REPORT UPON THE SCYPHOMEDUSAE COLLECTED BYTHE UNITED STATES BUREAU OF FISHERIESSTEAMER "ALBATROSS" IN THE PHILIPPINE IS-LANDS AND MALAY ARCHIPELAGO.Br Alfred Goldsborough Mayer,Director of the Department of Marine Biology of the Carnegie Institution) ofWashington.INTRODUCTION.In 1909 Dr. Hugh M. Smith, the then Acting United States Com-missioner of Fisheries (now Commissioner), gave to the author forstudy a part of the collection of medusae now under consideration inthis article, and the authorities of the Smithsonian Institution kindlypermitted the use of a table in the building in Washington whileengaged upon this work. In 1910 the remainder of the collection wassent to Princeton University, where the author occupied a researchroom in Guyot Hall. Thus the collection was studied under the mostadvantageous conditions, and it is a pleasure to express my sense ofindebtedness to the authorities of the United States Bureau of Fish-eries, the Smithsonian Institution, and Princeton University for theopportunities I have enjoyed.Previous reports upon this collection were published by the Car-negie Institution of Washington in Publications 109 and 212.Thirty-one Scyphomedusae were obtained by the Albatross in thePhilippines and adjacent regions, of which 10 were new to science,the list of species being as follows:CARYBEIDAE. RHIZOSTOMATA.Carybdea rastonii Haacke. Cassiopea andromeda, var. baduensisCarybdea alata, var. grandis Agassiz Mayer.and Mayer. Cephea octostyla (Forskal).Chiropsalmus quadrigatus Haeckel. Cephea cephea, var. coerulea Van-hoffen.GORONATAE. Cotylorhiza paciflca Mayer.Catostylus purpurus Mayer.Periphylla hyacinthina Steenstrup. Catostylus toumsendi Mayer.Lmuche unguiculata, var. aquila Catostylus mosaicus (Quoy and Gai-Mayer. mard).Atolla bairdii forma wyvillei Haeckel. Lychnorhiza bartschi Mayer.Atolla bairdii forma gigantea Maas. Lychnorhiza bomensis Mayer.Atolla bairdii forma valdiviae Van- Mastigias papua (Lesson),hoffen. Mastigias occellata (Modeer).175
176 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.SEMAEOSTOMAE. RHINOSTOMATA?Continued.Pelagia panopyra Peron and Lesueur. Phyllorhiza luzoni Mayer.Chrysaora melanaster Brandt. Versura maasi Mayer.Dactylometra africana Vanhoffen. Lobonema smithii Mayer.Sanderia malayensis Goette. TJiysanostoma thysanura Haeckel.Discomedusa philippina Mayer. Lorifera lorifera, var. pacificaAurellia aurita (Linnaeus). (Schultze).Aurellia labiata Chamisso and Eysen-hardt.Also in 1914 S. F. Light x describes 7 species from the Philippinesother than those obtained by the Albatross. Mr. Light's additionalspecies are : Dactylometra quinquecirrha; Cassiopea polypoides; Cas-siopea medusa, new species; Acromitus maculosus, new species;Lobonema mayeri, new species; Lobonemoides gracilis, new species;and Rhopilema visayana, new species.Thus 38 Scyphomedusae are already known from the Philippines,which is thus one of the richest regions of the world for Scypho-medusae, and stands in marked contrast with the tropical Pacificcoast of Queensland, Australia, of which only 10 species have beendescribed.This appears to be another illustration of the influence of a greatocean current, the rich region of the Philippines being in the sweepof the Japan Stream, whereas there is no well-defined current alongthe southern shore of Papua or off the Barrier Reef of Queensland.As is well known, H. B. Bigelow, in his report upon the Siphono-phorae of the Albatross, shows how abundant these forms are in theHumboldt current off the coast of South America and how poor theregion is in the mid-Pacific to the westward of this great current.My studies, made while assistant upon Dr. Alexander Agassiz's expe-ditions to the tropical Pacific, in 1899, as well as upon the CarnegieInstitution of Washington expedition of 1913 to Torres Straits andPapua, show that the whole great belt of the South Tropical Pacific,from the western edge of the Humboldt current to the shores ofAustralia, is poor in pelagic life. A number of local medusae appearin some of the large island groups, as in Fiji, but the region as awhole is poor in forms peculiar to itself, and even those of wide dis-tribution are, generally speaking, found only occasionally over thisgreat desert of ocean.STRUCTURE, PHYSIOLOGY, HABITS, AND DEVELOPMENT OF THESCYPHOMEDUSAE.The Scyphomedusae are the large jellyfishes, commonly called thesea blubbers, in which the body is umbrella shaped, the mouth parts
