the con-
cavity of the subumbrella. The 4 subgenital pits are narrow, slit-like and with their open-
ings constricted in the middle by a knob-shaped protuberance upon the floor of the subum-
brella. The 4 invaginated, genital sacs are small and separated completely one from another.

The cruciform central stomach giyes rise to 16 radiating canals, 8 of which extend to the
sense-organs and 8 are intermediate in position; all reach the bell-margin. A network of
anastomosing vessels places the outer halves of the radial-canals in connection one with
another, and extending inward between the 16 radial-canals are 16 blindly ending areas
composed of a network of vessels. There is no distinct ring-canal at the bell-margin, but
the connections between the radial-canals are wider along the inner edge of the zone of anas-
tomosing vessels than elsewhere. 16 deltoid areas of circular muscles alternate with the
radial-canals and are more or less completely separated in the radii of the canals themselves.

700 MEDUS OF THE WORLD.

The canal-system of the mouth-arms is as follows: 4 vessels arise from the perradial angles
of the lower side of the central stomach and extend downward into the arm-disk in the 4
perradii. These 4 primary canals bifurcate and the 8 branches extend down the 8 mouth-
arms. The 16 lateral vessels which go into the scapulets arise from these 8 mouth-arm canals.

The central mouth is always present in very young medusz but it usually disappears
inthe adult. When this mouth is present, or more or less vestigial, we often find a correspond-
ing complexity in the mouth-arm canals, for in this case the 4 perradial canals from the corners
of the lower part of the central stomach extend downward beyond the points of origin of the

Fic. 422.—Rhizostoma pulmo. From life, by the author, at Naples Zoological Station, December, 1907.

1, oral view of bell with mouth-arm removed. One half of the surface shows sector of circular muscles, and
the other half shows the muscles removed to reveal the canal-system. 1, side view of mouth-arms
with sections of same. 11, section of arm-disk immediately below scapulets. 1v, oral view of arm-
disk showing persistent central mouth opening. v, sense-organ from exumbrella side.

8 mouth-arm canals; upon reaching the cruciform, central mouth, each of these 4 canals
forks and the 8 branches extend outward along the 8 lines of the frilled mouths, becoming
confluent with the 8 main, mouth-arm canals in the 3-winged parts of the mouth-arms. A
detailed description of the canal-system is given by A. Brandt, 1870.

The gelatinous substance of the disk is creamy-yellow to milky, or rusty-yellowish and
translucent. Marginal lappets dark cobalt or violet to blue. The sensory-clubs are tipped
with orange, which colors the concretions. The frilled mouths are dull orange, yellow, or
brownish-yellow, and the outer parts of the terminal knobs are tinged with the same color.
Gonads yellowish, all other parts translucent.

RHIZOSTOMA8—RHIZOSTOMA, 701

This medusa is common in the Mediterranean. It is found throughout the year, but is
most abundant from June until August, becoming ripe in August and September. Very small
medusz are often found in June. Mature individuals are occasionally seen in midwinter.

Claus has studied the development of the pelagic ephyra. When 3.5 mm. in diameter
the ephyra has a central, cruciform mouth, the 4 lips of which are lined by a row of knobbed
tentacles as in Aurellia. There are 8 pairs (16) of velar lappets and 8 pairs (16) rhopalar lap-
pets. 16 radial-canals and a simple, circular canal. In this stage the 8 adradial canals end
in the ring-canal, but the 8 others go to the bell-margin. The velar lappets develop in pairs,
as in the Wecomedues: and in Stomolophus, not singly as in Aurellia or Cotylorhiza. The 8
mouth-arms arise from paired, terminal folds of the 4 primary rays of the cruciform, cen-
tral mouth. A detailed description of the young ephyra is given by Claus, 1883.

The rhythmical pulsation of this medusa has been seidied by von Uexkiull, who found
that if the marginal sense-organs be mechanically confined the pulsation is hindered. His
conclusion, however, that the stimulus which produces pulsation is mechanical in nature and
may be likened to that produced by the clapper of a bell in striking against the margin seems
to me improbable. Pressure upon the nerve-center might readily interfere with the activity
of the sense-club, and any confinement which cuts off the supply of soluble calcium from the
sea-water would soon cause pulsation to cease.

A very sug ggestive and important series of studies of the nature of the pulsation stimulus
in this medusa and in Cotylorhiza tuberculata was carried out by Bethe, 1903-1909. He finds,
in 1903, that under normal conditions hundreds of pulsations follow quite regularly one after
another, with only an occasional pause of brief duration. The medusa pulsates almost inces-
santly. According to Bethe there are many analogies between the pulsation of this medusa
and that of the vertebrate heart. For example, the “all or none” principle applies to
medusz, as does also the phenomena of the extra systole and corresponding compensation-
period of rest. The medusz also show a refractory stage during systole in which they are
insensible to stimuli, as was demonstrated by Marey, 1876 (Travaux du lab., p. 73), for the
vertebrate heart.

There is an increase in the time that elapses between stimulation and response, and also
in the duration of the pulsation itself as the temperature is lowered from 25° to 13° C. The
pulsation-stimulus is neryous in nature, being transmitted by the diffuse, nervous network
of the subumbrella. Indeed, there are areas of the subumbrella which are wholly without
muscles; nevertheless the pulsation-stimulus passes freely over these to the muscular areas
beyond. Bethe gives a good series of figures showing the histological character of the nerve
plexus which forms a network between the epithelium and the deep-lying, muscular layer of
the subumbrella. The sense-organs are physiologically speaking only highly differentiated
parts of the nerve-plexus of the subumbrella.

Under normal conditions the pulsation-stimulus originates in the marginal sense-organs,
yet in medusz (such as Cotylorhiza) which have an inner zone of radial and an outer zone of
circular muscles in the subumbrella the radial-muscles contract before the circular, although
they are farther away from the sense-organs. This is due, as Bethe shows, to the fact that the
latent period (1. e., the time that elapses between stimulus and response) is longer for the cir-
cular than for the radial muscles.

The pulsation is a reflex due to a constantly present stimulus, and the refractory stage
produces periodicity (rhythm) in the responses. The nerves can not send forth a new con-
traction-stimulus until a definite period of rest has elapsed.

Bethe, 1908, 09 (Pfliiger’s Archiv. fiir ges. Physiologie, Bdn. 124 and 127), has continued
his studies of the rhythmical pulsation of Riizoctorm pulmo at Naples. He finds that artificial
sea-water will not sustain life and pulsation as well as does natural sea-water unless a small
amount of CaCo, be added to the solution. This improvement of the artificial sea-water is not
due he believes to the addition of Ca or to the neutralization of an acid, but is caused by the
presence of the undissociated molecules of CaCo,. It will be recalled that Rogers, 1905
(Journal Experimental Zool., vol. 2, p. 249), found that the addition of small amounts of CaCo,
to solutions containing the pulsating heart of the crab Brachynotus had a beneficial effect. He
attributed this, however, to the neutralization of free acid in the solutions.

702 MEDUS OF THE WORLD.

NaCl in the absence of the other salts of sea-water at first stimulates and later retards
pulsation, and this effect is commonly reversible. According to Bethe, the absence of calcium
in the presence of the other salts of sea-water causes all movement to cease but normal
pulsation is restored by restoring the calcium to its normal amount. In a slight excess
in from 2 to 5 c.c. of $ molecular CaCl, in 100 sea-water calcium operates for a long time
to accelerate and strengthen pulsation. In larger excess such as 10 CaCl, in 100 sea-water
it retards pulsation and produces an abnormal duration of systole. Magnesium chloride
and sulphate produce a marked primary retardation, without any final acceleration. Potas-
sium tends gg | to stimulate the rhythmical movement in Rhizostoma. Thus, accord-
ing to Bethe, Na, K, and Ca of the sea-water are primarily stimulants for pulsation, but are
antagonized by Mg which alone is an inhibitor of pulsation, and thus a balanced solution is
formed. Na and Mg exert their effect primarily through the marginal sense-organs, and Mg
also acts upon the general system of muscles. Na, on the other hand, affects the nervous sys-
tem. Potassium exerts its influence through the marginal sense-organs but has little or no
direct influence upon the muscles and nervous network. Calcium in slight excess exerts all
or nearly all of its effect through the marginal sense-organs.

Aluminium is an inhibitor of pulsation and is relatively more powerful in this respect than
is magnesium. MgSO, is a less powerful inhibitor than MgCl, and the order of efficiency as
stimulants of the sodium salts is beginning with the most powerful Naz SO,, NaCl, Nal,
NaBr, NaNO,, and of the potassium salts K,SO,, KCl, KNO,. The rhythmical move-
ment of the whole medusa is controlled by the marginal sense-organs.

The action of the cations of the sea-water upon Rhizostoma appear, from Bethe’s work, to
be similar to their effect upon Casstopea, and I believe that the two medusz are controlled in
the same manner. (See Casstopea xamachana.)

I think, therefore, that Bethe is mistaken in concluding that potassium and calcium
stimulate pulsation. It is true in both RArzostoma and Cassiopea that pulsation endures longer
in NaCl+ KCl or in NaCl+ CaCl, than in pure NaCl, but its rate is slower. Indeed the rate
of pulsation in NaCl is so abnormally rapid that the medusa soon comes to rest exhausted, and
this exhaustion is partially prevented by the subduing effects of K or Ca and thus the pulsation
endures longer but at a slower rate. The apparent stimulation upon adding calcium in excess
is not due to the direct effect of the Ca, but to the fact that Ca counteracts the stupefying
effects of Mg, thus permitting the sodium to act more effectively as a stimulant. Thus the
sea-water is a balanced fluid, the stimulating effect of the Na cation being offset by the stupefy-
ing effects of Mg, K, and Ca.

Hargitt, 1904, has studied regeneration in R. pulmo, and he finds that it possesses a
moderate capacity, in confinement, for replacing lost rhopalia. Often 2 rhopalia regenerate .
in the place of the one which has been cut away. The medusa has but little power to regenerate
mouth-arms or gastric lobes.

Hesse, 1895, has made an elaborate study of the histology of the marginal sense-organs
and of the neryous network of the subumbrella. Definite strands of nerve-fibers extend
radially inward from the marginal sense-organs under the subumbrella epithelium. These
radial-fibers bend circumferentially at the zone of the ring-canal, forming a wide ring-like
band of nerve-fibers on the inner side of (centripetal to) the ring-canal. These nerve-fibers
arise from bipolar ganglia which are found in considerable numbers in the ectoderm along
the lines of the radial-canals which extend to the sense-organs. The bipolar ganglia are
derived from ectodermal cells of the subumbrella.

The ectodermal cells of the marginal sense-organs bear cilia, and some of them are
sensory while others are supporting cells. The sensory pit on the exumbrella side of the
sense-club is also lined with a ciliated, ectodermal epithelium containing sensory and support-
ing cells, and also ganglion cells which send numerous nerve-fibers downward through the
gelatinous substance into the entodermal core of the sense-club, where they form a felting
of fibers extending into the region of the entodermal concretions.

The inner sensory pit on the subumbrella side of the sense-club is also lined with an
epithelium similar to that of the exumbrella pit, and it also sends nerve-fibers into the sense-
club. The sensory and ganglion cells of these 2 pits and of the sense-club constitute the
central nervous system of the medusa.

RHIZOSTOMA2—RHIZOSTOMA. 703

Rhizostoma pulmo var. lutea Eschscholtz.

Orithyia lutea, Quoy ev Gaimarp, 1827, Annal. des Sci. Nat., Zool., tome 10, p. 175, planche 4B, fig. I.

Rhizostoma lutea, Escuscuourz, 1829, Syst. der Acalephen, p. si.

Pilema stylonectes, HArcKEL, 1880, Syst. der Medusen, p. 595.

Rhizostoma luteum, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 43—Grenacner und No tt, 1876, Abhandl.
Senckenberg Naturforsch. Gesell., Bd. 10, p. 42, taf. 8.

See the tabular synopsis of species of Rhizostoma.

This medusa, from the Straits of Gibraltar, is intermediate in most of its characters
between R. pulmo of the Mediterranean and R. octopus of the Atlantic coasts of Europe.
Indeed the 3 forms are so closely related that we may consider them to be local varieties
one of another. R. lutea is distinguished chiefly by its very long, terminal appendages on the
mouth-arms.

Rhizostoma pulmo var. octopus Oken.

Urtica marina, etc., octopedalis, Boriase, 1758, Nat. Hist. of Cornwall, p. 258, plate 25, figs. 15-17.

Medusa octopus, Linné (Gmelin), 1788, Systema Nature, Ed. 13, Pars. 6, p- 3157-

Medusa bleu, Cuvier, 1799, Journal de Physique, Chimie, d’Hist. Nat. et des Arts, Paris, tome 49, p. 437, 1 planche.

Cassiopea Perle Riixasore cuvieri, PERoN ET Lesueur, 1809, Annal. du Mus. Hist. Nat. Paris, tome 14, pp. 357, 362

Casstopea lunulata, Escuscuortz, 1829, Syst. der Acalephen, p. 44.

Rhyzostoma octopus, Oxen, 1835, Allgemeine Naturgesch., Bd. 5, p. 218.

Cassiopea anglicat+ C. rhizostomoidea+ Rhizostoma sepioides, Tiresius, 1829, Nova Acta Acad. Cur., Leop. C., tome 15, pp. 273,
283, tab. 71.

Rhizostoma coeruleum, Cuvier, 1817, Régne Animal., tome 4, p. a

Holigocladodes lunulatus, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, p. 155.

Pilema octopus, Harcket, 1880, Syst. der Medusen, p. 593 (list a authors).

Rhizostoma octopus, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft.3, p. 43; 1906, Nordisches Plankton Acraspede Medus.,
Nr. 11, p. 63, fign. 33, 34; 1908, deutsche Siidpolar Exped., 1901-03, Bd. 10, Zool. 2, pp. 28, 47.—Dau, 1893, Kommiss. zur
wissenschaft. Untersuch. deutsch, Meere Kiel, Ber.6, p. 172.—Browne, 1905, Proc. Roy. Soc. Edinburgh, vol. 25, p. 776.

This form is closely allied to R. pulmo of the Mediterranean, but it ranges into the
cold waters of the Atlantic coasts of Europe, being found off the shores of France, England,
Scotland, Belgium, Holland, and Germany. Mature individuals occur in great swarms in
September and October and are often cast ashore in vast numbers. The medusa differs
from R. pulmo in having g6 to 112 marginal lappets instead of 80. The lappets are more
pointed than in R. pulmo. The simple, upper part of the arm is shorter than the lower, 3-winged
part, whereas in the typical R. pulmo the reyerse is usually the case. The terminal clubs are
widest near their outer ends, whereas in R. pulmo they are widest near their bases. In other
respects it appears to be identical with R. pulmo.

Vanhotten, 1906, describes the young ephyra, 3 mm: wide, in the stage wherein the 4 rays
of the cruciform, central mouth are beginning to fork at their outer ails There are 4 small
clusters of gastric cirri. 16 radial-canals. The 8 rhopalar radial-canals extend to the marginal
sense-clubs and the 8 adradial canals end in the ring-canal, which is at a considerable distance
inward from the bell-margin. This ephyra is thus similar in all respects to that of R. pulmo.

It is somewhat remarkable that R. octopus is not found among the Azores, Canaries, or
other island groups of the Atlantic. Borlase states that in 1758 it was sometimes eaten by
man in Cornwall.

Rhizostoma pulmo var. corona Eschscholtz.

Medusa corona, Forskat, 1775, Descript. Anim. Itin. Orient., p. 107.
Rhizostoma corona, Escuscnoutz, 1829, Syst. der Acal., p. 52.-—VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p- 43-

Rhizostoma cuvieri, EHRENBERG, 1835, Abhandl. Berlin Acad., p. 184.
Pilema corona, HaEcKEL, 1880, Syst. der Medusen, p. 594.

See synoptic table of the species of Rhizostoma.
This imperfectly known Red Sea medusa appears to be closely related to R. pulmo of the
Mediterranean, but is said to be distinguished by having 140 to 180 marginal lappets,

instead of 80 as in R. pulmo.
Rhizostoma pulmo var. capensis Lesson.

Cephea capensis, Quoy er Gaimarp, 1824, Voyage de I’ Urante, p. 568, planche 84, fig. 9.
Rhizostoma capensis, Lesson, 1843, Hist. Zooph. Acaléphes, p. 417-

Pilema capense, Harcket, E., 1880, Syst. der Medusen, p. 645.

Rhizostoma capense, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 43-

704 MEDUS.® OF THE WORLD.

Quoy and Gaimard give an artistic, but evidently inaccurate, figure of this medusa, and
they fail to describe it. Their record is only interesting in that it indicates that a Rhizostoma
is found in the South Atlantic.