1 Philippine Journal of Science, vol. 9, Section D, No. 3.
SCYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS." 177occupying the position of the handle of the umbrella. The animalswims by means of periodic expansions and contractions of the mar-gin of the umbrella. If we look carefully, we will see that thismargin is notched at regular intervals, forming a series of lappets,and that 8, or sometimes 16, of these notches are deeper than theothers and contain each a minute finger-shaped or club-shaped senseorgan which may be provided with an eye, but which always con-tains a mass of crystals or concretions concentrated at its outer end.These little sense organs are so small that they appear to the nakedeye as mere pigmented specks set within the niches at regular inter-vals around the margin; but though small they are of vital impor-tance to the jellyfish, constituting its principal nerve centers; and ifwe cut them off, the animal commonly becomes paralyzed and is nolonger able to pulsate spontaneously. Mayer believes that the crys-tals or concretions within the sense clubs consist largely of calciumuric oxalate, and this makes it appear probable that sodium oxalateis constantly forming in the sense club and that the calcium chlorideof the sea water, when it enters the sense clubs, is precipitated, form-ing calcium oxalate, and in this manner setting free sodium chloride(common salt), which is a powerful stimulant for the nerves, thuscausing the pulsating reaction.Thus in these animals it is found that a stimulus which is con-stantly present causes periodic contractions followed by periods ofrest which are perhaps due to fatigue. In the case of the scypho-medusa Cassiopea this stimulus is internal, for the sea water itselfneither stimulates nor inhibits the sense organs of the animal, thestimulating effect of the interaction of its sodium, potassium, andcalcium being offset by the inhibiting effect of its magnesium uponthe motor centers.But the marginal sense organs do more than merely produce thepulsation stimulus, for Dr. L. K. Cary finds that if they be removedand the medusa be then wounded in any manner the first stages inregeneration are delayed, but if even a single sense organ be presentregeneration proceeds at once and with normal rapidity.In general, if the area of tissue enervated by a sense organ belarge it pulsates more rapidly than if the area be small, and althoughthe ratio is not strictty proportional to the area of tissue, for accord-ing to Dr. L. R. Cary (1917), the rate is reduced one-half when thearea is reduced to one-sixteenth. But young, small jellyfishes pul-sate more rapidly than large, old ones ; yet if we graft two jellyfishestogether the small active one will force the large one to pulsate atits own rate, which will be even more rapid than the normal rate ofthe small one, due to the large area of tissue the sense organs nowcontrol.101825??Bull. 100?17 12
178 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.Eimer and Romanes found that if the marginal sense organs beremoved the jellyfish is paralyzed and responds only by single con-tractions to external stimuli. Later, in 1906, Mayer found that ifthe sense organs be removed and we cut a ring-shaped or circuit-shaped strip of tissue from the concave part of the bell, we maythen start a contraction wave proceeding in one direction through thecircuit through which it travels continuously, being, indeed, en-trapped by the circuit of tissue from which it can not escape. Thismovement is almost machinelike in its regularity, its rate being about440 cm. a second, and in 1917 McClenden showed that this noteremained practically unchanged even though the nerves were arti-ficially stretched.It is interesting to see that the pulsation stimulus in jellyfishes isconducted by the nerves, whereas in the vertebrate heart it is con-ducted by the muscles. There is, however, as Parker showed, afundamental likeness between nervous and muscular activity, for inmost essential features, such as the compensating pause followingan extra pulsation and the refractory stage during systole, latentperiod, reaction to temperature, etc., the jellyfishes behave as doesthe vertebrate heart. In Europe Romanes, Bethe, and von Uexkull,.and in America Loeb, Harvey, Cary, McClenden, and Mayer havebeen most active in these studies.Recently Mayer finds that nerve conduction in Cassiopea is achemical reaction in which the cations of sodium, calcium, and po-tassium take the active part, while magnesium is nonessential. Thusif we dilute the sea water with 0.4 molecular magnesium chloridethe rate of pulsation declines only very slightly more than if wediluted it with distilled water, thus demonstrating the inert natureof magnesium in respect to the rate of nerve conduction. It is alsointeresting to see that this decline in rate is proportional to thedecline in the concentration of the cations of sodium, calcium, andpotassium which surround the nerves and not to the electrical con-ductivity of the sea water as a whole. Thus if we dilute with dis-tilled water we decrease the electrical conductivity in nearly thesame ratio as the dilution, while if we dilute with 0.4 molecularmagnesium chloride the electrical conductivity remains nearly con-stant while we simply reduce the concentration of the sodium, cal-cium, and potassium cations. It is therefore these cations whichare alone essential to the maintenance of the rate of nerve conduction.The previous work of Mayer, 1906, Meltzer and Auer, 1908, andespecially Osterhout, 1916, make it appear that sodium and calciumtogether combine with some proteid element forming a sodium-calcium-ion proteid, and this compound takes an essential part innerve conduction. Mayer in 1916 showed that the rate of nerve con-duction in the medusa Cassiopea has a temperature coefficient 2.5 times
SOYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS." 179
as great as that of the electrical conductivity of the sea water, andthis suggests that the sodium-calcium-ion proteid may have a hightemperature coefficient of ionization ; for if the rate of nerve conduc-tion is proportional to the concentration of the reacting cations, thiswould account for the high temperature coefficient of the rate ofnerve conduction.Also the sodium-calcium-ion proteid is probably colloidal in char-acter and being in an alkaline medium it is doubtless negative elec-trically. Thus, under the influence of the decided negative potentialwhich accompanies the reaction of nerve conduction, the surfacetension on the particles of this colloid must be augmented and thismay reduce the size of the particles in the manner stated by Mayer,Schaeffer, and Terroine, 1907.Granted this were the case, the solubility of the particles would beaugmented; for the surface decreases only as the square, while thevolume declines as the cube of the radius. Thus more ions of sodiumand calcium would pass into solution in a given time from the manysmall than from the few large particles ; the mass remaining constant.Nerve conduction is therefore probably a very complex chemicalreaction, in which also an enzyme action may possibly be involved aspostulated by Harvey, 1911.Goldfarb showed that Cassiopea regenerates more rapidly in 90per cent of sea water (90 parts of sea water mixed with 10 parts ofdistilled water) than it does in normal sea water. Nerve conditionis, however, most rapid in slightly concentrated sea water.In all Scyphomedusae, excepting the Ehizostomae, tentacles arefound at the bell-margin, and usually grow out from between thenotches of the rim. These tentacles are hollow, the stomach cavitybeing continued into them as it is also into the sense-clubs; indeed,the sense organs are only highly modified tentacles.Powerfully developed circular, or a combination of circular andradial, muscles are found in the under, concave part of the umbrella(the subumbrella), and here also there is a network of nerve fibersconnecting the muscles with the sense clubs. Curiously, there are nomuscles, and probably no nerves, over the outer convex part of theumbrella (the exumbrella).Those who have handled jellyfishes know that they are capable ofinflicting a sharp sting, the tentacles being especially active in thisrespect. Closely clustered over the surface of the tentacles and otherparts of the jellyfish there are minute cells, each containing a hollowtube coiled rope-like within the cell. Upon excitation these littletubes are turned inside out and shot forward, and being more orless barbed they penertate the skin, causing a sharp sting due in part,it appears, to formic acid. Thus it is that these large jellyfishes are
180 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.among the most persistent enemies of the fishes, for many an in-cautious victim is ensnared among their deadly tentacles only to beparalyzed and finally drawn upward into the mouth of the jellyfish.It is, however, a poor rule which does not work in both directions,and certain kinds of small fishes often accompany jellyfishes, swim-ming in and out among the dangerous tentacles, even biting off smallpieces of the jellyfish itself and occasionally themselves falling aprey to the stings, but in general enjoying a peculiar protection fromthe attacks of larger fishes who dare not venture too near the jelly-fish.The mouth, or mouths, of the jellyfish may be surrounded by veil-like lips or, if the mouths are numerous, as in the so-called Rizo-stomae (root-mouthed) jellyfishes, by complex frills lined by minutetentacles which at intervals bend to and fro and sweep, as it were,for food, for the jellyfishes are all carnivorous. In the higheranimals the intestine is a tube which lies suspended within the bodycavity, but Huxley showed that the jellyfishes have no body cavity,and consist simply of a stomach and an outside with a merestructureless lamella, or a solid mass of jell}7 , between the stomachwall and the outer skin layer of the animal.This gelatinous substance, which according to S. Hatai may beallied in composition to chitin or cartilage, may serve as a store offood for the animal in case of starvation, and Oassiopea can live atleast 42 days without food, the weight of the jellyfish decliningto less than one-hundredth of its original magnitude. The loss ofweight after the first day of starvation, and thus after all undigestedfood has been discharged through the mouths of the jellyfish, followsa simple law which shows that the loss of weight each day is propor-tional to the weight of the animal at the beginning of that day andthus the lighter it becomes the less weight lost.Thus, if W be the original weight of the jellyfish and y its weightafter x days of starving, then y=W {l?a) x \ where a is a constant,less than unity. This shows that the source of energy during starva-tion is chiefly the very large and simply organized gelatinous sub-stance which, being very voluminous and heavy, is far more impor-tant than the mere thin layer of cellular elements possessed by thejellyfish. For, as is well known, vertebrates in starving first con-sume glycogen, then mainly fats and lypoids, and finally proteid ele-ments. Due, however, to the large volume of its gelatinous sub-stance, the jellyfish chiefly consumes this during, starvation, although,as Doctor Hatai showed, the starvation is in other respects simi-lar to that of vertebrates. However, jellyfishes have on hand a con-stantly ready source of sustenance, and can, so to speak, feed upontheir own bodies if deprived of prey. Mayer, 1914, carried out some
SCYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS. ' ' 181work upon this subject, but the more recent study of S. Hatai is farmore elaborate and convincing, and should be consulted by all stu-dents of the subject.Indeed the rate of growth, and the ultimate size that an individualjellyfish attains before becoming mature, is a measure of its successin obtaining food, and it is interesting to see that the largest jelly-fishes are those of the cold seas where the floating animal life ismore abundant than in the Tropics. In common with the Corals,Sea-anemones, Alcyonaria, Siphonophores, and Hydromedusae theScyphomedusae are, so far as is known, exclusively carnivorous anddo not feed upon plant life. Thus it is possible that the more rapidgrowth rate of Pacific corals in comparison with those of the At-lantic may be due to the better food supply derived from the deeplagoons of the Pacific, whereas in the Atlantic the lagoons are shal-low and the water is charged over these vast flats with a precipitateof calcium carbonate which collects upon all floating animals andtends to smother them ; whereas in the Pacific this does not appear tobe the case.Jellyfishes are, then, all carnivorous, and, while few devour othersof their own species, they often greedily feed upon other sorts ofmedusae. These animals are, indeed, an important factor in destroy-ing the eggs of cod and other fishes whose spawn floats in the sea.The prey is seized by the mouth, and after being held and par-tially digested in the stomach, the remnant is ejected through themouth.The central stomach is a space in the middle of the umbrella, butthis always gives rise to an outwardly radiating system of pouches ortubes which may form a complex network of vessels under the mus-cular layer of the concave side of the umbrella. As this system ofpouches is connected with the stomach, and nutrient fluids derivedfrom the food circulate through it, it is often called the gastrovas-cular system, for it is both a sort of " chymiferous system " as well asa digestive and circulatory space.In all the larger jellyfishes, or Scyphomedusae, we find withinthe stomach four clusters of tentacle-shaped organs of unknownfunction, placed at the broad sides of the cruciform mouth. Thesmaller jellyfishes, or Hydromedusae, lack these stomach-tentacles orgastric cirri, as they are often called ; and in still another structuraldetail do they differ from the Scyphomedusae, for the Hydromedusaehave a diaphragm-like membrane (velum) extending inward fromthe bell-margin and partially closing the opening of the umbrella,but the larger jellyfishes (Scyphomedusae) do not have a diaphragmof this sort, although it is true that the Carybeidae, or sea-wasps,appear to have such a diaphragm, but it is not strictly comparablewith that of the Hydromedusae.