This form is from Table Bay, coast of South Africa, in March. It may be identical with
R. octopus or R. pulmo. Quoy and Gaimard’s figure shows an evenly rounded, oval bell,
higher than a hemisphere, with smooth or finely granular surface and with numerous
rounded lappets, as in R. pulmo. The lower arms are apparently longer than the upper,
as in R. octopus, and the terminal clubs are bluntly pointed, taper from base to tip, and are
about as long as the upper arms.

The colors are as in R. pulmo, but the bell and terminal clubs appear to be of a deeper
blue. Size (?) Number of lappets (? ) The medusa has not been seen since the days of Quoy
and Gaimard.

Genus RHOPILEMA Haeckel, 1880.

Rhopilema, Harcxen, 1880, Syst. der Medusen, p. 596.—Maas, O., 1903, Scyphomedusen der Siboga Expedition, Monog. 11,
p. 72-—C aus, 1883, Organisation und Entwick. Medusen, p. 60.—von LENDENFELD, 1884, Proc. Linnean Soc. New South
Wales, vol. 9, p. 291-—Kisninouye, 1890, 91, Zool. Magazine, Tokyo, vol. 2, p. 47; vol. 3, p. 53; 1899, Zoolog. Jahr-
biich., Abth. Syst., Bd. 12, p. 205.

Rhizostoma (in part), VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft-3, pp- 31, 43-

Nectopilema, Fewxes, 1887, American Journ. Sci., ser. 3, vol. 33, p. 120.

GENERIC CHARACTERS.

Rhizostomata scapulata with 8 separated, 3-winged, adradial mouth-arms which bear
numerous filaments or club-shaped appendages.

The central stomach gives rise to 16 radial-canals placed in intercommunication by a
network of anastomosing vessels. A ring-canal may or may not be present.

This genus is closely related to Rhizostoma (Pilema of Haeckel), but in Rhizostoma a
single club-shaped appendage arises from the lower end of each of the 8 mouth-arms and
there are no other clubs or filaments; while in Rhop:lema there are many appendages upon
each mouth-arm. The type species is R. rhopalophora of the Indian Ocean, and the same
medusa appears to haye been described by Kishinouye from Japan. “Nectopilema” of
Fewkes is identical with Rhopilema.

The edible medusz of China and Japan belong to the genus Rhopilema.

The terminal clubs upon the mouth-arms of Rhizostoma are merely the downward, mouth-
free extension of the axial shaft of each arm. These terminal clubs are triangular in cross-
section, as are the mouth-arms themselves, and they contain a continuation of the axial-canal
of the arm. They are not homologous with the vesicular, club-shaped mouth-filaments of
Rhopilema.

Rhopilema esculenta Kishinouye.

Rhopilema sp., Kisuinovuye, 1890, Zool. Magazine, Tokyo, vol. 2, p. 47, plate 2.

Rhopilema esculenta, Kisutnovre, 1891, Zool. Magazine, Tokyo, vol. 3, p. 53; 1899, Zoolog. Jahrbucher, Bd. 12, Abth. Syst.,
p- 205, taf. 13, fign. 1-5.

(2) Rhopilema rhopalophora, Haecker, 1880, Syst. der Medusen, p. 596.

The umbrella is more than 450 mm. wide, about 330 mm. high when contracted, but
nearly hemispherical when expanded. It is about 50 mm. thick at the center, but gradually
becomes thin toward the margin. Exumbrella smooth, but the marginal lobes are furrowed
with numerous, minute, longitudinal (radial) grooves. 8 marginal sense-organs, each with
an exumbrella sensory pit which displays radiating furrows in its floor. In each octant of
the margin there are 14 to 20 oval, velar lappets between 2 very small, lanceolate, ocular
lappets. The ocular lappets are only about one-fourth as long and as wide as the velar lappets.

In the subumbrella the circular muscles are well developed and unitary, and the coronal
furrow is not distinct. The arm-disk is very thick and prismatic, about one-third as wide as
the bell-diameter and somewhat longer than wide. The 4 oral pillars are quadrate and the
4 subgenital ostia are somewhat heart-shaped and as wide as the pillars. There is a rough,
prickly protuberance upon the floor of the subumbrella opposite the opening of each sub-
genital ostium. Altogether, therefore, there are 4 of these protuberances alternating with the 4

RHIZOSTOM2—RHOPILEMA. 705
arm-disk pillars, and thus interradial in position. 4 separate genital cavities. 8 pairs (16)
simitar-shaped scapulets arise from the adradial sides of the arm-disk. Their upper sides
are convex and bear frilled mouths and numerous, hollow filaments which are about two-thirds
as long as the scapulets themselves. The lower sides of the scapulets are concave and devoid
of mouth-openings or filaments.

The 8 adradial mouth-arms are stout, triangular, and pyramidal, and exclusive of their
appendages they are about two-thirds as long as the diameter of the umbrella. The upper
parts of these arms are coalesced with each other along half theirlengths. These upper parts

Way tan
Hendry der
: ; Wh hy uh
iol I Wily

at » I I

Hy lt!
\Wi ll \\

il

ct AW QY, MN) jl
AN OQ Gy, Ml”
Nw Ws = uy Y Vy,

F
Fic. 423.—Rhopilema esculenta. After Kishinouye, in Zoolog. Jahrb., Abth. Syst., Bd. 12.

A and B, side and front views of mouth-arms. C, sagittal section of medusa. D, oral view of arm-disk.
E, rhopalium. F, subumbrella showing canal-system (above) and muscular sectors (below).

of the arms are free of mouths and are about as long as the scapulets, while the lower parts are
twice as long as the upper and bear numerous, frilled mouths and more than 100 appendages.
There are 2 kinds of appendages—filamentous and fusiform. The fusiform appendages are
longer than the filaments and may be three-fourths as long as the diameter of the umbrella,
the 5 longest being found at the center and 1 at each perradial angle of the arm-disk. The
filaments are much more numerous than the fusiform appendages. There are no definite
terminal clubs at the lower ends of the mouth-arms.

The central stomach gives rise to 16 radial-canals, 4 perradial, 4 interradial, and 8
adradial. These canals extend to the bell-margin and are connected one with another by
means of an indistinct ring-canal which is about midway between the margin and the periphery

706 MEDUS OF THE WORLD.

of the stomach. On its inner side the ring-canal gives rise to an anastomosing network of
vessels which fuse with the perradial and interradial canals, but not with the adradial. On
its outer side the ring-canal gives off another network which fuses with all 16 radial-canals.

4 main canals arise from the lower part of the stomach in the 4 principal radii. These
main canals fork and each fork extends down one of the 8 adradial mouth-arms, where they
branch many times and go to the numerous frilled mouths.

The color of the medusa is usually blue, but occasionally dark-red. ‘The mouth-frills
are brown and the mouth-arm appendages are milky-white, or nearly transparent. The
gonads are yellow, the male being lighter in color than the female.

This medusa is abundant in the Inland Sea of Japan, and is also found off the coast of
China.

It is the custom in Japan to preserve it with a mixture of alum and salt or between the
steamed leaves of a kind of oak. It is then soaked in water, flavored with condiments, and
when so prepared constitutes an agreeable food.

Rhopilema rhopalophora Haeckel, from the Indian Ocean east of Madagascar, is closely
allied to, if not identical with, this Japanese medusa, but it is said to have a large, terminal
club at the end of each arm. This club is fusiform, triangular in cross-section, and as long
as the whole lower-arm itself. There are 144 lappets, the velar ones being rectangular, and
the bell is 100 mm. wide and hemispherical. In other respects it appears to be similar to
Kishinouye’s medusa, although Haeckel’s description is too brief to be satisfactory.

Rhopilema hispidum Maas.

(?) Pilema clavigera, HarcKeL, 1880, Syst. der Medusen, p. 595.

Rhizostoma hispidum, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 32, 43, taf. 5, fign. 1, 2-
Rhopilema verrucosa, Kisninovye, 1899, Zoolog. Jahrbiicher, Bd. 12, p. 208, 1 fig.

Rhopilema hispidum, Maas, 1903, Scyphomedusen der Siboga Expedition, Monog. 11, p. 73, taf. 9, fign. 78-81.

Bell hemispherical or higher than a hemisphere, and may become about 250 to 340 mm.
in diameter. Walls very thin; the exumbrella is thickly besprinkled with small, sharp-pointed,
conical projections. ‘The 8 marginal sense-clubs have no ocelli, but above each is a large,
sensory pit with radiating furrows. 80 marginal lappets. ‘The 8 sense-organs are each flanked
by a pair of very small, narrow, lanceolate lappets and there are typically 8 velar lappets in
each octant of the bell-margin; these velar lappets are oblong, rounded, and 3 times as long
and 5 times as wide as the ocular lappets.

The arm-disk is of the usual 8-sided form. The 4 interradial subgenital ostia are,
according to Maas, not quite so wide as the perradial columns of the arm-disk between them;
but according to Kishinouye the subgenital ostia in his ““Rhopilema verrucosa,” which appears
to be identical with R. hispidum, are 3 times as wide as the perradial columns. The 4 genital
cavities are only partially and irregularly fused and do not form a unitary geni‘al space, as in
Mastigias and Crambessa, nor are they completely separated into 4 cavities, as in Cassiopea
(see Maas, 1903). The 8 mouth-arms are two-thirds as long as the diameter of the umbrella.
They are fused one with another in the upper thirds of their lengths and are free in their
lower two-thirds. There are 16 scapulets, 2 of which arise from the abaxial (outer) side of
each of the 8 upper arms. Each scapulet is simitar-shaped and forked at its outer end, and
is about half as long as the radius of the umbrella. There are frilled mouths and elongate
filiform appendages upon the upper side of each scapulet.

The lower arms are 3-winged or Y-shaped in cross-section, one wing being inward and
axial, the other wings being lateral and directed outwardly. Each of “these lateral wings
is of the shape of an equilateral triangle, and there are 4 elongate, sharp-pointed projections
from the abaxial angle of each wing. The pointed lowermost end of the lower arm terminates
in a large, club-shaped appendage, with a faceted, swollen end. This appendage is about
as long as the upper arm and there are other much swollen, club-shaped appendages which
arise between the frilled mouths of the 3 wings of each of the lower arms.

The canal-system of the umbrella consists of 16 radial-canals, 4 perradial, 4 interradial,
and 8 adradial, the adradial ones being nearer to the perradial than to the interradial canals.
All of the canals extend quite to the bell-margin, and all give off side branches which form
an anastomosing network. A definite ring-canal is not present. The circular muscle-system

PLATE 74.

Fig. 1. Rhopilema verrilli1z, mature medusa, 0.5 natural size. Bell shown
expanded. When in systole it is higher than a hemisphere. Off
Middletown, Pamlico Sound, North Carolina, November 16,
1904.

Fig. 1’. Rhopilema verrilli:. Marginal sense-organ seen from the exumbrella
side, showing the sensory pit in the exumbrella above the club.
The sensory-mass is orange. There is no ocellus.

Fig. 2. Crambione cooki, 0.66 natural size. Agassiz Expedition to the Great
Barrier Reef, off Cooktown, Queensland, Australia; May 4, 1896.

For figure 2 see page 677.

Drawn from life, by the author.

RHIZOSTOMAS—RHOPILEMA. 707

of the subumbrella consists of 16 triangular areas which alternate with and are widely separated
by the 16 radial-canals. ‘

This medusa was described by Vanhéffen from Hongkong, China, and later by Maas
from the Malay Archipelago. It appears to be identical with R. verrucosa Kishinouye, from
Japan. R. hispidum is possibly identical with Pilema clavigera Haeckel; but in Haeckel’s
medusa there are only 48 marginal lappets when the disk is 90 mm. wide, whereas in R.
hispidum of the same width there are 80 marginal lappets. The granular projections upon
the exumbrella in Haeckel’s medusa are bluntly rounded, whereas in R. hispidum they are
sharp and thorn-shaped. The terminal appendages of the mouth-arms appear to be some-
what longer in Haeckel’s medusa than in R. hispidum. Haeckel describes “* Pilema clavigera”
from a single alcoholic specimen from Hongkong, China.

Rhkopilema verrillii.
Plate 74, figs. 1, 1’.

Nectopilema verrillii, Fewkes, 1887, American Journ. Sci., ser. 3, vol. 33, p. 119, plate 4.

The disk is fully 350 mm. in diameter, hemispherical in contraction, but slightly flatter
than a hemisphere when expanded. The gelatinous substance is thick and ngid. The cen-
ter of the exumbrella is smooth, but over the lappets there are many shallow furrows and
the surface near the margin resembles sand-paper, being covered with numerous minute ele-
vations. There are 8 marginal sense-organs, each of which contains a terminal, entodermal
concretion-mass of red pigment granules. There is a simple, exumbrella sensory pit without
furrows; 64 marginal lappets. There are 6 large, oval lappets in each octant of the margin of
the disk, together with 2 small, oval lappets adjacent to the sense organs.

The arm-disk is cruciform and about half as wide as the bell, and the 4 perradial col-
umns are only three-fifths as wide as the heart-shaped, subgenital ostia. 8 short, tough,
gelatinous, lower mouth-arms, which arise from the arm-disk, are each about 180 mm. long
and Y-shaped in cross-section below. They branch sparingly and the very numerous mouths
are found upon their lower and inner sides, in furrows bordered by numerous, small, waving
cirri. There are about 25 to 60 blunt, translucent spindle-shaped appendages, which arise
from the lower sides of the mouth-arms, and are besprinkled with wart-like clusters of
nematocysts. The largest filaments arise from the principal crotches of the mouth-arms.

A pair of short flapper-like, lateral scapulets arise from the outer side of each of the
8 mouth-arms near its base, the mouth-openings of which are confined to their upper edges
and connect by a main duct in each scapulet with the axial-canal of the arm to which they are
attached. There are no appendages upon the scapulets. 4 short, gelatinous, perradial columns
connect the arm-disk with the subumbrella. There are 4 deep, heart-shaped clefts or genital
ostia between these 4 columns, but the bottom of each cleft is bridged over by a delicate
membrane. The 4 separate gonads develop within this membrane and the stomach is bor-
dered on the sides by the 4 stout, perradial columns and the 4 interradial membranes.
The 4 genital sacs are separated one from another; indeed the gonads are somewhat protru-
sive in old medusz.

There is a blunt wart-like papilla upon the subumbrella surface at a short distance
beyond the opening of each genital ostium, and this partially constricts the opening, giving
it a heart-shaped outline. The circular muscles of the subumbrella are very powerful, but
are almost interrupted in the radii of the 16 radial-canals in old meduse although they are
entire near the margin in young animals. Thus in old medusa there are 16 partially isolated
arcades of circular muscles as in Rhizostoma pulmo. There are no radial muscles.

The stomach is wide and cruciform, and corresponds in outline with the cruciform
arm disk under which it lies. Its exumbrella roof is plane, but the subumbrella floor dips
downward into the center of the arm-disk, and 4 open, perradial, gutter-like furrows extend
from this central depression down the center of each perradial column to the outer edge of
the stomach. 4 perradial ducts arise from the depressed center of the stomach at the
middle of the arm-disk, and these soon bifurcate giving 8 adradial ducts which extend down
each of the 8 mouth-arms giving off side branches to the numerous mouths.

~I
—

MEDUS® OF THE WORLD.

The axial ducts of the 16 small, flapper-like, lateral scapulets connect directly with
the axial duct of the arm to which they are attached. The large, central stomach gives rise
to 16 main radial-canals which extend outward to the 8 sense-organs and to intermediate
points on the bell-margin. These 16 main radial-canals give rise to numerous side branches
which anastomose in a network of vessels as in Rhizostoma pulmo. There is no definite ring-
canal in the adult.

The gelatinous substance of the medusa is translucent, dull, milky-yellow. The mouths
are rich yellow with chocolate-red blotches of pigment scattered at intervals at the bases of
the cirri. The ring-muscles of the subumbrella are a decided yellow, and according to Fewkes
the radial-canals are chocolate or rich chestnut in color. In the specimens studied by me,
however, they were yellow. The gonads are dull milky-yellow. There are numerous, small,
gastric cirri upon the gonads.

Fic. 424.—Rhopilema verrillii. Drawn by the author from a specimen obtained by Professor Verrill at
Outer Island near Branford, Connecticut.

Drawing one-half natural size. 6 of the mouth-arms are cut off close to their bases, and the scapulets are
cut off from 4 of them in order to show 2 of the subgenital ostia. The muscular arcades are
shown in one-half of the subumbrella and the canal-system in the other half. Thus in old
medusa there are 16 partially isolated arcades of circular muscles as in Rhizostoma pulmo.
There are no radial muscles.