182 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.The sexual organs of the larger jellyfishes (Scyphomedusae) areentodermal and are found in the stomach, peripheral to, and closelyassociated with, the four clusters of gastric cirri. The sexes areusually separate, the animals being either male or female, although,in rare instances, as in Chrysaora, they are hermaphroditic, or malewhen young and female when old. When mature at the breedingseason the males and females usually come to the surface in greatnumbers and may congregate in vast swarms many square miles inarea. The larvae or eggs may then be cast out into the water by thebreaking down of the stomach wall, or the larvae may undergo apart of their development within the stomach, or mouth parts, ofthe mother, finally to be cast out through the mouth, or set freefrom the disintegrating bodies of the dying parents.In any event the larvae soon develop into minute pear-shapedcreatures about as large as a pin's head, their bodies being coveredwith vibrating cilia, which enable them to spin and progress throughthe water. For a few days, or even weeks, they may remain thusswimming near the surface and may be drifted hundreds of miles bytide and ocean current. Soon, however, the little pear-shaped planula,as it is called, settles down head first upon the bottom and fastensitself to some fixed object. Then for the first time the mouth devel-ops at that which was the posterior end of the planula, and tentaclesgrow out so that the mouth is soon surrounded by 16 or more of theseorgans which serve to capture the minute Crustacea and other organ-isms upon which the little polyp feeds. Thus it remains sedentaryfor a long period, growing all the time and superficially resembling asmall sea-anemone. Finally a series of constrictions develop at reg-ular intervals around the sides, and the creature appears as if it werecomposed of a series of disks set one upon the other. The margin ofeach disk soon develops eight cleft lobes, and eight sense-clubs appearin the clefts. Then the uppermost disk, containing the mouth andthe crown of tentacles, is cast off and perishes, while the others areset free in succession and swim away as minute jellyfishes, soon todevelop tentacles and finally to become mature and repeat this pe-culiar process of development. After the last disk has been cast off,only the stump of the strobila, as it is called, remains, but this mayregenerate a new ring of tentacles and continue to grow, and possiblyto develop more jellyfishes at the succeeding season.There are many interesting variations of this typical process ofdevelopment. Often the strobila, instead of giving off a series ofdisks, develops only a single constriction, and every alternate tentaclechanges into a sense-club, while the other tentacles may be whollyabsorbed, so that they disappear. In this case only a single ephyraor larval jellyfish is set free. This form of development is especiallycharacteristic of the Rhizostomae or multi-mouthed jellyfishes, such
SCYPHOMEDUSAE COLLECTED BY STEAMER "ALBATROSS." 183
as Cassiopea. In the free-floating Pelagia, however, the planulalarva never becomes attached, but remains swimming through thewater until it develops directly into a jellyfish. Thus it is that thesejellyfishes are quite independent of the land and are widely distrib-uted over the tropical and warm oceans; but this is exceptional, formost of the Scyphomedusae must spend their early days attached tosome fixed object and usually in relatively shallow water near somecoast.Some of these coastal medusae are, however, widely distributedover the world, one of these being the large semi-transparent Aurel-lia aurita of our own bays and harbors, which appears so commonlyduring the summer, and may be recognized by its four horseshoe-shaped, milky or pink-colored genital organs. This form occursfrom pole to pole.Such adaptability to wide range of temperature is very rare amongjellyfishes, and is known only in Aurellia aurita and according toVanhoffen in Nausithoe puncata among the scyphomedusae, and Sol-mundella among hydromedusae ; these forms occurring in seas of alltemperatures.Nevertheless, even tropical medusae are much more injuriouslyaffected by a slight rise in temperature than are the jellyfishes of thetemperate regions, and we may say that most tropical forms livewithin 12? C. of their heat-death-temperature, and even tropical formscan withstand cooling better than they can resist heat. To use an en-gineering expression we might say that the medusae of temperateregions have a larger " factor of safety " in respect to temperature.Harvey showed that, upon heating, the rate of conduction of thenervous stimulus which causes pulsation increases in an arithmeticalratio, so that its " curve " is a straight line. At from 34? to 39?,however, the curve makes a sudden bend downward and the rate de-clines sharply. This decline may, in part, be due to the formationof carbon dioxide (C02 ) in the tissues, for Winterstein showed thatthe rate of oxygen consumption in jellyfishes is 3^ times more rapidat 30? to 35? than it is at about 12? C. Indeed, experiments madeby the author in 1917 support the idea that high temperature causesacid to accumulate in the tissues and this causes death throughacidosis.The large, rich rosin-brown colored cyaneas of our New Englandcoast are not found in the Tropics, but closely allied species reappearin the South Temperate Zone, so that somehow they have managed,perhaps in the glacial epoch, to cross the warm zone of the Tropics,or they may have succeeded in crossing the Equator in the cold, deep,underlying drift that moves toward the warm regions over the seabottom from both the northern and southern polar seas.
184 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.The following table shows the range of the 174 species of Scypho-medusae, not counting varieties, described in Mayer's Medusae of theWorld.Confined to the Tropics 77Ranging from Tropics to Temperate regions 16Ranging from Tropical to Polar regions 2Confined to the North Temperate region 32Confined to the South Temperate region 11Ranging from Temperate to Polar seas 6Confined to the Arctic Ocean 7Confined to the Antarctic Ocean 7Deep sea medusae all of wide range 16Total 174In this table the Mediterranean is designated " North Temperate "and the Red Sea " Tropical." In the " Tropics " the surface tempera-ture in summer is above 70? F. (21? C). The summer temperatureof " Temperate Region " rangesfrom 70? F. to 50? (21? to 18?C), and that of the " Polar Seas "below 50? F. (10? C).
'i'/i^I r \ \ \ ^Uk) As is shown, 44 per cent of themi jMlOll'il^Wn^^ known Scyphomedusae appear tobe confined to tropical waters andonly two species, Aurellia auritaby the author. Naples Zoological and possibly Nausithoe punctataSTATI0N - (fig. 1) range from Tropical toPolar Seas. Indeed most of the scyphomedusae are confined withinrather narrow temperature ranges, the only considerable number ofwidely ranging forms being the deep sea species whose habitat hasabout the same temperature whether under the Equator or in thePolar regions. Incidentally the table seems to indicate our deficiencyof knowledge of the South Temperate regions for it is probable thatthe numbers of species in the North Temperate are more nearly equalto those found in the South Temperate Zone.We may distinguish five main divisions or orders of the largerjelly fishes of scyphomedusae.1. The Carybdeidae or Cubomedusae, with bell almost rectangularand with flexible tentacles mounted upon the ends of gelatinous wing-shaped expansions. They have four knob-shaped sense-clubs, whichare very large and set within niches on the sides of the bell, with theeyes all directed inward, apparently to command a view of the mouth.These medusae have a marginal diaphragm which partially closesthe opening to the umbrella cavity. The Carybdeidae are confinedto tropical and warm seas and usually remain upon or near the bot-
SCYPHOMEDUSAE COLLECTED BY STEAMER "ALBATBOSS. ' ' 185torn until they become mature, when they often swim to the surfaceto cast out their eggs and sperm. They are dull milky-yellow, orlivid in color, and their tentacles sting so sharply that they are com-monly called " sea-wasps." None are known north of Cape Cod,but they are well represented in the Tropics throughout the world.2. The Stauromedusae are attached forms which do not pulsate.The body is pear-shaped or pyramidal and the jellyfish fastens itselfto seaweed or rocks by means of a stalk at the aboral end of the body.There are usually 8 clusters of knobbed tentacles which are developedat the pointed ends of eight marginal lobes, and 8 large " anchors "or suckerlike clubs upon the margin may alternate with the tentacles.These forms are confined to cold and temperate seas, and are knownfrom the Arctic and Antarctic Oceans, although they have neverbeen found in the Tropics.3. The Coronatae are usually deep-sea forms distinguished by theirpeculiar dark purple or reddish-brown color, which is very charac-teristic of the invertebrate animals of the deep sea, due possibly tothe fact that the red rays from the sun are rapidly absorbed as theypenetrate into the ocean, so that at depths greater that 1,000 fathomsthere is practically no red light, and here red-colored animals mustappear black and be well concealed in the general darkness of theirabysmal realm. The Coronatae are characterized by having a deepringlike furrow cutting into the thick gelatinous wall of the sides ofthe umbrella; and peripheral to this are gelatinous thickeningsin the radii of the tentacles and sense-organs. The gelatinous wallof the bell is much reduced in thickness at the ring-furrow, the thinpart acting as a hinge to permit the creature to close its bell duringcontraction. These forms are common in the deep waters of thePhilippines, as indeed they are upon the bottom of all seas 500fathoms or more in depth. Most of the species, such as Atolla andPeriphylla, do not normally come to the surface, but a closely alliedform, the little Nausithoe, is one of the most universally present sur-face forms of all warm seas, while the mature Linuche occur in vastnumbers, all rapidly pulsating and resembling little brown thimbles,the swarm often covering square miles of ocean in tropical regions.4. The Semaeostomata are the common large jellyfishes of ourcoasts, such as Aurellia, Cyanea, and Dactylometra. They have ten-tacles and a single cruciform mouth provided with veillike lips,and there is no ring-furrow cutting into the surface of the umbrella,such as is seen in the Coronatae. They are common in bays andbrackish estuaries: and are the largest and most conspicuous of alljellyfishes in temperate regions.5. The Rhizostomae are the large jellyfishes of the East Indianand tropical seas, where they are commonly as conspicuous as are the
186 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.Semaeostomeae in our cold northern waters. They have numerousmouths and no marginal tentacles, and their gelatinous substance isoften as rigid as newly-formed cartilage. Often they occur inswarms in the harbors, swimming strongly against the tide, and aftera storm great numbers are found stranded upon the shore. The rigidgelatinous substance of these jellyfishes is very characteristic and wasseen in fossil forms found in the Jurassic lithographic slates of Stein-heim in Bavaria, showing that in the age of the Reptiles these mosthighly differentiated jellyfishes existed. Indeed, so old are the jelly-fishes that their relationship to the echinoderms, sponges, and cteno-phores remains unknown; all intergrading forms, if such ever ex-isted, having died out long ago, as is often the case in very ancientorders.In an early stage of development the higher animals commonlypass through a condition in which they have only an outer cell layerand a cavity lined by cells destined to form the intestine. Theo-retically speaking, they are simply little 2-layered sacks, the outerlayer being the external skin with its nervous and sensory organs,and the inner layer being the stomach ; and thus the name gastrulais applied to this stage. Jellyfishes are essentially in the gastrulastage, even when adult. Yet so extraordinary are the foldings, out-growths, and adaptations that have arisen in