This rare medusa was first found by Prof. A. E. Verrill, in 1886, in New Haven Harbor,
Connecticut, during September. In 1889, Professor Verrill again found it in considerable
numbers among the Thimble Islands about 10 miles east of New Haven, in Long Island
Sound, where they were common in August and September. They then disappeared, but
were again found at the Thimble Islands during the summer and autumn of 1903, and
again in Branford Harbor, Connecticut, in September and October, 1909. I secured the
specimen figured on plate 74 at Middleton, Pamlico Sound, North Carolina, in November,

RHIZOSTOM2—RHOPILEMA, EUPILEMA, STOMOLOPHUS. 709

1904, where it was swimming near the surface on a calm morning. The fishermen
informed me that it is seen quite frequently in Pamlico Sound in autumn. I believe, there-
fore, that it is a southern form which occasionally establishes itself in Long Island Sound.

The following are the dimensions in mm. of a specimen of Rhopilema vervillii found by
Professor Verrill at Outer Island, near Branford, Connecticut, in Long Island Sound, on
September 12, 1909.

Diameter of contracted bell, 218. Width of largest velar lappets, 11.

Diameter of fully-expanded bell, 268. Length of largest velar lappets, 12.5.

Perradial diameter of arm-disk, 135. Length of scapulets, 36.

Interradial diameter of arm-disk, 76. Length of mouth-bearing part of scapulets, 35.

Width of each perradial column of arm-disk, 33. Length of lower arms, 65.
Circumferential width of subgenital ostium, 49 Width at widest distal part of lower arms, 54+.
Radial width of subgenital ostium, 27. Width of lower arms at points of origin from arm-disk,
Width of zone of circular muscles, 54. 26 to 32.

Width of ocular lappets, 3. Length of longest mouth-arm appendages, 26.

Length of ocular lappets, 5. 5.

Genus EUPILEMA Haeckel, 1880.

Eupilema, Harcket, 1880, Syst. der Medusen, p. §90.—Vannorren, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p- 43-

The type species is Euphilema scapulare Haeckel, from the Malay Archipelago. Cyanea
rhizostoma Brandt, 1838, may be of the same genus, but is too imperfectly described to be
determined.

GENERIC CHARACTERS.

Rhizostomata scapulata with 8 free, 3-winged mouth-arms, without filaments, clubs,
or other appendages. 16 radial-canals all connected by an anastomosing network of vessels
in the outer parts of the subumbrella.

This genus is closely related to Rhizostoma, but has no mouth-arm appendages.

Eupilema scapulare Haeckel.
Eupilema scapulare, HAEcKEL, 1880, Syst. der Medusen, pp. 582, 590.—VaANHOFFEN, 1888, Biblio. Zool., Bd. 1, Heft. 3, p- 43.

Bell 150 mm. wide, 50 mm. high, hat-shaped with rounded dome. 8 rhopalia, 144 lappets.
In each octant 8 pairs of long, narrow, rectangular, projecting, velar lappets, between 2 small,
oval, ocular lappets. Mouth-arms not quite as long as bell-radius. Scapulets simitar-shaped,
as long as free, upper part of arm. 17 radial-canals all connected by an anastomosing network
of vessels which extend inward to the zone of the bases of the arm-disk pillars. The form
of the scapulets shows that they are only the secondarily separated, uppermost lappets of the
dorsal wings of the mouth-arms. Color (?) Sunda Archipelago, Sumatra.

Very briefly described from a preserved specimen by Haeckel.

Genus STOMOLOPHUS L. Agassiz.

Stomolophus, Acassiz, L., 1862, Cont. Nat. Hist. U.S., vol. 4, pp- 138, 151-—Acassiz, A., 1865, North American Acal., p. 40.—
Craus, 1883, Organisation und Entwick. Medusen, p. 60.—von LENDENFELD, 1884, Proc. Linnean Soc. New South Wales,
vol. 9, p- 292.—VANHOFFEN, 1898, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 30, 42-—Maas, 1903, Scyphomedusen der
Siboga Exped., Monog. 11, p. 80.

Brachiolophus+ Stomolophus, Harcket, 1880, Syst. der Medusen, pp. 597, 598-

The type species is S. meleagris L. Agassiz, which ranges from South America to the
mouth of Chesapeake Bay, United States, and is apparently found also on the Pacific side
of the Isthmus of Panama.

GENERIC CHARACTERS.

Rhizostomata scapulata with a central mouth-opening. With a well-developed, tube-like
manubrium formed by the fusion of the lateral edges of the 8 primitive mouth-arms. Only
the extreme ends of the mouth-arms are free and they branch complexly. 8 pairs of scapulets.
4 separate invaginated gonads. The central stomach gives rise to 16 radial-canals, which
are all connected by a marginal network of anastomosing vessels. 8 marginal sense-organs.

Brachiolophus Haeckel is only a young stage of Stomolophus.

710 MEDUS2 OF THE WORLD.

Stomolophus meleagris L. Agassiz.
Plates 75 and 76, figs. 1-3.
Cephea rhizostoma, Gisnes, 1847, (non Lamarck) Fauna of South Carolina.
Stomolophus meleagris, AGassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, pp. 138, 151; Ibid., 1860, vol. 3, plate 14, figs. 1-8.—
Acassiz, A., 1865, North Amer. eehoe Pp. 40.
Stomolophus meleagris+ S. agaricus ?, Harcxet, E., 1880, Syst. der Medusen, p. 599.
Stomolophus chunit, VANHOFFEN, E., "1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 31, 42, taf. 3, fign. 4, 5; taf. 4, fig. 1.—

Trinct, G., 1906, Annuar Mus. Zool. Napoli, ser. 2, vol. 2, No. 9, pp. 1-4.

Bell about 180 mm. in diameter, half-egg-shaped, higher than a hemisphere, the gelatinous
substance thick and rigid; no marginal tentacles; 8 marginal sense-organs, 4 radial and 4
interradial. Each sense-club is deeply set within a niche between the ocular lappets and is
protected above by a partial web between the lappets. Sense-club hollow and spindle-shaped,
terminating in a knob-like end which contains an entodermal mass of deeply pigmented
concretions. A deep, 3-sided, furrowed pit projects inward from the surface of the exumbrella
just above the base of each sense-club (see [c] fig. 2’, plate 75). About 128 marginal lappets,
16 in each octant, those flanking the sense-organs being about 3 times as long as the others.
The velar lappets have rounded edges, but the ocular lappets are longer and sharp-pointed.

A rigid, thick-walled mouth- pabe. or manubrium, projects downward from the center
of the subumbrella and extends 40 to 50 mm. beyond level of bell-margin. This manubrium-
like tube is formed by the lateral coalescence of the 8 adradial mouth-arms, which are fused
along their adjacent sides, their extreme ends only being free. These free ends of the mouth-
arms bifurcate and flare outward at lower end of manubrium. A deep groove, or gutter,
extends along the ventral side of each of the 8 mouth-arms and branches dichotomously twice
and extends outward over the lower sides of the 16 free ends of, the mouth-arms. The free
edges of this branching groove are complexly branched and folded and are lined by a row of
numerous small knobbed tentacles, which move constantly in such manner as to drive food
particles into the mouth-grooyes. In addition to the central mouth, there are numerous slit-
like lateral mouths which are situated upon 16 knife-blade shaped scapulets attached to the
upper part of,and occupying more than half of the length of, the manubrium. The free edges
of these lateral mouths are complexly crenulated and lined with small, incessantly waving
tentacles exactly as are the mouth-grooves at the free extremity of the manubrium (fig. 2,
plate 76). The 8 main mouth-grooves of the manubrium lead into a 4-cornered, central
cesophagus which extends upward into the wide, central stomach. 16 lateral branches, 4
from each side of this four-cornered cesophagus, extend outward to the slit-like mouths in
the 16 scapulets. These radiating tubes arise from the cesophagus near the level of the
upper parts of the scapulets. The central stomach is wide and lenticular, and occupies the
midst of the umbrella. It gives rise peripherally to 16 radial-canals which in their outer
halves give off many side branches which in turn form an anastomosing network placing all
of the radial-canals in connection one with another, as in the genus Rhizostoma. No ring-
canal.

There are 4 deep, cylindrical subgenital pits, and a blunt papilla is found on the sub-
umbrella surface just beyond the opening of each genital ostium. The gonads are found in
4 folded regions in the wall of the subumbrella at the bottom of the subgenital pits. There
are 16 semi-elliptical areas of circular muscles in the subumbrella, 2 between each succes-
sive pair of sense-organs. Alternating with the circular-muscle areas there are 16 narrow,
triangular strands of weakly developed radial-muscle-fibers, the broad end of each triangle
abutting against the side of the manubrium, and the narrow, pointed end being directed
outward. 8 of these occupy the radii of the marginal sense-organs and 8 are intermediate in
position. The bell-margin pulsates incessantly with remarkable strength and rapidity.

The gelatinous substance of the bell is of a milky bluish or yellowish color and the ento-
dermal parts are dull yellow. The surface of the exumbrella is reticulated with brown pigment
which is especially dense near the margin. There are numerous white or yellowish spots in this
brown marginal zone. The mouth-frills are brownish-pink.

Young medusa.—\ haye captured an immature medusa of this species in which the bell
was 3 mm. in diameter and the entire animal 5 mm. in length (fig. 3, plate 75). The bell
was flatter than a hemisphere and the surface of the exumbrella was covered with wart-like
clusters of nematocysts, among which there were numerous, brown-colored pigment cells.

PLATE 75.

All figures are of Stomolophus meleagris.

fig. 1. Mature medusa, 0.66 natural size. In the ocean off Fernandina,

Florida, January 3, 1905.

ig. 2. Young medusa, 1.5 times natural size. Charleston Harbor, South
Carolina, August 28, 1897.

. 2’. Sagittal section of the medusa shown in figure 2.

. 2”. Cross-section of the fused mouth-arms of the medusa shown in
figure 2, cut at the level of the 16 scapulets.

ig. 2’. Marginal sense-organ of the medusa shown in figure 2, showing

the exumbrella pit above the sense-organ. There is no ocellus.

ig. 3. Young medusa, 5.5 mm. long and 3 mm. in diameter, showing the

lateral mouths (scapulet mouths) beginning to break through the

walls of the primitive cesophagus. Charleston Harbor, South

Carolina, September 9, 1898. An oral view of this medusa is

shown in figure 1, plate 76.

Drawn from life, by the author.

PLATE 75

MAYER

Avoenk ta

PLaTE 76.

All figures are of Stomolophus meleagris.

Fig. 1. Oral view of a young medusa 3 mm. in diameter. A side view of this
medusa is shown in figure 3, plate 75.

Fig. 2. Sagittal interradial section of mature medusa shown in figure 1, plate
74. Also cross-sections of the mouth-arm-tube at three different
levels. g, gonad; p, subumbrella papilla in the radius of the sub-
genital pit; S, stomach cavity; sg, subgenital pit.

a
°Q

ig. 3. Oral view of a quadrant of subumbrella, omitting the mouth-parts.
The sector AB, shows the canal-system of the subumbrella, and
BC two of the 16 sectors of circular muscles.

Drawn from life, by the author.

MAYER

ce

Cc ‘

A
sii

iN

ASS

eo Gi [

= ait UAT f
SquGUSCGG0 Ng
PAAMANTVANAM

PLATE 76

AHoon Ce

RHIZOSTOM.2—STOMOLOPHUS. 711

There were 8 marginal sense-organs and 48 marginal lappets (fig. 1, plate 76). The lappets
flanking the sense-organs were about twice as long as the others. The ocular lappets were,
however, simple while the others were bifurcated and evidently in process of division. The
central mouth was situated at the extremity of a long 4-cornered proboscis which possessed 4
bifurcated lips. The free edges of these lips were lined by a row of short, slender, knobbed
tentacles which maintained a constant motion. In addition to the principal or terminal mouth
there were 8 small, tube-like, lateral mouths arranged in 4 pairs, the beginnings of the scapulets.
These mouths arose from the sides of the manubrium near its base and were interradial in
position (7. ¢., go° from the radii of the 4 principal lips), and in addition to these lateral mouths
there were 4 pairs of hernia-like projections upon the surface of the manubrium. These pro-
jections alternated in position with the already functional, lateral mouths, and would no doubt
soon have broken through and formed another set of such mouths (fig. 1, plate 76). The
functional mouths were each surrounded by 8 tentacles which were similar in structure to the
tentacles lining the free edges of the principal mouth.

The medusa was quite transparent except for a trace of brown pigment in the ectoderm
of the exumbrella and the dark-red pigment of the sense-organs.

I found it in Charleston Harbor, South Carolina, on September g, 1898.

The resemblance between this young rhizostomous medusa and the adult condition in
the Semzostomez is very striking. The terminal mouth was used, indeed, for the capture of
food, an operation which was facilitated not only by the flexibility of the lips but also by the
incessant motion of the tentacles.

This species is very common along the sandy coasts of North and South Carolina and
Georgia. In April, 1910, mature meduse were abundant at Tortugas, Florida. It does not
extend north of the mouth of Chesapeake Bay. It is found along the northern coast of South
America, and at many places in the Gulf of Mexico, and I believe it to be identical with S.
chuntu Vanhoffen, from the Bay of Panama on the Pacific side of the Isthmus. It often occurs
in yast swarms, occupying an area which is sometimes over 100 miles in length. Mature
individuals are abundant in winter and spring off the coast from Florida to South Carolina.
It is not often seen in brackish harbors, but is practically confined to pure ocean water off
the coast. At most it enters only the mouths of harbors.

I can see no difference between S. chuni Vanhoffen and S§. meleagris. S. chunit is
described as being only 90 mm. wide and with only 112 marginal lappets. In all respects it
resembles a half-grown S. meleagris. Indeed, Trinci, 1906, records S$. chuni: from the Gulf
of Paria between Trinidad and Venezuela, Atlantic coast of South America, and it appears,
therefore, that “‘S. chunii”’ must occur on both Atlantic and Pacific sides of the Isthmus of
Panama. It is probable that the medusa has remained unchanged since the closure of the
Isthmus in Mesozoic times.

Haeckel’s Brachiolophus collarts is only a younger stage of the same medusa with a bell
80 mm. wide and with 80 marginal lappets. It is described from the Galapagos Islands.

Stomolophus meleagris var. fritillaria.

Stomolophus fritillaria, HaecKet, 1880, Syst. der Medusen, p. 598, taf. 35, fign. 1-9—von LenpenFeLp, 1884, Proc. Linnean
Soc. New South Wales, vol. 9, p. 292—VanuorreN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, pp. 31, 42.

This form appears to be smaller than S§. meleagris, the bell being only about 80 mm. in
diameter and 60 mm. in height. The marginal lappets are more numerous, being 208 in
number. The ‘‘manubrium” or fused tube of mouth-arms extends only about one-fourth
the bell-height beyond the level of the margin, instead of about one-third the bell-height, as
in S. meleagris. This medusa is found at Suranim on the Atlantic coast of South America.
Color (? )

The only valid distinctions between this medusa and S. meleagris are in its large number of
marginal lappets, and in the cleft in the middle of each octant of velar lappets. Also the 16
scapulets are hidden well up under the bell instead of extending down to about the level of
the bell-margin. It may be regarded as a southern variety of S. meleagris. Haeckel describes
it from 3 alcoholic specimens in the Copenhagen museum.

712 MEDUS OF THE WORLD.

RHIZOSTOMATA SIMPLICIA Vanhoffen, 1888.

Archirhizide, Haxcker, 1880, Syst. der Medusen, p. 565.—Craus, 1883, Organisation und Entwick. Medusen, Leipzig.—von

LenpENFELD, 1888, Zeit. fiir wissen. Zool., Bd. 47, p. 210.

Rhizostomata simplicia, VANHOFFEN, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 39-

These are probably only immature or torn and regenerating forms which are rendered
still more unnatural through shrinkage in alcohol. I present this account of them merely in
the hope that some may oe cadiecaael At present they are wholly apocryphal. A des-
cription of the genera follows:

Archirhiza Haecxer, 1880. 8 free mouth-arms, 4 separate, subgenital cavities.

Haplorhiza Haecxet, 1880. 8 free mouth-arms. A unitary, subgenital cavity.
Cannorhiza Harcke, 1880. Mouth-arms fused along their sides, forming a mouth-cylinder.

Haeckel is the only naturalist who has seen any of these forms. They are all small and I
incline to the belief that they are merely immature stages or injured and regenerating
specimens of various other rhizostome in the condition preceding the development of the
ultimate ramuli of the mouth-arms. Vanhdéffen (1902, Wissen. Ergeb. Valdivia Exped., Bd.
3, Lfg. 1, p. 52) believes them to be merely mutilated medusz with the branches and appen-
dages of the mouth-arms lost or reduced. I have recorded them merely because they may still
have a place in literature if not in the ocean.