their two body layersduring the vast time they have existed upon the earth that, ulti-mately, simple as they are, no class of the animal kingdom exhibitsa more surprising variety of forms than do the jellyfishes and theirclose allies the Siphonophorae.It is interesting to observe that the large jellyfishes, Scyphome-dusae, which have gastric cirri and no marginal diaphragm or velum,are probably only very remotely related to the small jellyfishes, thehydromedusae, which have a velum and lack gastric cirri. Indeed,we have good reason to believe that the jellyfish-shape and peculiarlocomotion through pulsation have been derived independently in thetwo groups. The Scyphomedusae are probably allied to the actiniansor sea-anemones, while the hydromedusae have probably been derivedfrom hydroids. In fact a jellyfish shape and pulsating body have beenacquired independently in widely different kinds of animals, such asPelagothuria, a holothurian which bears a wonderfully close resem-blance to a jellyfish and swims actively through the water in thetropical Pacific; and in Craspedotella, a minute unicellular marineanimal, which would certainly have been mistaken for a jellyfish hadit not been of microsopic size.Indeed, there is reason to lead us to believe that the bell of theNarcomedusae is a mere outgrowth from the sides of the pyriformlarva, and has thus been acquired in a manner quite different from
SCYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS." 187that of the other hydromedusae. Thus the umbrellalike bodies ofjellyfishes have probably been acquired in at least three differentways within the group itself. Although nearly one-half of the knownforms of Scyphomedusae are confined to the Tropics, yet many ofthese are rare, although, curiously, when found they usually appearin swarms. Thus one may explore the Florida Eeef for 20 years andnot find a single Pelagia or Stomolophus, yet if an individual befound there are almost certainly dozens or even thousands in theneighborhood. It is as if they had all remained floating side by sidethroughout their lives, or at least throughout the period when theycome to the surface from some well-defined region of the bottomwherein they have spent their early days.Apart from the harbors and semibrackish estuaries one rarely findsin the Tropics great swarms of medusae of a few species such ascharacterize the coastal waters of the temperate regions.There are, however, exceptions to this rule, such as the enormousnumber of Linuche, which when mature rise suddenly and simul-taneously to the surface to cast out their genital products and thento sink and die. For areas of square miles the tropical ocean is be-sprinkled with these little brown thimbles, darting in a rapid jerkingmovement.Similarly the pale milky Cubomedusae with their long pink tenta-cles rise, when mature, from the depths to congregate along tropicalshores for the few days or weeks of the breeding season.Metschnikoff observed that many medusae cast out their eggs onlyat certain definite times of the day or night. One of his Mediter-ranean species, for example, laid its eggs always at about 3.30 o'clockin the afternoon, and Conklin observed that Linuche casts its eggsonly at about 8 o'clock in the morning, but other Scyphomedusae arenot so regular, and indeed may retain the developing young in theirmouth folds for days or weeks before they escape into the ocean.DESCRIPTION OF GENERA AND SPECIES.Genus CARYBDEA Peron and Lesueur, 1809.Carybdea Peron and Lesueur, 1809, Ann. Mus. Hist. Nat., Paris, vol. 14, p.332.?Mayer, 1910, Medusae of the World, vol. 3, p. 506.Generic Characters.?Carybdeidae, with 4 simple, interradial ten-tacles and pedalia. Velarium supported by 4 bracket-like frenulae.Velar canals present. Stomach small and 4-sided.CARYBDEA RASTONII Haacke.Carybdea rastonii Haacke, 1887, Jena, Zeitsch. fur Naturwissen, vol. 20,p. 591, pi. 35, figs. 1-15.?Mayer, 1910, Medusae of the World, vol. 3,p. 508.
188 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.There are two specimens of this medusa in the Albatross collec-tion from the Philippines. They are apparently mature and theirdimensions in millimeters are as follows
:
Subig Bay,Luzon, sur-face, Jan. 6,1908. Cat.No. 27937,U.S.N.H.
Height of bell I 34Width of bell 20Length of pedalia j 11Width of pedalia at baseWidth of pedalia at widest part ' 6. 5Height of sensory niche above velar margin j 6Length of flexible shafts of tentacles ' 30
Taal Anchor-age, BalayanBay, Luzon,surface, Feb.20, 1909.Cat. No.28714,U.S.N. M.
3323.59.545.5666Two other specimens (Cat. No. 27935, U.S.N.M.) were taken bythe Albatross at Nasugbu, Luzon, January 15, 1908, and still another(Cat. No. 27936, U.S.N.M.) at Mansalay,Mindoro.This is the commonest Cubomedusa ofthe tropical Pacific, being widely distrib-uted from South Australia to the Ha-waiian Islands. It can be recognized byits prismatic, 4-sided bell, the 4 pedaliabeing about one-fourth to one-third as longas the bell-height, flat and spatula-shapedand about three-fifths as wide as long. Thesense-clubs have each 2 large median eyesand 4 small lateral ocelli. There are 4short, branched, non-anastomosing velarcanals in each quadrant of the velarium,16 in all. Four very small branched tuftsof gastric cirri. This medusa is veryclosely allied to Garybdea marsupialis ofthe Mediterranean, but is distinguished byhaving only 16 instead of 24 to 30 velar canals, and by its somewhatmore slender pedalia. In both forms the bell is dull milky yellowand the flexible parts of the tentacles are pink. In common withother Cubomedusae it comes to the surface when mature and is thenabundant in harbors. The young usually remain in deep water ator near the bottom.
Fig. 2.