Genus “ARCHIRHIZA” Haeckel, 1880.
Archirhiza, Haxcket, 1880, Syst. der Medusen, p. 565.—Vanuorren, 1888, Bibliotheca Zoologica, Bd. 1, Heft. 3, p. 39-

GENERIC CHARACTERS.

Rhizostomata simplicia with 8 simple separate, unbranched mouth-arms. With 4 separate,
subgenital cayities. With 16 radial-canals, some or all of which may give rise to anastomosing
side branches. The ring-canal gives off an anastomosing network of vessels which ramify
through the marginal lappets. The mouths have no appendages and are found only on the
ventral sides of the mouth-arms. 8 rhopalia.

Haeckel founded this genus for Archirhiza primordialis from Bass Strait, Australia. Later
he describes another medusa, 4. aurosa, from New Zealand, which is apparently only a later
stage in the growth of his 4. primordialis. Indeed, I suspect that both of these medusz are
immature, or “‘reconstructed” from fragmentary specimens.

Archirhiza aurosa Haeckel.

Archirhiza primordialis (young ?), Harcxer, 1880, Syst. der Medusen, p. 565, taf. 36, fign. 1, 2.—Hamann, 1881, Jena Zeit.
fiir Naturw., Bd. 15, p. 245 (anatomy of mouth-arms).—Vannorren, 1888, Bibliotheca Zoologica, Heft. 3, p- 39-
Archirhiza aurosa (mature ? ), HaEcKet, loc. cit., p. 645.

Young medusa (?).—Bell flatly and evenly rounded, hemispherical in contraction, 2 to 3
times as wide as high when expanded. 40 mm. wide. Exumbrella finely granulated as in
Aurellia. 8 rhopalia, perradial and interradial. 48 marginal lappets. In each octant 2 large
median, flanked by 2 smaller velar lappets, and with 2 still smaller, rhopalar lappets flanking
the sense-organs. All of the lappets are pointed. Diameter of arm-disk two-thirds as wide
as bell-radius. 8 simple, separate, fleshy, unbranched, recurved mouth-arms arise in 4 pairs
on either side of each perradial corner of arm-disk. These mouth-arms lack appendages,
but there is a zigzag row of fringed mouths along the ventral side of each arm. These 8
lines of mouths of the mouth-arms fuse into 4 perradial lines over the mouth-arm-disk. The
mouth-arms are shorter than the bell-radius and are simitar-shaped, but fleshy and blunt at
their ends. 4 separate interradial genital sacs are invaginated into the stomach cavity, so the
arm-disk displays 4 interradial subgenital ostia. The central stomach gives rise to 16 radial-
canals, of which the 8 perradial and interradial canals give off branching side branches near
the bell-margin. The 8 adradial canals are simple. All 16 canals and their side branches fuse
with a well-developed ring-canal at the zone of the rhopalia, and on its outer side this ring-
canal gives off a close-meshed network of small vessels which anastomose through the marginal
lappets. Color (?) Found in Bass Strait between Australia and Tasmania.

RHIZOSTOMAZ—ARCHIRHIZA, HA PLORHIZA, CANNORHIZA, lis

It is highly probable that the medusa described above is only the young of the form
called Archirhiza aurosa, by Haeckel, from New Zealand. This is larger, being 50 mm. in
diameter. There are 80 instead of 48 marginal lappets. In each octant 8 oval, pointed velar,
between 2 small, triangular, rhopalar lappets. Mouth-arm conical, more pointed than in
A. primordialis, and 1.5 times as long as bell-radius. All 16 of the radial-canals give off
anastomosing side branches.

Altogether, the differences between /. “primordialis” and A. aurosa are precisely such
as one would expect to occur during the growth of the medusz. They are probably only young
stages of some species of Catostylus.

“Genus HAPLORHIZA” Haeckel, 1880.
Haplorhiza, HAarcket, 1880, Syst. der Medusen, p- 604.—VanuorreNn, 1888, Bibliotheca Zoologica, Heft. 3 P- 39-

GENERIC CHARACTERS,

Similar to Archirhiza but with a unitary subgenital cavity instead of 4 cavities as in
Archirhiza.

This single, subgenital cavity or “porticus” is cruciform in shape and formed by the
fusion of the 4 primitive interradial genital sacs and the breaking down of the walls in the
fused regions, thus forming a cross-shaped cavity beneath the stomach. It is in communication
with the outer world through the 4 genital ostia, and is separated completely from the central
stomach. Its side walls constitute partitions between it and the central stomach and they

contain the genital products.
Haplorhiza simplex Haeckel.

Haplorhiza simplex, Harcxer, 1880, Syst. der Medusen, p. 604.

Bell flat, shield-shaped, 40 mm. wide, 20 mm. high. 8 thopalia. 48 marginal lappets.
In each octant 4 large, quadratic velar, between 2 tongue-shaped, projecting, ocular lappets.
8 simple, cylindrical mouth-arms, as long as bell-radius. 4 interradial, subgenital ostia, not
quite as wide as the arm-disk-pillars between them. 16 radial-canals and a ring-canal. Bass
Strait, southern Australia.

This medusa is very briefly described by Haeckel, but in all respects it appears to be an
immature stage of some other rhizostomous medusa such as Catostylus. Its 8 simple mouth-
arms lack appendages and resemble those of the immature Archirhiza primordialis Haeckel.
The mouths are confined to the ventral sides of the mouth-arms.

Haplorhiza punctata Haeckel.

Haplorhiza punctata, HAEcKEL, 1880, Syst. der Medusen, p. 604.

Bell flatly rounded, 40 mm. wide, 20 mm. high. 8 rhopalia. 176 marginal lappets, in
each octant 10 pairs of small, rounded velar lappets between 2 small, rhopalar lappets. 8
simple, cylindrical mouth-arms, hardly half as long as bell-radius, arise in pairs from the 4
perradial angles of the wide 4-cornered arm-disk. The frilled mouths are confined to the
ventral sides of the mouth-arms and there are no appendages. The 4 subgenital ostia are
3 times as wide as the columns between them.

Exumbrella dark violet-brown, besprinkled uniformly with round white points.

Arnheims Land, coast of northern Australia. This medusa is apparently immature.

“Genus CANNORHIZA”’ Haeckel, 1880.
Cannorhiza, HarcKet, 1880, Syst. der Medusen, p. 605.—VaNHOFFEN, 1888, Bibliotheca Zoologica, Heft. 3, p. 39.

The type species is Cannorhiza connexa, Haeckel, from the neighborhood of New Zealand,

South Pacific.
GENERIC CHARACTERS.

Rhizostomata simplicia with 8 simple, unbranched mouth-arms, the sides of which are
fused one to another forming a hollow mouth-arm-cylinder, and leaying only the lower ends
of the arms free. There were neither clubs nor filaments upon the mouth-arms. There is a
unitary subgenital porticus. 8 rhopalia. The central stomach gives rise to 24 branching radial-
canals which fuse with a ring-canal. Peripherally the ring-canal gives off a network of vessels

which ramify through the lappets.

714 MEDUS OF THE WORLD.

Cannorhiza connexa Haeckel.

Cannorhiza connexa, Harcxet, 1880, Syst. der Medusen, p. 605, taf. 40, fign. 1-8.—Vanuorren, 1888, Bibliotheca Zoologica,

Heft. 3, p- 39-

Bell flatly rounded, exumbrella surface finely granulated, 80 mm. wide, 30 mm. high.
Gelatinous substance tough and horny in consistency. 8 rhopalia. 80 marginal lappets.
In each octant are 8 small, rectangular, velar lappets between 2 small, oval, pointed, rhopalar
lappets. The 8 mouth-arms are fused along their sides forming a mouth-arm- cylinder. Each
mouth-arm is cylindrical, somewhat longer than the bell-radius. The short, free, lower end
of each mouth-arm is simple, inbranched, curved outward and somewhat upward. The
lateral fusions of the 8 adradial mouth-arms are marked by 8 longitudinal furrows, the 4
perradial being shallower than the 4 interradial. The frilled mouths are developed only on
the lower sides of the mouth-arms and there are neither filaments, clubs, nor other appendages.
The 4 interradial, subgenital ostia are not quite as wide as the perradial spaces between them,
and the arm-disk is not quite as wide as the bell-radius. There is a cruciform unitary sub-
genital porticus and the 4 gonads are in the side walls of this space, the cruciform roof being
thick and gelatinous. The 4 genital membranes are much folded. The cruciform central
stomach lies above the subgenital porticus from
which it is completely separated by the lateral
genital membranes and gelatinous, cruciform
roof of the porticus.

A wide canal extends downward from each
of the 4 perradial corners of the stomach, through
the columns, into the arm-disk, where they fuse
at the center, forming a small cavity from which
arise the 4 pairs of canals leading down the lower
sides of the 8 adradial mouth-arms. Each mouth-
arm canal gives off numerous short side branches
which lead to the frilled mouths. 24 radial-
canals arise from the margin of the cruciform
stomach and extend outward into the subum-
brella. The 8 perradial and interradial canals
Fic. 425.— Cannorhiza connexa,” after Haeckel, in Das ai about twice as wide as the others. The 4

Soak der Meducen: perradial canals are very short and the 4 inter-
j radial very long. All 24 canals give off anas-
tomosing side branches and then fuse with the ring-canal. The ring-canal, in turn, gives
rise on its outer side to a fine-meshed network of narrow, anastomosing vessels which ramify

through the lappets. Found near New Zealand, South Pacific. Described in detail by Haeckel.

poe a

“Stomatonema reticulatum’”’ Fewkes.

Stomatonema reticulatum, Fewxes, 1884, American Naturalist, vol. 18, p. 300.

Bell 36 mm. wide, with thick walls, thinner at margin. There are no marginal tentacles.
There are 8 marginal sense-organs which bear some resemblance to those of Aurellia. Shape
and number of marginal lappets (?) The 8 mouth-arms arise from the arm-disk by 4 attach-
ments and are bordered on their lower sides by a double row of mouths which also extend
half-way up the upper side of each mouth-arm. 4 large, globular, ovarian sacs lie in the
interradii of the arm-disk alternating with the 4 primary branches of the mouth-arms. A
number of radial-canals arise from the central stomach and fuse with a narrow zone of anas-
tomosing vessels at the bell-margin. Fewkes does not state how many radial-canals there are
in this medusa, nor does he give any account of the marginal lappets, musculature, relative
sizes of the parts of the medusa, color, gonads, or mouth-arm-appendages. He mentions a
single specimen found in Montevideo Harbor, Atlantic coast of South America.

The description given by Fewkes is unfortunately too fragmentary even for generic
determination, but apparently the medusa is related to Haeckel’s Aurosa, but is distin-
guished by having mouths on the dorsal as well as on the ventral sides of the 8 mouth-arms.

. FOSSIL MEDUS®.

~I
i
on

FOSSIL, MEDUSAE.

Fossil medusa, or in many cases fossils supposed to be those of medusa, have been
described from the lower Cambrian and Devonian, and especially from the Jurassic litho-
graphic slates of Bavaria wherein their preservation is so perfect that in some cases, as in
Paraphyllites distinctus, they can be classified accurately with relation to living forms.

Owing to the uncertainty with which we must regard many of these fossils | have deemed
it best to group them together and not to attempt the fruitless task of classifying them, except-
ing in cases wherein their state of preservation warrants such a procedure.

A thorough review accompanied by excellent illustrations of all fossil medusze known
previous to 1898 is given by Walcott in Monographs U. S. Geological Survey, vol. 30, 1898,
and this work should be consulted by all students of the subject, for the account here given
does not attempt to present detailed descriptions.

Medusina radiata Walcott.

Astylospongia radiata, LiNNARSSON, 1871, K6ngl. svensk. Vet.-akad. Handl., vol. 9, No. 7, p. 13, plate 2, figs. 15, 16.

Medusites radiatus, Natuorst, 1881, Kongl. svensk. Vet.-akad. Handl., vol. 19, No. 1, p. 25, plate 6, figs. 1, 2.—(?) Pompecky,
1896, Jahrbuch, K.-k. geol. Reichsanstalt, Bd. 45, p. 501, taf. 14, fig. 3.

Medusina radiata, Wacotr, 1898, Monographs U. S. Geol. Surv., vol. 30, p. 56, plate 28, Lie ze

This fossil medusa from the lower Cambrian sandstone of Sweden is probably one of the
AEquoride and bears a general resemblance to the genus Zygodactyla. Bell 40 to 60 mm. wide.
Central stomach 25 to 30 mm. wide. 130 to 150 radial-canals which occasionally branch.
Subumbrella with radiating string-of-pearl-like papilla as in Zygodactyla. Nathorst describes
these papille as being upon the radial-canals, but to me it seems more probable that they are
interradial as in Zygodactyla.

Pompeckj’s somewhat similar medusa from the middle Cambrian beds of Bohemia is

probably an 4quorea, and not a Zygodactyla.
Medusina princeps Torell.

Protolyellia princeps, Torett, 1870, Lunds Universitets Arks-Skcift, 1869, No. 8, p. 10.

Astylosponiga radiata, LaNNaRssoN (in part), 1871, Kéngl. svensk. Vet.-akad., Handl., Bd. 9, Nr. 7, p. 13, taf. 2, fig. 15.
Medusites favosus, NaTuorRsT, 1881, Kongl. svensk. Vet.-akad. Handl., Bd. 19, Nr. 1, p. 25, taf. 25, taf. 5, fign. 5, 6 (?).
Medusites princeps, MarrHew, 1890, Trans. Royal Soc. Canada, vol. 8, p. 140.

Medusina princeps, Waicotr, 1898, Monographs U. S. Geol. Surv., vol. 30, p. 54, plate 28, fig. 1.

This fossil from the lower Cambrian of Sweden is of doubtful affinities. Nathorst’s
conclusion that it is one of the Cyaneide because the exumbrella floor of the stomach exhibits
polygonal facets appears to me to be too venturesome, for the radiating furrows of the sub-
umbrella resemble the radial-canals of A’quorea.

Medusina deperdita Walcott.

Acalepha deperdita, Beyricu, 1849, Zeitschrift deutsch. Geol. Gesell., Bd. 1, p. 437, taf. 39, fig. 1.
Medusites deperditus, HarcKe1, 1865, Zeitschrift fiir wissen. Zool., Bd. 15, p. 506, taf. 39, fig. 1.
Medusina deperita, Waucort, 1898, Monograph U. S. Geol. Surv., vol. 30, p. 91, plate 44, fig. 1.

It is possible that this fossil medusa from the Jurassic limestone of Eichstadt may belong
to the genus Cunoctantha, but there are no tentacles and we have only the 8 stomach-pouches
upon which to hazard this inference The bell is70mm.and the diameter through the stomach-
pouches 45 mm. wide.

Paraphyllites distinctus Maas.

Paraphyllites distinctus, Maas, 1906, Neu. Jahrb. Min. Geol., Pakiontol., Bd. 2, p. 90, 4 fign.

The preservation of this fossil from the Jurassic lithographic slates is so perfect that its
affinities with living medusz can be determined. It is therefore described on page 549 in
connection with Paraphyllina to which it is closely related.

716 MEDUSZ OF THE WORLD.

Cannostomites multicirrata Maas.
Cannostomites multicirrata, Maas, 1902, Palaontographica, vol. 48, p. 303, taf. 23, fig. 1, text-fig.

This is a fossil from the Jurassic lithographic slates of Bavaria. A single specimen was
studied by Maas. It may possibly be allied to 4tolla. Only the subumbrella is known. Bell
about 100 mm. wide. An open, central mouth with 4 perradial, gelatinous columns. Sub-
umbrella simple with 4 interradial sickle-shaped, notched, and swollen gonads. Marginal
ring-muscle entire and powerfully developed as in Atolla. Numerous marginal lappets of
varying sizes arranged in multiples of 4. Short, simple tapering tentacles arise from notches
between the lappets.

Atollites minor and Atollites zitteli Maas.
Atollites minor and A. zitteli, Maas, 1902, Paldontographica, Bd. 48, p. 319, taf. 23, fign. 5 und 6.

These fossils are from the lower chalk of Carpathia from the Warnsdorf strata. There
are more than 10 marginal lappets in high relief, quite variable in size and number. A small,
plain center of the exumbrella and an intermediate zone with raised streaks radiating out-

wardly. The meduse may be allied to 4tolla. They are well figured by Maas.

Acraspedites antiquus Haeckel.

Medusites antiquus, Harcket, 1865, Zeit. fiir wissen. Zool., Bd. 15, p. 509, taf. 39, fig. 2.

Acraspedites antiquus, HAECKEL, 1869, Zeit. fiir wissen. Zool., Bd. 19, p. 559; 1880, Syst. der Medusen, p. 647.—von Amnon, 1886,
Abbandl. Math.-phys. Classe Kénigl. bayer. Akad. Wissen., Bd. 15, p. 157-—Watcorrt, 1898, Monographs U.S. Geol.
Sury., vol. 30, p. 75, plate 44, fig. 2.