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Carybdea rastoniifrom the HawaiianIslands. Two views ofits sense-clubs and aquadrant of the velarium,showing the branchedvelar canals.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 189CARYBDEA ALATA, var. GRANDIS Agassiz and Mayer.Charybdea grandis Agassiz and Mayer, 1902, Mem. Museum Comp. Zool.at Harvard College, vol. 26, p. 153, pi. 6, figs. 26-31.Carybdea alata, var. grandis Mayeb, 1910, Medusae of the World, vol. 3,p. 511, fig. 329.The bell of this medusa becomes 230mm. high, but the largest specimen ofthis cruise was obtained by the Alba-tross in Borneo and is only 166 mm.high. This tropical Pacific form maybe distinguished by its short, wide-flaring pedalia and by having only 1or 2 median eyes upon each sense-cluband no lateral eyes. When young, how-ever, there are 2 large median and 2small lateral eyes, but the latter appearto fuse later with the median eyes.There are 24 velar canals which areshort, branched, and non-anastomosing.C. moseri Mayer is only a half-grownstage of this medusa. In C. alata thereare 6 eyes in each sense-club, and thepedalia are longer and narrower thanin the large variety grandis.The dimensions and characters of thetwo largest of the three specimensfound by the Albatross are as follows:
Fig. 3.? Carybdea alata, half-grown MEDUSA FROM THE HA-WAIIAN Islands. Two views ofITS SENSE - CLUBS AND A QUAD-RANT OF THE VELARIUM, SHOWINGTHE STRAIGHT SIMPLE VELARCANALS.
Station 5361, Feb. 9, 1909,Manila Bay, Luzon, 12 D. 5594, Sept. 30. 1909,fathoms, 2 specimens. off Mount Putri,Cat. No. 28713 U.S.N.M. Borneo.
Height of bellWidth of bellLength of pedaliaWidth of pedalia at widest partWidth of pedalia at baseHeight of sensory niche above velar marginNumber of eyes in each sense-clubGonads
120 ! 166.88 ' 144.38 along inner side i 77 along outer side.
I 36 along inner side.16.5 ! 29.1416 31.2 median, no lateral eyes 2 median, no lateraleyes.Small, immature Small.
190 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.Genus CHIROPSALMUS L. Agassiz, 1862.Chiropsahnus L. Agassiz, 1862, Contr. Nat. Hist. U. S., vol. 4, p. 174.
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Mayeb, 1910, Medusae of the World, vol. 3, p. 515.Generic characters.?Carybdeidae with four interradial, branch-ing pedalia which give rise to a number of tentacles. Four wide per-radially situated stomach pockets in the subumbrella, each of whichgives rise to finger-shaped,unbranched, hernialikepouches, which projectinto the bell cavity. Wide,marginal pouches and nu-merous canals in the ve-larium. Eight leaf-shapedgonads.CHIROPSALMUS QUADRIGATUSHaeckel.Chiropsahnus quadrigatusHaeckel, 18S0, Syst. derMedusen, p. 447.
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Mayeb,1910, Medusae of theWorld, vol. 3, p. 516, fig.331.?Light, 1914, Philip-pine Journ. Science, vol.9, p. 197.The 44 specimens of thismedusa were collected bythe Albatross from thePhilippines, but not oneseems to be mature.Light (1914) found thismedusa at Culion Bay, Cu-lion, and at Palawan, andthus it ranges widely in the Philippines. Light's specimens were largerthan those found by the Albatross, being at least 200 mm. in diameterand usually with seven tentacles to each padalium. The tentaclesare 1.5 meters or more in length and have lavander-colored bands ofnematocysts. Light reports that the sting of this medusa is verysevere and may even be fatal to man. The Dactylometra of thePhilippines is also a dangerous form, but Lobonema, which some ofthe members of the Albatross expedition believed to be virulent, isnot capable of inflicting a very severe sting.1
"IG. 4. CHIROPSALMUS QUADRIGATUS FROM THRPhilippine Islands. A, B, Views of sense-club. O, View looking down on the cruci-form mouth. D, Gastric cirri. E, BranchedPEDALIUM WITH SEVERED TENTACLES.
1 Light, 1914, Philippine Journal of Science, vol. 9, pp. 291-295.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 191The following is a record of specimens of Ghiropsalmus quadri-gatus obtained by the Albatross in the Philippine Islands
:
Locality.
192 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.into 4 pairs of rhopalar, and 4 pairs of tentacular lappets. A deepannular furrow separates the dome-like apex of the exumbrellafrom marginal zone of bell. Between this ring-furrow and thelappets is a zone of 16 pedalia, 12 in the tentacular and 4 in therhopalar radii, and these are separated one from another by 16deep, radiating clefts which extend down the mid-axial lines ofthe lappets. There are four deep, interradial subgenital pits in thefloor of the subumbrella, lined above their edges by rows of internalgastric cirri. The large central stomach extends peripherally out-ward into the subumbrellain the four perradii. Thesefour openings lead into awide ring-sinus in the sub-umbrella, which in turnsends out a radiating vesselin the radius of each tenta-cle and rhopalium, 16 in all.These vessels fork beforereaching the tentacles orrhopalia, and their diverg-ing ends curve around theedges of the lappets andform a marginal ring-canal.I believe that Periphyllahyacinthina can not be sep-Fig. 5.
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Periphylla hyacinthina from the arated specifically from P.Hawaiian Islands, showing range in slope J, O de C ab OStry cha. TheOF THE BELL. *shape of the bell is quitevariable, and when large the medusa usually becomes relativelyflat and domelike, whereas it is relatively high and conical whenyoung. P. hyacinthina is said to be densely pigmented with pur-ple-brown so that the gonads can not be seen through the bellwalls, whereas P. dodecabostrycha is said to be less densely coloredand semitranslucent. This distinction does not always apply, andcertainly the degree of pigmentation appears to be quite independ-ent of the shape of the bell, whether flat and domelike or high andpointed. Browne (1910), in his study of the Scyphomedusae of theNational Antarctic Expedition, concludes that P. dodecabostrychais probably only a large-growth phase of P. hyacinthina, and withthis opinion I am heartily in accord.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 193PERIPHYLLA HYACINTHINA Steenstrup.Periphylla hyacinthina Steenstbup, 1837, Act. Mus. Hafniensis.