This is the vague impression of a medusa in the Jurassic white coral limestone, litho-
graphic slate of Eichstadt, Bavaria. It is about 140 mm. in diameter and appears to have 8
marginal lobes. The sculpturing of the bell may place it among the Coronata, but Haeckel
is inclined to classify it with the Pelagidz.

Semaeostomites zitteli Haeckel.

Semaeostomites zitteli, HanckeL, 1874, Jena. Zeitsch. fiir Naturw., Bd. 8, p. 323, taf. 11; 1880, Syst. der Medusen, p. 647.—VoN
Ammon, 1886, Abhandlung Math.-phys. Classe K6nig]. bayerischen Akad. Wissen., Bd. 15, p. 157.—Watcort, 1898,
Monographs U. S. Geol. Surv., vol. 30, p. 70, pl. 39, text-fig. 17.

This semzostomous medusa is found fossil in the Jurassic lithographic slate of Solenhofen,
Bavaria. According to Haeckel the disk is 80 mm. wide, mouth with 4 lips each about 80 mm.
long and 10 mm. wide. Central stomach cruciform. 4 interradial, elliptical, genital pouches
each 8 to 10 mm. wide. 16 unbranched (?) radial-canals, 4 perradial, 4 interradial, and 8
adradial. A ring-canal at some distance inward from the margin. 120 to 128 marginal lappets,
and an equal number of tentacles each about 30 mm. long. This medusa is apparently one
of the Ulmaridz allied to Discomedusa.

Eulithota fasciculata Haeckel.

Eulithota fasciculata, Harcke1, 1869, Zeit. fiir wissen. Zool., Bd. 19, p. 549, taf. 42, fign. 1,2; 1880, Syst. der Medusen, p. 647.—
von Amon, 1886, Abhand]. Math.-phys. Classe Konig]. bayer. Akad. Wissen., Bd. 15, p. 157.—Watcorr, 1898, Mono-
graphs U. S. Geol. Surv., vol. 30, p. 73, plate 45, figs. 3, 4-

This is a fossil medusa from the Jurassic lithographic slate of Solenhofen, Bavaria.

16 marginal lappets. 8 clusters, each composed of at least 4 tentacles, which arise from
8 perradial and interradial (?) thickened pads between the lappets. These pads may represent
the 8 rhopalia. 16 crescentic gonads around the stomach-margin. 16 interlobular radial-
canals. 4(?) simple, short lips. The affinities of this medusa are uncertain but its nearest
living ally appears to be Sthenonia or Poralza.

Myogramma speciosum Maas.

Myogramma speciosum, Maas, 1902, Palaontographica, Bd. 48, p. 298, tafn. 22, 23.—von Ammon, 1908, Geonostischen Jahres-
heften, Jahrg. 19, p. 174, fign. 2, 3-

This fossil is from the lithographic slate of Solenhofen. Discomedusz with flatly rounded
bell. 4-rayed in the organization of the central stomach. 8-rayed at the bell-margin. In the

FOSSIL MEDUS&, 717

subumbrella a wide marginal ring-muscle, an intermediate zone of 16 feathered arcades as in
Casstopea, and an inner zone of ring-muscles around the central stomach.

At the bell-margin there appear to be numerous thickly set, short, branched, tree-like
tentacles. The mouth-parts are lost. Maas studied 3 imperfect specimens of this remarkable
medusa which appears to be distinguished from all other Scyphomedusz by its branched
marginal tentacles. His best preserved specimen was 300 mm. wide.

Genus MEDUSINA Walcott, 1898.

Medusina, Warcott, 1898, Monograph U. S. Geol. Sury., vol. 30, p. 49.

Walcott proposes the name Medusina to designate all fossil medusa whose generic char-
acters can not be determined.

Among the most obscure are M. quadrata, M. bicincta, M. staurophora, M. circularis,
and M. porpitina from the Jurassic lithographic limestone of Bavaria, and M. atava from the
Permian. All of these are well described by Walcott, Joc. cit., pp. 93 to 96. Beyond the fact
that these fossils are apparently medusz not much can be said of them, and in most cases we
can not be certain as to whether they are Scyphomedusez or Hydromeduse.

Medusina costata Walcott.

°
Spatangopsis costata, TORRELL, 1870, Lunds Universitets Ars-Skrift, 1869, No. 8, p. 11.
A gelacrinus lindstromt, LINNARSSON, 1871, Kongl. svensk. Vet.-akad. Handl., Bd. 9, Nr.7, p- 11, taf. 1, fign. 6-7, taf.2, fign. 10-14.
Medusites lindstrémi, Scumivr, 1888, Mém. Acad. Imp. Sci. St. Pétersbourg, sér. 7, tome 36, p. 27, planche 2, figs. 34, 35-—WaL-
corr, 1891, Tenth Ann. Report, U. S. Geol. Sury., Part 1, plate 56, figs. 1-1c.
Medusites costatus, MattHEW, 1890, Trans. Royal Soc. Canada, vol. 8, p. 142.
Medusina costata, Watcotr, 1898, Monographs, U. S. Geol. Surv., vol. 30, p. 49, plates 29, 30.
This fossil from the lower Cambrian sandstone of Sweden, at Lugnas and Timmerdala

in Mount Billingen, is probably an durellia. The best description is presented by Walcott.

Medusina geryonides Huene.
Medusina geryontdes, HuENE, 1901, Neues Jahrbuch fiir Mineralogie Geol. und Palaeontol., Bd. 1, p. 1, taf. 1, fign. 1, 2.—Fucus,

gor, Centralblatt fiir Mineral. Geol. und Palaeontol., Jahrg. 1go1, p. 166.

This fossil medusa is from the Murchison sandstone of Wiesensteig in Wiirtemburg. It
is about 28 to 30 mm. in diameter. In the center there is a crater-like elevation with an irreg-
ular, flat, 6-sided knob at its middle. Surrounding this crater-shaped center there is a zone of
12 radiating concavities, the deepest being 2.5 mm. deep. Huene suggests that this medusa
may have been related to the Geryonidz, but this is wholly problematical, and it appears to
me that it might equally well be a cast of Brooksella. Fuchs casts doubt upon its being a
fossil medusa.

Laotira cambria Walcott.
Laotira cambria, Watcotr, 1896, Proc. U.S. Nat. Mus., vol. 18, p. 613, plate 32, figs. 1-8; 1898, Monographs U.S. Geol. Surv.,

vol. 30, p. 32, plates 5-19, 21-23.

This is a fossil from the middle Cambrian shale of Coosa Valley, Alabama, and is supposed
to be that of a medusa. It is a remarkably variable form and quite distinct from any living
medusa. It probably reproduced by fission as does Gastroblasta.

Walcott defines it as Discomeduse with a lobate umbrella with 4 to 12 lobes in simple
forms and with a large number in the compound forms. No tentacles and no central mouth-
openings in the adult. A simple radial-canal in each lobe of the umbrella and in the inter-
radii. Oral arms represented by interradial lobes attached to the central axis and to the central
lobes. Described in full detail by Walcott, 1898. It shows a tendency to intergrade in its
simplest forms with Brooksella conjusa from the same formation.

Dactyloidites asteroides Walcott.

Buthotrephis (?) asteroides, Fircu, 1850, Trans. New York State Agricult. Soc., vol. 9, p. 863.

Dactyloidites bulbosus, Hatt, 1886, 39th Ann. Report State Museum Nat. Hist. New York, p. 160, plate 11, figs. 1, 2.

Dactyloidites asteroides, Watcort, 1891, 10th Ann. Report U.S. Geol. Surv., Part 1, p. 605, plates 57, 58, fig. 61; 1898, Mono-
graphs U. S. Geol. Surv., vol. 30, p. 41, plates 24-28.

This fossil is from the lower Cambrian terrane of Eastern New York at Penrhyn Quarry,
Middle Branville, Washington County, and from St. Albans, Vermont. It is an irregular,

718 MEDUSZ OF THE WORLD.

star-like fossil, and sometimes two stars are joined by a single lobe. It appears to have been
eregarious, for Walcott records 42 specimens on a slab of slate 37 by 62 inches. There is of
course no proof that these fossils are those of medusz.

Rhizostomites admirandus Haeckel.
Rhizostomites admirandus, Haxcket, 1866, Neues Jahrbuch fiir Min. Geol. und Paleontol., p. 261, taf. 5; 1869, Zeit. fiir wissen.

Zool., Bd. 19, p- 557; 1880, Syst. der Medusen, p. 647.—Branpt, 1871, Mém. Acad. Imp. Sci., St. Pétersbourg, sér.7,

tome 16, p. 1, planche 1, figs. 1-4.—von Amon, 1886, Abhandl. Math.-phys. Classe, Akad. Wissen Miinchen, Bd. 15,

p- 123, 158, 163, taf. 2, fig. 2.—Watcort, 1898, Monographs U.S. Geol. Surv., vol. 30, p. 76, plates 40, 42.—Maas, 1902,

Palaeontographica, Bd. 48, p. 306, taf. 23, fig. 2——von Amon, 1908, Geonostischen Jahresheften, Jahrg. 19, p. 174, fig. 1-

yon Ammon, who has made the most thorough study of this fossil from the Jurassic
lithographic limestone of Solenhofen and of Eichstadt, Bavaria, has decided that it is
identical with Haeckel’s Rhizostomites lithographicus, and that Leptobrachites trigonobrachius
is probably the same medusa turned over on its side. He also concludes that Haeckel’s Hexa-
rhizites insignis is only a 6-rayed aberration of the same medusa. A thorough review of the
literature of this subject and excellent figures are presented by Walcott, 1898, Joc. cit.

According to von Ammon and Walcott the disk in Rhzzostomites is round and as large
as 400 mm. in diameter, with 4 to 8 principal lobes and about 128 small marginal lappets of
various sizes, and indentations of the bell-rim marking the places of the 8 marginal sense-
organs. No marginal tentacles. A wide zone of circular muscles in the subumbrella, unbroken
in the rhopalar radii. 16 radial-canals, 8 rhopalar and 8 inter-rhopalar; and a circular canal in
the external third of the umbrella. A strongly marked, circular depression between the muscle-
zone and the arm-disk may indicate an inner ring-canal. 4 not very wide subgenital ostia,
with 4 opercula forming lappets. Probably 8 long, thin mouth-arms with crinkled appen-
dages, and apparently with a tassel-shaped tuft at the lower end.

This is undoubtedly a Rhizostomous medusa which appears to belong to an extinct genus
related to the modern Rhizostomata triptera or lorifera. Maas, 1902, gives a remarkably clear
photograph of the margin showing one of the sense-organs, and he discusses the probable
form of the gonads.

Brooksella alternata Walcott.
Brooksella alternata, Waucort, 1986, Proc. U. S. Nat. Mus., vol. 18, p. 612, plate 31, figs. 1-5; 1898, Monographs U. S. Geol.

Surv., p. 23, plates 1-4.

This fossil from the middle Cambrian shale from Coosa Valley, Alabama, is supposed to
be that of a medusa. They are 40 to 50 mm. in diameter. From 5 to 20, usually 5 to 8, more
or less distinct marginal lobes. No tentacles. A simple radial-canal in each lobe of the
umbrella. Oral plate quadripartite with 4 oral arms arising from it. Central stomach well
developed, but apparently there was no central mouth-opening.

This form was possibly allied to the Rhizostoma and may have had habits similar to those
of Cassiopea.

Brooksella confusa Walcott.
Brooksella confusa, Wa.cott, 1896, Proc. U. S, Nat. Mus., vol. 18, p. 612, plate 31, figs. 7a-b; 1898, Monographs U. S. Geol.

Sury., vol. 30, p. 30, plate 3.

In Brooksella confusa the marginal lobes do not unite at the center of the bell but join
irregularly, whereas in B. alternata they radiate from the center of the disk. This fossil is
found in the middle Cambrian shale of Coosa Valley, Alabama.

Brooksella rhenana Kinkelin.
Kinkexin, 1903, Bericht Senckenberg Naturf. Gesell., Theil 2, p. 89, taf. 1, fign. 1, 2

An 8-lobed medusa from the middle Devonian of Ruplach. Only one specimen, showing

its exumbrella, was found, and this appears to me to be identical with, or at any rate very
closely related to, Walcett’s Brooksella alternata.

APEENDEX:

PREOCCUPIED GENERIC NAMES.

The establishment of the International Commission upon Zoological Nomenclature and
the general recognition which the code that controls its decisions has won for itself among
naturalists makes it more than ever desirable that the validity of the generic names we now
use should be firmly established. Accordingly, the tenability of each and every generic name
adopted in this work has been made the subject of a thorough search, and I am somewhat
surprised to find that certain names which have been used for generations without question
of their priority are actually preoccupied for other groups of animals and can not be applied
to meduse. Unfortunately I did not carry out this investigation until after volumes I and II
were printed. A list of the generic names which can not be applied to medusz follows:

Corynitis (page 71, Vol. I)=Linvillea nom. nov.

Both Corynitis and Corynetes are preoccupied, the former having been applied to Arach-
nids in 1854, and the latter to Coleoptera by Herbst, 1792. Wagner’s Plotocnide is vaguely
described and figured, but it appears to me to be a Protiara, and Browne’s T 1aricodon, while
it may be a “‘Corynitis,” is too imperfectly known to be determined. We must therefore give
to this genus a new name, and I propose Linvillea in honor of Dr. Henry R. Linville, whe
found the hydroid. The type species is therefore Linvillea agassizt.

Slabberia (page 73, Vol. I) =Dipurena.

Slabberia is preoccupied by Oken, 1815 (Lehrbuch der Naturgesch., Theil 3, Zool., p. 828),
for Slabber’s Medusa marina, which is an Obelia and therefore wholly different from the
medusa to which the name Slabberia was applied by Forbes, 1846. We must therefore drop
Slabberia in the sense in which I have used it and substitute for it the generic name Di purena

McCrady, 1857.
Turris and Tiara=Clavula.

Both Turris and Tiara are preoccupied and can not be used for meduse. The name
Clavula may be applied to these meduse, as has been explained on page 491, Volume II.

Laodicea (see page 201, Vol. I).

According to L. Agassiz, 1842-46, Nomenclator Zoologicus, the generic name Laodicea
was used by Lamouroux, 1816, Hist. Polypiers Coralligenes, and this statement of Agassiz’s
is copied in Scudder’s Universal Index to Genera in Zoology, 1882, p. 167. Upon referring to
Lamoroux’s work, however, I can not discover that he used the name Laodicea, and believe
that Agassiz is mistaken, and that Laodicea of Lesson, 1843, may be retained for meduse.
Laodice is preoccupied, having been used by Gemminger, 1871, for Coleoptera, before Haeckel,
1879, applied it to meduse, but this does not interfere with the use of Laodicea. Indeed,
several medusa genera escape by so narrow a margin; for example, 4malthea takes precedence
over Amalthaea, yet the latter, differing as it does by a single letter, may be used. Similarly
Chrysaor takes precedence over Chrysaora, yet both may be used.

719

20 MEDUS OF THE WORLD.

J

HYDROMEDUS.

Maas, 1909 ( eee Math. phys. Klasse der K. Bayer. Akad. der Wissenschaft, Miin-
chen, Suppl. Bd. 1, Abhandl. 8), gives a description of 23 hydromedusz from Japan obtained
upon Doflein’s ee

The old species are Cytais vulgaris, T1ara papua, Proboscidactyla flavicirrata var. stoloni-
fera, Spirocodon saltatrix, Euchetlota poradasica: Phialidium pacificum, Phialidium discoida,
Mesonema pensile, Gontonemus vertens var. depressum, Olindioides formosa, Liriope rosacea,
Rhopalone ma velatum, Aglaura hemistoma, A2gina rosea, Solmundella bitentaculata, Cuninn
peregrina, and Solmaris rhodoloma.,

The new forms are called Sarsta japonica, Nemopsis dofleint, Turritopsis nutricula var.
pacifica, and Willsta pacifica. A Zanclea and an Obelia are possibly new.

It is interesting that Euchezlota paradoxica, known hitherto only from the Florida-Bahama
region, should be reported from Japan.

One of the most valuable features of Maas’s paper is his redescription, accompanied by an
excellent account of the post-embryonic development of Spirocodon saltatrix. Reviews of his
account of this, and of the new forms, are presented in this Appendix.

An important paper upon Arctic Hydromeduse and Scyphomeduse is that of Hartlaub,
1909, Croisiére Océanographique Belgica dans la Mer du Gronland, Meduses, 18 pp., planches
76-77. Unfortunately this has reached me too late to be reviewed for this work.

Pennaria tiarella (see page 25, Vol. I).
Pennaria tiarella, Harcitr, G. T., 1909, Bull. Mus. Comp. Zool. at Harvard College, vol. 53, p. 164, 5 plates, 44 figs.

Hargitt studies the development of Pennarta tiarella and finds that the odcyte nucleus
dissolves within the germinative vesicle before the nuclear membrane is ruptured. The linine
network of the germinative vesicle extends to the nucleolus, so that an interchange of substances
may possibly occur between the chromatin and the nucleolus.

The two polar bodies appear to be formed at about the time of the liberation of the medusa,
by a process of mitosis of which Hargitt gives a detailed account.

Fertilization usually occurs after the polar bodies have been formed. ‘The cytoplasm is
very active at this time, forming protuberances upon the surface of the egg. The male and
female pronuclei unite by apposition.

The first cleavage spindle seems to form from the cytoplasm. The cytoplasmic division
is delayed, the second nuclear division being completed before the first cleavage furrow has
cut half through the egg.

Hargitt disagrees with Beckwith, 1909, and finds that the polar bodies are seen only at or
near the time of the liberation of the medusz, about 7 p. m. He also finds, contrary to Beck-
with, that the nucleolus disappears within the germinative vesicle before the dissolution of the
nuclear membrane, whereas Beckwith states that it is cast out into the cytoplasm.

Corymorpha nutans (see page 31, Vol. I).
Corymorpha nutans, Mararp, 1907, Bull. Museum Paris, p. 563—Torrey, 1907, Science, vol. 25, p- 734-

Malard finds Corymorpha nutans when dredging in deep water northeast of the Ile Tati-
hou, coast of France, and Torrey observes spontaneous fission in the stems of the hydroid.

Sarsia rosaria (see page 59, Vol. I).

Sarsia rosaria, Kisninouye, 1910, Journ. College of Sci., Tokyo, vol. 27, art. 9, p. 24.

Kishinouye finds this medusa off the Kurile Islands, north of Japan. A. Agassiz records
it from the Pacific coast of North America.

Sarsia japonica Maas.
Sarsia japonica, Maas, 1909, Abhandl. Akad. Wissen., Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 6, taf. 1, fig. 1.
Bell 12 to 15 mm. high, 4 to 6 mm. wide, with flatly rounded apex. Bell walls 2 mm.

thick. Manubrium nearly cylindrical, four-fifths to five-sixths as long as the depth of the bell-
cavity. Gonads scattered irregularly over the manubrium from the base to near the mouth.

APPENDIX. 721
Tentacle-bulbs large pyramidal, without ocelli and without nettling warts. Tentacles tapering,
shorter than bell-height when contracted, their proximal parts with scattered nematocysts,
and distal two-thirds with ring-like nettle-batteries. 4 straight, slender radial-canals without
an axial-canal above the stomach. Gonads and tentacle bulbs yellowish-brown, other parts
colorless. From Todohokke and Hokkaido, Japan. Hydroid unknown. It is probably an
Arctic form.

It is distinguished from S. resplendens and §. brachygaster by haying no ocelli, and from
S. apicula by having no axial-canal and by its bluntly rounded apex. Its nearest relative is
S. fammea, with which, indeed, it may prove to be identical.

Fic. 426.—Corymorpha pendula. After L. Agassiz in Contributions to the
Natural History of the United States, vol. 4.

Eleutheria (see page 93, Vol. I).

Eleutheria, Bape, 1907, Das Seewasser-Aquarium, Magdeburg, 2 fign.—Krumepacn, 1907, Beitrage zur Kenntniss der Medusa
Eleutheria, Breslau, 47 pp-

Krumbach gives a detailed account of Eleutheria. Unfortunately I have not seen his
paper and am unable to review it.

Cladonema radiatum (see page 90, Vol. I).
Cladonema radiatum, Binver, 1908, Bull. Muséum, Paris, p. 385.—Ricuters, 1908, Zool. Anzeiger, Bd. 33, p. 687.

Binder gives a good description of the hydroid from the Atlantic coast of France, and
Richters records it from Helgoland, German Ocean.

22 MEDUS® OF THE WORLD

Urashimea globosa Kishinouye.

Urashimea globosa, Kisuinovure, 1910, Journal College of Sci., Tokyo, vol. 27, art. 9, p. 27, plate 5, figs. 27-29.
Young medusa, Urashimea macrotentaculata, Ibid., p. 28, fig. 30.

Bell 17 mm. high, 15 mm. wide, globular with very thick walls. Numerous meridional
bands of nematocysts arranged more or less definitely in 4 perradial groups. There are 4
interradial, hollow spaces between the exumbrella and the subumbrella, and these have many
pointed processes on the aboral side. Unfortunately Kishinouye’s description is lacking in
detail and his figures only add to my confusion respecting the nature of these spaces. Are
they entirely cut off from the gastrovascular system? ‘The radial-canals have many minute
processes on the aboral side.”

There are 4 tapering tentacles longer than bell-height and with numerous, short, capped
filaments on all sides. Each tentacle with an abaxial ocellus at its base. Mouth with 4 tri-
angular lips. The gonads are 4 pouches hanging down from the horizontal parts of the radial-
canals near the stomach. Each gland is broadest at its “axial extremity” and exhibits two
longitudinal folds. Found at Saghalin and at Monbetsu in Kitami, Hokkaido.

This remarkable medusa is so briefly described and figured that I can not venture to
define its generic characters. Kishinouye states that it is one of the Cladonemide. From
Japan and Saghalin Island.

Urashimea macrotentaculata is apparently a young specimen of the same medusa from
Kuno in Suruga Bay, Japan.

Turritopsis pacifica Maas.
Turritopsis nutricola var. pacifica, Maas, 1909, Abhandl. Akad. Wissen, Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 14, taf. 1, fign.

6-8; taf. 2, fig. 9.

Bell of adult medusa 8 to g mm. high, 5 to 6 mm. wide. This form is distinguished by
the number and arrangement of its tentacles, 120 to 150 of which anise, not in a single row,
as in the Atlantic Turritopsis, but in 3 or even 4 rows, one above the other; the number of
rows increases with age. The tentacles are tapering, their entoderm chordate, and each has
a small, projecting ocellus on the abaxial side of its swollen bulb, whereas the ocelli of the
Atlantic Turritopsis nutricula are on the axial (inner) sides of the tentacles. There are 4
diffuse interradial gonads. The manubnum and radial-canals are as in T. nutricula. Gonads
and stomach orange, ocelli red, entoderm of tentacles greenish. Ten specimens, from Sagami
Bay near Misaki, Japan, in Ociober.

This form differs so markedly from the American medusa that we may safely call it a
distinct species.

Rathkea octopunctata (see pages 175, 177, Vol. I).
Cytais octopunctata, MArKow, 1908, Zool. Anzeiger, Bd. 33, p. 664.

Markow finds that this medusa is very abundant near Sebastopol between February and
April, from the surface to a depth of not more than 7 feet. Budding medusz were abundant
from January 27 to February 12 in water of 6.1° to 6.9° C. The rediscovery of this medusa
in such abundance in the Black Sea makes it practically certain that it was described by
Brandt, 1838, under the name Rathkea blumenbachi, and that this name is merely a synonym
of Rathkea octopunctata, which is the type of the genus. Brandt’s figure shows pinnately-
branched oral tentacles, but this is evidently a mistake.

Rathkea octopunctata var. grata (see page 179, Vol. I).

Lizzia shimiko, Kisuinoure, 1910, Journal College of Sci., Tokyo, vol. 27, art. 9, p. 25, plate 5, fig. 24.

I believe this to be identical with the Arctic variety of R. octopunctata, commonly called
R. grata. Kishinouye found it to be quite common in winter at Misaki, Hamana Inlet, and
in Omura Bay, Japan. He says that the bell is 2 mm. wide and that the 8 basal bulbs are
red and each gives rise to 3 or 4 tentacles. The manubrium and medusa-buds are pinkish.
Kishinouye’s description and figure might equally well have been derived from a study of
R. octopunctata yar. grata, from our Massachusetts coast, the Japanese and American meduse
apparently being identical in all respects.

The medusa is so abundant in Japan that it has received the vulgar name “shimiko.”’

APPENDIX, 723

Nemopsis dofleini Maas.

Nemopsis dofleint, Maas, 1909, Abhandl. Akad. Wissen., Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 11, taf. 1, fign. 4 und 5.
Favonta nipponica, Kisurnovuye, TgTo, Journal College of Sci., Tokyo, vol. 27, art. 9, p. 26, plate 5, fig. 25.
(2) Favonia sulcata, Kisuinovye, Ibid., p. 26, plate 5, fig. 26 (contracted specimen from Korsakoff).

It is probable that Nemopsis dofleint Maas is identical with Favonia nipponica Kishinouye
and that the pair of small median tentacles of each perradial cluster was lost in the 9 preserved
specimens studied by Maas. These are very brittle and are often lost in large specimens of
the American Nemopsis, especially after preservation in formalin. Kishinouye finds that the
marginal tentacles.arise in two rows from the “epaulets,” whereas Maas finds them ina single,
closely crowded row. The appearance of two rows is often due to contraction or to crowd-
ing. Bearing these differences in mind, | present the descriptions of both authors in order

that they may be compared in detail.

Nemopsis dofleini.

Nemopsis “‘nipponica.”

Size in mm.

Shape of bell.

Shape of tentacular epaulet
from which each group of

marginal tentacles arises.

Number of tentacles in each
perradial marginal cluster.

Number of dichotomous
branchings of the oral tenta-
cles.

Size and shape of gonads.

Color.

Where found.

20 to 22 high, 12 to 15 wide.

4-sided, prismatic with flatly rounded apex and
thick walls.

Cleft in the middle. The two halves wing-
shaped.

40 to 60, simple, tapering, set in one row. Ten-
tacles shorter than bell-height. No median
clavate pair of tentacles observed. Ocelli at
the tentacle bases.

5 to 7.

In the form of a double fold along the radial-
canals almost reaching the ring-canal. A per-
radial separation between each half of each
gonad.

Stomach, gonads, and tentacle-bulbs yellowish,
ocelli dark brown.

Bay of Tokyo, Japan.

17 high, 15 wide.

As in N. dofleini.

As in N. dofleini.

About 50 in two rows. A median pair of small
clavate tentacles. Ocelli at tentacle bases.

About Io times.

As in N. dofleini but shorter, being, however,
more than half as long as radial-canals.

Tentacle-bulbs and lips orange, ocelli brown.
Male gonads bluish, female pale yellowish.

Bays of Tokyo and Mikawa, very abundant in
spring.

Willsia pacifica Maas.
Willia pacifica, Maas, 1909, Abhandl. Akad. Wissen., Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 17, taf. 3, fig. 16.

Bell flat, 2 to 3 mm. high, 5 to 7 mm. wide; 6 regularly spaced radial-canals arise from
the stomach and branch so that about 618 terminal canals reach the bell-margin. Manu-
brium flat, 6-sided, with 6 complexly-folded lips. 96 to 108 tentacles as numerous as the
terminal branches of the radial-canals. 96 to 108 short, narrow nettling streaks upon the
exumbrella, alternating with the tentacles. No ocelli. Ring-canal rudimentary. Color (?)
Coast of Japan in September. One specimen.

Polyorchis karafutoensis Kishinouye.
Polyorchis karafutoensis, Kisutnouye, 1910, Journal College of Sci., University Tokyo, vol. 27, art. 9, Pp. 30, plate 5, fig. 31.

This medusa differs from Polyorchis penicillata in its greater size, being 60 mm. high and
50 mm. wide. Also its gonads are dichotomously branched, many of the marginal tentacles
are forked, and the ring-canal gives off centripetal branches.

There are about 40 gonads, 10 on each radial-canal, and these are longer than the manu-
brium, which latter is of the size and shape seen in P. penicillata. There are about 120 marginal

724 MEDUS.® OF THE WORLD.

tentacles, said to arise in several rows from the bell-margin. Each of the 4 radial-canals
gives off from 14 to 16 long, lateral branches which branch at their outer ends but do not
anastomose. Many short, usually unbranched, centripetal canals arise from the ring-canal
and end blindly.

A single specimen was obtained at Korsakoff, Saghalin Island, on September 19, 1906.

In the character of its canal-system this medusa is intermediate between Polyorchis and
Spirocodon, but the bell-margin is simple, not cleft into lappets, and the tentacles are spaced
at equal distances apart around the margin.

Spirocodon saltatrix Tilesius (see page 220, Vol. I).
Spirocodon saltatrix, Maas, 1909, Abhandl. Akad. Wissen., Miinchen, Suppl. Bd. 1, Abhandl. 8, p. 18, taf. 2, fign. 10-13.

Maas gives by far the best published description of this medusa and corrects several
errors of former students, especially in respect to the character of the gonads.

When the bell is only 12 mm. high and 5 mm. wide, with high, slightly bulging sides and
dome-like apex, the gelatinous substance is thick, being thicker in the perradii than in the
8 adradii. The circular muscles of the subumbrella are entire. There are 8 clusters, each
with about 20 tapering tentacles. | The stomach is a long, simple tube with 4 distinct lips

CO TIE LYS Fe ZF

Un

Fic. 427.—Spirocodon saitatrix. After Maas in Abhandl. Akad Wissen., Miinchen.
A, young medusa with small gonads and tentacles, still in 8 clusters. B, half-
grown medusa, showing one of the gonads. C, full-grown medusa, tentacles
and canals omitted to show the form of one of the gonads.

APPENDIX. 725

which are at a level about one-eighth of the height of the bell-cavity above the margin of the
bell. There is a well-developed peduncle above the stomach. The 4 radial-canals each give
off about 20 branched but non-anastomosing side branches in the subumbrella. Over the
peduncle the canals do not branch. There is a ring-canal with 4 short interradial, branched,
blindly ending centripetal canals. In this young stage the gonads are not apparent.

The gonads begin to develop along the 4 radial-canals on the peduncle close to the base
of the stomach when the medusa is about 18 mm. high and 15 mm. wide (see text-fgure
426a). The radial-canals at this point begin to elongate more rapidly than the portion of
the peduncle upon which they lie, and thus they begin to loop outward into the subumbrella
cavity. Finally these freely-projecting canals twist spirally and hang downward into the bell-
cavity, the canal extending around the free edge of a mesentery (figs. 4268 and c). In later
stages the bell-margin develops 8 indentations, 4 perradial and 4 interradial, with 8 adradial
convexities between. The 8 clusters of tentacles of the young medusa spread laterally as new
tentacles develop, until finally the tentacles become congruous entirely around the bell-margin.

The specimens studied by Maas were from the shore at Yokohama and from the Bays
of Tokyo and Sagami, Japan. ‘The largest were 40 mm. high and 30 mm. wide.

According to Maas the 4 interradial vessels are the only blindly ending canals which
arise from the ring-canal, this being contrary to the observations of other authors.

Obelia congdoni Hargitt (see page 248, Vol. II).

Obelia hyalina, ConGpoN, 1907, Proc. American Acad. Arts and Sci., Boston, vol. 42, p. 468, figs. 7-9.
Obelia congdoni, Harcirt, C. W., 1909, Biol. Bulletin, Woods Hole, vol. 17, p- 375-

Hargitt believes this to be specifically distinct from O. hyalina Clarke. The branches
of the stem do not arise in the axils of the hydrothece as in O. hyalina. The gonangia are
larger, being about 4 times the length of the hydrothece, and the opening is not simple, but
there is a neck with everted rim. Moreover, the colony is 20 to 30 mm. high and profusely
branched instead of being about 12 mm. high and but little branched. The newly liberated
medusa has 24 tentacles, but within 10 or 12 hours it has 30 to 36. The hydroid is found
upon drifting Sargassum and is a tropical form.

Staurophora mertensii (see page 291, Vol. II).

Staurophora discoidea, Kisninovye, 1910, Journal College of Sci., Univ. of Tokyo, vol. 27, art. 9, p- 29.

Kishinouye describes this medusa from Japan and Saghalin Island. I believe it to be
identical with S. mertensi:. He states that it closely resembles S. mertensi1, but that there
are about 30 folds on each side of a limb of the gastric cross, instead of 17 as in S. mertensit.
These folds of the genital glands vary greatly in number and increase with age in the Atlantic
Staurophora; they afford therefore an insufficiently definite criterion upon which to base
specific distinctions.

Cubaia gemmifera.

Scolionema gemmifera, KisHinovYe, 1910, Journal College of Sci., Tokyo, vol. 27, art. 9, p- 31, plate 5, figs. 32, 33-

This appears to be an immature Cubaza. The largest of Kishinouye’s specimens was
4 mm. wide and medusiform buds were beginning to develop upon its gonads. The distal
ends of the tentacles beyond the adhesive disks are longer than in any species of this genus
hitherto described. The manubrium is light red with brown mouth. Of the 16 tentacles,
8 were with “suckers” and 8 smaller ones were without them. 8 lithocysts. Entoderm at
base of tentacles greenish and in the distal parts reddish. Found at Misaki, Japan, in winter.

Craspedacusta sowerbii Lankester (see page 363, Vol. II).
Craspedacusta sowerbii, Decisions of International Commission on Zool. Nomenclature, 1910, Science, vol. 31, p. 150.
The International Commission on Zoological Nomenclature publishes its unanimous

decision that the name of this medusa is Craspedacusta sowerbit Lankester, not Limnocodium
victoria Allman.

MEDUS.® OF THE WORLD

~I
bo
oO

Microhydra ryderi (see page 366, Vol. II).
Microhydra ryderi, Goerre, 1908, Mitt. philomath. Gesell., Strassburg, Bd. 4, Jahrg. 16, p. 35, 1 taf.

Goette records the finding of this hydroid in the neighborhood of Strassburg. Hitherto
it has been known only from Tacony Creek near Philadelphia. Unfortunately I have not
been able to see his paper.

Genus Limnocnida (see page 370, Vol. II).
Limnocnida, Gravier, 1908, La Méduse du Tanganyika et du Victoria Nyanza. Sa dispersion en Afrique, Résult. Scientifiques
voyages en Afrique Edouard Foa, Paris, pp. 601-611.

Gravier gives an account of the dispersion of this genus in Central Africa.

Egina citrea (see page 451, Vol. II).
gina pentamera, Kisuinovye, 1910, Journ. College of Sci., Tokyo, vol. 27, art. 9, p. 32, plate 5, fig. 34.
This is a 5-rayed 42gina citrea from Misaki and Suruga, Japan, in winter. Vanhoften,
1908, called attention to the frequent occurrence of 5-rayed aberrations of 4!gina. Kish-
inouye’s medusa is about 20 mm. wide and 10 mm. high, with thick gelatinous bell having a

flat top and sloping sides. The mouth is a simple round opening and the 10 genital sacs
are nearly quadrate. The 5 tentacles are each about twice as long as the bell-radius. Color ( ?)

SCYPHOMEDUS.
Carybdea rastonii (see page 508, Vol. III).
Charybdea mora, Kisuinovye, 1910, Journ. College of Sci., Univ. of Tokyo, vol. 27, art. 9,-p. 6, plate 1, figs. 4-9.

This form from Japan appears to be identical with C. rastoni of the Pacific. It may
possibly be distinguished, however, as a local variety by the large nettling warts over its
exumbrella and its relatively long pedalia, these being about two-fifths as long as the height
of the umbrella. I have, however, seen specimens of C. rastoni: with pedalia one-third as
long as the bell-height.

Carybdea alata (see page 508, Vol. III).
Tamoya virulenta, Kisuinouye, 1910, Journal College of Sci., University Tokyo, vol. 27, art. 9, p. 6, plate 1, figs. 4-9.

This form, from the Inland Sea of Japan, is apparently identical with C. alata. Kish-
inouye describes large specimens 100 mm. high and 60 mm. wide. He finds from 6 to 8
dendritic velar canals in each quadrant, whereas I have not seen more than 6 in specimens of
C. alata. A variation of this sort may be expected, however, in specimens of such great size
as those found by Kishinouye.

Haliclystus octoradiatus (see page 534, Vol. III).
Haliclystus octoradiatus, WietrzyKowsk1, 1909, Comptes Rendus Acad. des Sci., Paris, tome 149, p. 746 (development).

Wietrzykowski gives the best account yet published of the early stages of Haliclystus.
The planula i is about 1164 long, 18 wide. The ectoderm forms a continuous sac of flat,
hexagonal cells, apparently without cilia. There are generally about 16 entodermal cells
arranged in a single row. After 1 to 4 days of free life, the planulz settle down upon
their anterior ends and become hemispherical. They are apt to settle down in clusters and
feed upon Nauplius larvae, which they capture by means of their nematocysts. The mouth
breaks through by the perforation of the ectodermal sac at the summit of the larva. The
larva then gradually becomes vaguely 4-lobed and about 150 in diameter, and a tentacula-
form bud develops at the summit of each of the 4 lobes. These buds become detached and
resemble the original planula, which developed from the egg, and go through developmental
stages similar to those of the mother-larva, fixing themselves by their anterior ends and in
turn giving rise to buds, as did their mother.

At the time of formation of the primary buds, one sees a well-developed i invagination ofa
glandular character at the center of the adherent surface of the larva. This is the beginning
of the pedal zone. The body then elongates, becoming filiform, and then 2 tentacles, 180°
apart and exactly similar in structure to the knobbed tentacles of the adult, develop on opposite

APPENDIX. 127

sides of the mouth. The hypostome then elongates. This stage, with two well-developed
tentacles and the hypostome, persists for several days.

A third tentacle similar to the first two then develops and the three tentacles set them-
selves 120° apart, giving the polyp a triradial symmetry. Finally, a fourth tentacle develops
and the larva has 4 knobbed tentacles go° apart. No later stages were observed at Roscoff.
France, where these studies were undertaken by Wietrzykowski.

Genus Thaumatoscyphus Kishinouye, 1910.

Thaumatoscyphus, Kisutnouye, 1910, Journ. College of Sci., Tokyo, vol. 27, art. 9, p. 2.

The type species is T haumatoscyphus distinctus Kishinouye, from the most northeastern
island of Chishima, Kurile Islands, Japan.

GENERIC CHARACTERS.

Stauromedusz closely allied to Haliclystus and Stenoscyphus, but with a unitary coronal
muscle in the exumbrella; with rudimentary adradial lobes and small, non-adhesive perradial
and interradial tentacles. 4 interradial pits in the subumbrella. Peduncle 4-chambered.
8 adradial gonads. Gastric cavity as in Eleutherocarpide.

This genus is distinguished from all other Stauromedusze by its exumbrella coronal muscle.
This structure is so remarkable, being unknown in any other Scyphomeduse, that its existence
requires confirmation, for contraction in preservation may have produced the furrows which
Kishinouye observes and believes to be the outlines of strands of muscle fibers. He cut no
sections.

Thaumatoscyphus distinctus Kishinouye.

Thaumatoscyphus distinctus, KisHtNouyYe, 1910, Journ. College of Sci., Tokyo, vol. 27, art. 9, p. 2, plate 1, figs. 1 and 2

Body goblet-shaped, 30 mm. high. Calyx 15 mm. wide and half as high as height of
entire Sania 8 short, adradial lobes, each with about 40 short, captate foncales growing in
a lanceolate tract on the aboral side of each lobe. The renraclesi in the proximal part of “the
tentacular tract have very large, swollen stalks and degenerate distal knobs. These swollen
stalks serve as adhesive organs.

The 8 perradial and interradial tentacles are small, cylindrical, and without well-developed
distal knobs, although their ends are captate with a median depression at the tip. These
tentacles bear black pigment at their bases and along the median line. They are not adhesive
organs. The peduncle 1 is more or less quadrate, About as long as the calyx and 4 times as
long as wide. It is 4-chambered.

Four deep, interradial infundibula in the subumbrella. The subumbrella is beset with
large, spherical, wart-like clusters of nematocysts, those near the margin and middle parts
of the mesogonia being the largest, and about 1 mm. in diameter.

The coronal muscle is a broad, undivided band, the greater part of which 1s said to hie
in the exumbrella beyond the clusters of tentacles. 4 broad but weakly developed per-
radial areas of radial muscles extend from the pyloric region through the stomach wall. The
interradial muscles are better developed and extend from the aboral end of the peduncle to
the bell-margin. Each interradial muscle band is divided at its distal end into two short
limbs which extend to the bases of the adradial clusters of tentacles.

The cesophagus is short, somewhat quadrangular, and with deep longitudinal folds.
The 4 lips are folded. The central stomach-cavity is long and prismatic and there are 8
adradial rows of simple, long, gastric cirri. There are 8 adradial lanceolate gonads, each con-
sisting of 7 or 8 oblong follicles. The abaxial surface of each gonad is black and can be seen
through the translucent wall of the body.

Two specimens found in August, 1903, from Shimushiri, Kurile Islands, Japan.

Unfortunately Kishinouye appears to have cut no sections and he bases his statement
of the existence of an exumbrella coronal muscle upon the presence of annular folds in the
external surface of the body-wall. This appearance may well be due to unnatural contraction
in the killing fluid. He studied two preserved specimens. Even if this coronal muscle does not
exist, the medusa may still be called T haumatoscyphus, for it is distinguished from Stenoscyphus
by its adradial lobes, and from Haliclystus by having 4 subgenital pits in its subumbrella.

28 MEDUS OF THE WORLD.

Genus Parumbrosa Kishinouye, roro.
Parumbrosa, Kisninovre, 1910, Journ. College of Sci., Univ. Tokyo, vol. 27, art. 9, p. 19.

The type species is Parumbrosa polylobata Kishinouye from Toyama Bay, Japan.

GENERIC CHARACTERS.

Ulmaride similar to the genus Discomedusa, but with 64 marginal lappets instead of 32.
This genus is evidently derived from Discomedusa by the bifurcation of its marginal lappets.

Parumbrosa polylobata Kishinouye.
Parumbrosa polylobata, Kisninouye, 1910, Journ. College of Sci., Univ. Tokyo, vol. 27, art. 9, p. 19, plate 4, figs. 20-23.

Bell 160 mm. in diameter, flat, about 4 times as wide as high. Gelatinous substance
of delicate consistency. Exumbrella finely and uniformly graduated. 64 narrow, lanceolate,
pointed marginal lappets. 6 velar lobes between every 2 divergent ocular lobes. The velar
lobes are 3 times as long as wide, but the ocular lobes are only about half as long as the velar
and about twice as long as wide. Each pair of ocular lobes is, however, mounted upon a
common basal projection which causes them to project beyond the contour of the velar lobes.

There are 24 tentacles and 8 sense-organs
arranged so that 2 marginal lappets are placed
between a tentacle and a sense-organ or between
two successive tentacles. The adradial tentacles
are the longest. There are powerful muscle fibers
on the axial side and transverse bands of nemato-
cysts on the abaxial side of each tentacle.

The subumbrella is nearly smooth with weakly
developed muscles. The canal-system is as in
Discomedusa philippina (see page 607) except that
the perradial and interradial canals are less com-
plex in their branching and there is but a single,
blindly ending side branch from the ring-canal in
each lappet, instead of two as in D. philtppina.
D. philippina may, indeed, be only the young of
P. polylobata, and later the 32 lappets may divide to form 64. The large size of the gonads
and complex branching of the interradial and perradial canals in D. philippina, however,
cause me to hesitate before drawing this conclusion. The bluntly rounded lappets of D.
philippina are also very different in shape from the long pointed ones of P. polylobata.
In any event P. polylobata was evidently derived philogenetically if not ontogenetically
from some such medusa as D. philippina.

The cesophagus of P. polylobata is about as long as the bell-radius, is 4-sided and pris-
matic, and the richly folded, lanceolate lips are as long as the mouth-tube. They are thick
and keeled along the midrib, and their margins bear numerous minute filaments. The 4 long,
narrow gonads are about 5 times as wide as the perradial spaces between them. The medusa
is colorless and nearly transparent. It was found in large numbers in a haul of a shrimp-trawl
in Toyama Bay, Japan, in June, 1907, from a depth of about 65 fathoms.

Kishinouye gives excellent figures of the medusa.

Fic. 428.—Parumbrosa polylobata. After Kishinouye, in
Journal College of Science University of Tokyo.

FOSSIL MEDUS&.
Ephyropsites jurassicus von Ammon.
Ephyropsites jurassicus, YON AMMON, 1908, Geonostischen Jahresheft, 1906, Jahrg. 19, p. 169, taf. 3, 2 fign.

This is one of the Coronate closely allied to Nausithoé. It is from the Upper Jurassic
limestone of Pfalzpaint. Bell 150 mm. wide with a distinct annular furrow and a pedal zone
40 mm. wide. 8 tentacles and 8 rhopalia. 16 pedalia in the radii of the tentacles and sense-
organs. The tentacular pedalia are twice as wide at the zone of the tentacles as the rhopalar
pedalia. A median ridge upon each pedalium, and near the margin on the thopalar pedalia
a pair of short, radiating ridges, each of which gives off a divergent cross-furrow arising from
the inner end of each radiating ridge. ‘There are ring muscles in the subumbrella. A single
impression of this medusa was studied by von Ammon.

InpeEx To Vouume III.

Acalepha deperdita, 715
Acraspeda, 499
Acraspedites antiquus, 710
Aigina citrea, 720
Agassiz, Alexander, acknowledgment to, 499
Agelacrinus lindstromi, 717
Appendix, 719
Arcadomyaria, 634
Archirhiza, 712
aurosa, 712
primordialis, 712
Archirihizida, 633
Astylospongia radiata, 715
Atolla, 561
achillis, 566
alexandri, 566
batrdit, 563
forma valdivie, 565
' chunt, 566
gigantea, 505
verrillii, 567
wyuillet, 566
forma verrillit, 567
Atollida, 561
Atollites minor, 710
zittelt, 710
Atorella, 567
subglobosa, 508
vanhoffent, 508
Atorellide, 541, 567
Aurelia, 619
aurita, 623
clausa, 628
ceerulea, 623
colpota, 623
cruciata, 623
dubia, 623, 627
flavidula, 623
habanensis, 623
japonica, 623
labiata, 628
limbata, 628
maldivensis, 629
marginalis, 627
sex-ovalis, 623
solida, 627
vitiana, 623
Aurelina, 618
Aurellia, 619
aurtta, 623
forma marginalts, 627
dubia, 627
flavidula, 623
labiata, 628

maldivensts, 629

Aurellia solida, 627
water-content of, 622
Auricoma, 619
Aur sa 630
furcata, 650

Bathotrephis asteroides, 717

Bathyluca solaris, 585

Biblis, 619

Brachiolophus, 709
collaris, 711

Brooksella alternata, 718
confusa, 718
rhenana, 718

Budding in Cyanea, 599; Tzniolhydra roscoffen-
sis, 622; Cassiopea, 643; Cotylorhiza, 662;
Haliclystus, 726

Calicinaria cyathiformis, 524
Callinema, 612
ornata, 616
Cannorhiza, 713
connexa, 714
Cannostomites multictrrata, 710
Capria, 521, 539
sturdzit, 539
Caryhdea, 505, 500
reactions to light, 509
alata, 508, 510, 720
var. grandis, 511
var. mosert, 512
var. pyramis, 511
aurtfera, 510
haplonema, 513
marsuptalts, 507
murrayana, 512
rastont1, 508, 726
xaymacana, 509
Carybdetde, 501, 504
Carduella, 523
Casstopea, 636
andromeda, 037
var. acyclobbia, 640
var. malayensis, 039
var. maldivensis, 639
var. zanzibarica, 639
anglica, 703
borbonica, 659
borlasea, 703
canariensis, 659
corallifora, 659
depressa, 049
var. picta, 649
dieuphila, 636
forskalca, 637
729

730

Cassitopea frondosa, 641, 647
lunulata, 703
mertensit, O49
ndrosta, 050
ornata, 648

var. digitala, 648
pallasit, 647
picta, 649
polypotdes, 640
rhizostomoidea, 703
theophila, 636
xamachana, 641

Cassiopeidz, 633

Cassiopeja acycloblia, 640
andromeda var. cyclobalia, 640
mertensii var. ndrosia, 650

Catostylide, 633

Catostylus, 664
cruciatus, 667
mosatcus, 666
ornatellus, 670
orsint, 669
palmi pes, 667
purpurus, 671
stiphropterus, 670
stuhlmannt, 669
tagt, 068
tripterus, O71
turgescens, 671
viridescens, 670
wilkesu, 666

Cephea, 651
aldrovandi, 699
capensis, 703
cephea, 654

var. conifera, 655

var. dumokuroa, 656
conifera, 654, 655
cyclophora, 652
dumokuroa, 656
forskalea, 654
fusca, 654
mosaica, 666
ocellata, 680
octostyla, 652

var. cerulescens, 653
papua, 678
papuensis, 678
polychroma, 659
rhizostoma, 710
thizostomoidea, 654
tuberculata, 659
typhlodendrium, 658
vesiculosa, 663
wagneri, 659

Cepheidz, 663, 650

Charybdea, 506
arborifera, 508
hyacinthina,”544
mora, 726
moseri, 512

INDEX.

Charybdea obeliscus, 511
philippina, 512
Chaunostomidz, 633, 650
Chemical composition of Cyanea, 600; Aurellia,
626
Chirodropus, 505, 518
gorilla, 518
palmatus, 519
Chiropsalmus, 505, 515
buttendijkt, 515
quadrigatus, 516
quadrumanus, 515
zygonema, 517
Chrysaora, 577
aspilonota, 579
blossevillei, 581
calliparea, 582
chinensis, 582
convoluta, 581
cyclonata, 579
(dodecabostrycha) dubia, 546
fulgida, 581
gaudichaudii, 593
gilberti, 582
helvola, 581
var. calliparea, 582
var. chinensis, 582
heptanema, 579
hysoscella, 579
var. blossevillet, 581
var. fulgida, 581
blossevillet var. plocamia, 581
isosceles, 579
lactea, 583
lesueuril, 579
macrogona, 579
mediterranea, 579
melanaster, 582
var. gilberti, 582
pleurophora, 579
plocamia, 581
(polybostrycha) helvola, 581
spilhemigona, 579
spilogona, 579
Cladonema radiatum, 721
Claustra, 619
Cletstocarpida, 519
Coloring matter, chemical composition of: in
Pelagia, 572; Cyanea, 600; Cassiopea, 637
Collaspida, 541, 561
Collaspis, 561
achillis, 566
Commensal plant cells in Scyphomedusaw: Cas-
siopea, 637, 643, 646; Cotylorhiza, 661;
Catostylus, 665, 667
Commensalism between Scyphomedusz and fishes:
Sanderia, 591; Cyanea, 609; Cotylorhiza,
663; Catostylus, 667 wins
Commensalism between Scyphomeduse and
shrimp: Cephea, 658
Coronata, 541

INDEX,

Coronate, 501, 541
Corymorpha nutans, 720
Corynitis = Linvillea, 719
Cotylorhiza, 658
ambulacrata, 663
borbonica, 659
tuberculata, 659
Couthouya, 591
gaudichaudi, 593
Couthouyia, 591
pendula, 593
Crambessa, 664.
cruciata, 667
mosaica, 666
palmipes, 667
pictonum, 668
stiphroptera, 670
stuhImanni, 669
tagi, 668
triptera, 6071
viridescens, 670
Crambione, 676
cooki1, 677
mastigophora, 676
Cramborhiza, 672
flagellata, 673
Craspedacusta sowerbit, 725
Craspedotella, 499
Craterlophus, 521, 538
macrocystis, 538
tethys, 538
Crossostoma, 685
anadyomene, 686
Crystals in ectoderm of Nausithoé, 555
Cubaia gemmifera, 725
Cubomeduse, 504
Cyanea, 595
annaskala, 601
arctica, 591, 596, 597
behringiana, 596
calliparea, 582
capillata, 596
var. fulva, 600
var. nozaki1, OOT
var. versicolor, 600
citrea, 597
ferruginea, 596, 597
fulva, 596, 597, 600
imporcata, 597
lamarcki, 596, 597
muellerianthe, 601
nozakii, 601
postelsii, 596, 597
purpura, 601
rosea, 601
Cyanea versicolor, 596, 597, 600
Cyanetda, 501
Cyclomyaria, 634
Cyst-formation in: Cyanea, 599; Taeniolhydra
roscoffensis, 622

“J
co
pear

Dactylometra, 583
africana, 588
ferruginaster, 588
lactea, 583
longrcirra, 589
pacifica var. ferruginaster, 588
quinquecirrha, 585
var. pacifica, 589
Dactyloidites asteroides, 717
bulbosus, 717
Degenerate medusx: Stauromedusz, 520; Aurellia,
622; Taeniolhydra, 622
Depastrella, 523
Depastrum, 521, 523
cyathi forme, 524
tnabat, 525
Desmonema, 591, 595
annasethe, 601
chierchiana, 593
gaudichaudi, 593
rosea, 601
Desmostoma, 677
gracile, 681
Development of: Tripedalia, 514; Lucernaria,
526; Haliclystus, 534, 726; Nausithoé,
5553 Linuche, 559; Pelagia, 573; 574,
Chrysaora, 577, 581; Dactylometra, 586;
Cyanea, 599; Diplulmaris, 611; Aurellia, 622,
625; Rhizostome, 633; Cassiopea, 643;
Cotylorhiza, 661; Mastigias, 679; Phyllo-
thiza, 684; Rhizostoma, 701; Stomolophus,
710
Dianaea cyanella, 574
Diplocraspedon, 619
limbata, 628
Diplopilus couthouyi, 654
Diplulmarts, 609
antarctica, O10
Discomedusa, 606
lobata, 607
philip pina, 607
Discomedusx, 541, 569
Donacostoma, 604.
woodii, 604
Drymonema, 603
cordelio, 603
dalmatina, 603

gorgo, 604.

Edible meduse: Chiropsalmus, 517; Rhizostoma,
703; Rhopilema, 706

Eleutheria, 721

Eleutherocarpide, 519

Environment, effect of, upon: Dactylometra, 587;
Cyanea, 599, 600; Darkness upon Cassiopea,
643

Ephyra, 551
prometor, 551

Ephyridz, 550

Ephyroides rotaformis, 561

Ephyropside, 541, 550

732

Ephyropsis, 553
Ephyropsites jurassicus, 728
Eucrambessa, 677
milleri, 679
Eulithota fasciculata, 716
Eupilema, 709
scapulare, 709
Evagora, 619
Excretory system of Atolla, 561

Favonia sulcata, 723
nipponica, 723
Floresca, 605
pallada, 605
parthenta, 605
Floscula, 605
pandora, 606
promethea, 605
Flosculida, 604
Fosstl Medusa, 715

Grafting in Cassiopea, 646

INDEX.

Kishinouyea, 521, 531
nagatensis, 531

Laodicea, 719
Laotira cambria, 717
Leonura, 696
leptura, 696°
terminalis, 696
Leptobrachia, 696
leptopus, 606
lorifera, 694
Leptobrachidw, 633, 691
Limnoenida, 726
Limnocodium, 725
Linerges, 557
aquila, 560
draco, 560
mercurius, 558
pegasus, 558
Linergide, 550
Liniscus, 557
cyamopterus, 558
ornithopterus, 558
sandalopterus, 558

Habits of Meduse: Creeping habits of the planula Linuche, 557

of Lucernaria, 500, 526; egg-laying and aquila, 560
swimming habits of Linuche, 559; feeding uriguiculata 558
habits of Chrysaora, 581; of Cyanea, 600; of vesiculata, 558

Cassiopea 637, 647, 648; Cotylorhiza, 661

Heccaedecomma, 612
ambiguum, 615
Haltclystus, 521, 531
antarcticus, 530
auricula, 532
kerguelensts, 536
octoradtatus, 534, 726
salpinx, 535
Steynegert, 535
tenuis, 532
Halicyathus, 536
lagena, 537
Halimocyathus, 521, 536
lagena, 537
platypus, 537
Halipetasus, 651
scaber, 652
Haplorhiza, 713
punctata, 713
simplex, 713

Linvillea, 719
Lipkea, 521, 539
ruspoltana, 540
Lizzia shimiko, 722
Lobocrocis blossevillei, 581
Lobonema, 631, 688
smith, 689
Loborhiza, 664
ornatella, 670
Lorifera, 693
arabica, 694
flagellata, 695
lorifera, 604
var. pacifica, 695
Lucernarta, 521, 526
auricula, 532, 534, 537
australis, 530
bathyphila, 530
campanulata, 530
conyolutus, 530
cyathiformis, 524

Hermaphroditism of: Chrysaora, 581; Pseudo- fabricii, 537

rhiza, 683
Hidroticus rufus, 680
Himantostoma, 693

flagellata, 695

lorifera, 694

var. pacifica, 695

Holigocladodes lunulatus, 703
Homopnensis frondosa, 663

Kuragea, 589
depressa, 589

Lucernarta haeckelt, 529
infundibulum, 529
kiikenthalt, 529
octoradiata, 534
platypus, 537
pyramidalis (?)*, 528
quadricornis, 527
salpinx, §35
typica, 537
waltert, 520

Lucernarida, 519

Lucernosa, 526
bathyphila, 530
haeckeli, 529
kukenthali, 529
walteri, 529

Lychnorhiza, 672
bartscht, 674.
flagellata, 673
lucerna, 673

Lychnorhizidz, 633

Manania, 536
auricula, 537
Marsupialida, 504
Masttgias, 677
gracile, 681
ocellata, 6SO
orsini, 669
pantherina, 681
papua, 678, 681
var. physophora, 678
rosea, O81
papua var. stboge, 679, 680
var. siderea, 679
physophora, 678
Mastygias, 677
mulleri, 679
papua, 678
siderea, 679
Medora, 591, 595
capensis, 593
Medusa, 619
andromeda, 637
aurita, 623
bleu, 703
capillata, 596
cephea, 652, 654
corona, 703
frondosa, 647
hysoscella, 579
marsupialis, 507
noctiluca, 572
octopus, 703
octostyla, 652, 654
panopyra, 575
pelagica, 574
pulmo, 699
stelligera, 607
tuber, 659
tuberculata, 659
unguiculata, 558
Medusina, 717
costata, 619, 717
deperdita, 715
geryonides, 717
princeps, 715
radiata, 715
Medusites antiquus, 716
costatus, 717
deperditus, 715
favosus, 715

INDEX.

Medusites lindstr6mi, 717
princeps, 715
radiatus, 715

Melanaster, 577
mertensil, 582

Melusina, 604
formosa, 597

Microhydra rydert, 726

Microstylus, 651

Monocraspedon, 619

Monorhiza, 682
haeckelii, 683

Myogramma speciosum, 716

Nauphanta, 553
albatrossi, 555, 557
challenger, 556
polaris, 554
vettoris pisani, 554

Nauphantopsis, 548
diomedea, 548

Nausicaa, 553
pheacum, 556

Nausithoé, 553
albatrosst1, 557
albida, 554
challengert, 556
clausi, 556
marginata, 554
picta, 557
punctata, 554

rubra, 557
Nectophilema, 704

verrillii, 707
Nemopsis doflein1, 723
Neopelagia eximia, 590

733

Nervous system of: Lucernaria, 526; Rhizostoma,

702

Netrostoma, 651
ccerulescens, 653
dumokuroa, 656
typhlodendrium, 658

Obelia congdoni, 725
Octogonia, 553
Ocyroé, 619
Orithyia lutea, 703
Orythia incolor, 663

Palephyra, 551

antiqua, 551

indica, 553

pelagtca, 552

primigenia, 551
Paraphyllina, 549

intermedia, 549
Paraphyllinide, 541, 548
Paraphyllites distinctus, 549, 715
Parumbrosa, 605, 728

polylobata, 728

734

Patera, 604
cerebriformis, 604
Pelagia, 570
crassa, 576
var. sublaevis, 576
cyanella, 574, 576
denticulata, 576
discoidea, 576
flaveola, 576
neglecta, 574
noctiluca, 572
var, neglecta, 574
panopyra, 575
var. placenta, 575
papillata, 576
perla, 576
phosphora, 576
placenta, 575
quinquecirrha, 585
tahitiana, 576
unguiculata, 558
Pelagida, 569
Pelagothuria, 499
Pennaria tiarella, 720
Pertcolpa, 541
campana, 542
quadrigata, 542
tetralina, 542
Pericrypta, 541
campana, 542
Periphema, 543
Peromedusz, 541
Pertphylla, 543
dodecalostrycha, 549
humilis, 546
hyacinthina, 544

forma dodecalostrycha, 546

forma regina, 546
mirabilis, 546
Perirhiza, 561
nematophora, 654
Pertphyllide, 541
Periphyllopsis, 547
brauert, 546
Phacellophora, 612
ambigua, 613, 615
camtschatica, 613
ornata, 616
stcula, 613
Phanerocarpx, 499
Philogeny of Rhizostoma, 663
Phyllorhiza, 684.
punctata, 684
Pilema, 698
capense, 703
clavigera, 706
corona, 703
octopus, 703
pulmo, 699
stylonectes, 703

INDEX.

Polyclonia, 636, 647
frondosa, 647
theophila, 636
Polyorchis karafutoensts, 723
Polyrhiza, 663
cephea, 654
vestculosa, 663
Poralta, 617
rufescens, 017
Potta marina, 699
Preoccupied generic names, 719
‘ Procharagma prototypus, 508
Procharybdts, 505, 506
tetraptera, 500
Procyanea, 595
Protolyellia princeps, 715
Pseudoclytia pentata, 619
Pseudorhiza, 682
aurosa, 682
haeckelt1, 683
thocambaui, 678
Pulsation in: Scyphomedusz, 503; Aurellia, 620;
Cassiopea, 637, 645; Rhizostoma, 701;
Rhisoztomz, 632; Cotylorhiza, 661

Radiomyaria, 634, 650
Rathkea octopunctata, 722
var. grata, 722
Regeneration in: Stauromedusaw, 522; Haliclys-
tus, 532; Craterlophus, 538; Cassiopea, 637,
646; Rhizostoma, 632; Rhizostoma, 702
Rhacopilus, 664
cruciatus, 667
cyanolobatus, 667
Rhizostoma, 698
aldrovandi, 699
brachyra, 692
capensis, 703
coeruleum, 703
corona, 703
cuvierl, 699, 703
fulgidum, 581
hispidum, 706
leptopus, 696
lorifera, 694
lutea, 703
mosaica, 666
pulmo, 699
var. capensis, 703
var. corona, 703
var. lutea, 703
var. octopus, 703
rosea, 681
sepioides, 703
trigona, 663
Rhizostoma, 501, 631
relationships of, 500
Rhizostomata, 631
dichotdma, 634, 650
lorifera, 635, 691
pinnata, 634, 635

INDEX.

Rhizostomata scapulata, 635, 697
simplicia, 635, 712
triptera, 634, 663

Rhizostomidz, 631, 633, 697

Rhizostomites admtrandus, 718

Rhopilema, 704
esculenta, 704
hispidum, 706
rhopalophora, 704
verrillit, 707
verrucosa, 706

Sanderta, 590
malayensts, 590
Sarsia japonica, 720
rosarta, 720
Scyphomedusie: development of, 499, 501; geo-
graphical distribution, 501; reactions to
light, 509; relationship to Hydromeduse and
to Actiniana, 502
Semzostomata, 569
Semeaostomee, 501, 569
Semaostomites zittelt, 716
Slabberia=D1purena, 719
Semostome, 569
S patangopsts costata, 717
Sptrocodon saltatrix, 724
Spongtcola fistularts, 553, 554
Stauromedusa, 501, 519
Staurophora mertensi1, 725
Steganopthalme, 499
Stenoptycha, 595
Stenoscyphida, 525
Stenoscyphus, 521, 524
hexaradtatus, 525
tnabat, 525
mirabilis, 553, 554
Sthenonta, 611
albida, 611
Sthenonine, O11
Stings caused by medusz, 690
Stomolophidz, 633, 697
Stomolophus, 709
agaricus, 710
chunii, 710
meleagris, 710
var. fritillarta, 711
Stomatonema reticulatum, 714
Stylorhiza, 651
octostyla, 652
polystyla, 652
Swarms of Meduse: Carybdea, 511; Linuche,
559, 560; Cyanea, 600; Cassiopea, 643;
Cephea, 657; Catostylus, 665, 667; Stomo-
lophus, 711; Haliclystus, 726

Taeniolhydra roscoffensis, 622
T amoya, 505, 512
haplonema, 513
prismatica, 513
punctata, 509

Tamoya quadrumanus, 515
virulenta, 726
T essera, 522
princeps, 522
typus, 523
Tesserantha, 522
connectens, 523
Tesseraria, 521, 522
scyphomeda, 523
T haumatoscyphus, 727
distinctus, 727
T ysanostoma, 691
denscrispum, 692
thysanura, 692
Toreuma, 635
dteuphila, 636
gegenbauri, 636
thysanostoma, 636
Toxoclytus, 664
roseus, 681
tripterus, 671
turgescens, 671
Tripedalta, 505, 513
cystophora, 514
Turris and Tiara=Clavula, 719
Turritopsts nutricola var. pactfica, 722

pacifica, 722

Ulmaride, 604
Ulmaris, 606, 608
prototypus, 607, 609
Ulmaropsis, 609
antarctica, 610
drygalskii, 610
Umbrosa, 606
lobata, 607
Umbrosina, 605
Undosa, 608
undulata, 609
Urashimea globosa, 722
macrotentaculata, 722
Urtica marina, 579
Urtica marina—octopedalis, 703

Variations in Meduse: Lucernaria, 530; Hali-
clystus, 534; Craterlophus, 538; Dactylo-
metra, 583, 584; Sanderia, 591; Aurellia,
619, 626; Cassiopea, 640, 641, 643

Versura, 685
anad yomene, 686
maast, 687
palmata, 685
pinnata, 686

vesicata, 086
Willsia pacifica, 723

Zonephyra, 551
corona, 571
pelagica, 552
zonaria, 552

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