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Mayer,1910, Medusae of the World, vol. 3, p. 543, figs. 342 and 343.Periphylla hyacinthina forma dodecabostrycha Mayeb, 1910, Medusae ofthe World, vol. 3, p. 546.The Albatross obtained 21 specimens of this common deep-seamedusa among the Philippine Islands between depths of 338 and1,291 fathoms, as follows:
Station.
D 5201 and D 5203, Cat. No. 28704,U.S.N.M., off Limasaua Island.
Date.
1908.Apr. 10
1909.Mar. 2D 5373, near Marinduque Island,about 15 miles off shore. Cat.No. 28705, U.S.N.M.D 5379, about 37 miles off Mompog Mar. 4Island, near Marinduque Island. ICat. No. 28706, U.S.N.M.D 5471, from about 15 miles off j June 19Point, Light, eastcoast of Luzon.Cat. No. 2873S, U.S.N.M.D 5486, from 6 miles off Batobobo July 31Point, Panaon Island.D 5497, about 10 miles off Bantigui Aug. 3Island.
Depth.
D 5507, off northern Mindanao.Cat. No. 2S703, U.S.N.M.D 5628, Pariente Strait, about 7miles from St. Lamo Island.Cat. No. 2S707, U.S.N.M.D 5647, Buton StraitD 5652, Gulf of Boni
Aug. 5Nov. 30Dec 16Dec. 17
Fathoms.554-775
338
920
568
585960
4251,291
519525
Character ofbottom.
Gray sand andmud andgreen mud.Soft sand
Green mudand finesand.
....doGray mudGreen mud.
Size andnumber ofspecimens.
2 large, 190mm. wide,105 high.2 large
4 medium
.
7 large andmedium.1
2 medium.1 medium.
.do.
.do.
Variety.
Dodecabostrycha.
Do.
Do.
Hyacinthina.
Dodecabostrycha.Do.
Do.Hyacinthina.
Do.Do.
Genus LINUCHE Eschscholtz, 1829.Linuche Eschschultz, 1829, Syst. der Acalephen, p. 91.
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Mayeb, 1910,Medusae of the World, vol. 3, p. 557.Generic characters.?Coronatae with 8 rhopalia, 4 perridial, and4 interradial. Eight tentacles, 16 marginal lappets; with hernia-likesacs of the gastrovascular cavity protruding from the floor of thesubumbrella. Eight gonads grouped in four pairs close to the fourperradii. The central stomach opens by four perradial ostia into aring-sinus, which in turn breaks up into 16 branching, radiatingpouches in the lappets. A marginal ring canal is present in thePacific L. aquila, and according to Vanhoffen also in L. unguiculataof the Atlantic, but I believe it is not always present in the Atlanticform, for I am unable to demonstrate it by inflation of the poucheswith air in the living or recently preserved specimens.101825??Bull. 100?17 13
194 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.
'a O
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LINUCHE UNGUICULATA, var. AQUILA Mayer.Linerges aquila Haeckel, 1880, Syst. der Medusen, p. 496.Linuche aquila Mayek, 1910, Medusae of the World, vol. 3, p. 560, figs. 356B and C.Linuche unguiculata Vanhoffen, 1913, Zoologischen Jahrbiichern, Suppl.11, Heft 3, p. 429.Linuche ungiculata forma aquila Mayer, 1915, Publication No. 212, CarnegieInstitution of Washington, p. 174, fig. 1, B and O.This variety is widely distributed over the tropical Pacific and isclosely related to the tropical Atlantic L. unguiculata, with which itis identical in form and dimensions, being about 13 mm. high and 16mm. wide. It has 48 wartlike protuberances upon the subumbrellaarranged in two rows instead of in three, as is commonly the casein the Atlantic medusa. EightOn ?of the subumbrella sacs in thevariety aquila alternate withthe gonads and eight arisefrom the sides of the gonadsthemselves. Thus, in the Pa-cific medusa we have two zonesof protuberances? an innerzone of 16 large sacs and anouter of 32 small subumbrellasaccules. The 16 large sacs liein the midregions of the go-nads, while the 32 small sac-cules lie at the zone of theouter ends of the gonads. Inthe Atlantic form the areas ofbrown cells are commonly de-veloped only centrifugal to the zone of gonad, while in the Pacificvariety they occur between the gonads as well as beyond them. Amarginal ring canal is present. The Pacific variety is, however,found also in the Atlantic.Vast swarms of these Medusae are found among the Samoan, Fiji,and Paumotos Islands, and they extend to the coasts of Africa andto Queensland, Australia. They abound in Queensland in the springmonths, in Fiji in December, and at Singapore in April. I havestudied a large collection of these medusae taken in the PhilippineIslands at Mactan, near Cebu, on April 6, 1908, Cat. No. 27944,U.S.N.M., by the United States Bureau of Fisheries steamer Alba-tross. All were mature.Vanhoffen (1913) reported that he succeeded in demonstratingthat a marginal ring canal is present in the Atlantic L. unguiculata,although after many tests I have been unable to detect its presence in
BFIG. 6.?LlNDCHB UNGUICULATA. A, THEUSUAL ARRANGEMENT OP THE SUBUMBRELLAWARTS IN THE TROPICAL ATLANTIC FORM.B, The usual arrangement of the sub-umbrella WARTS IN THE PACIFIC FORM.G, Enlarged view of lips, gonads, andSUBUMBRELLA SACCULES IN THE PACIFICFORM FROM THE PHILIPPINES.
SCYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS.' ' 195living or freshly preserved animals, and am inclined to believe that thedelicate membrane separating adjacent pouches was broken in Van-hoffen's specimens, which had been preserved for a long time informalin. Moreover. Vanhoffen found that in some of the Atlanticmedusae from the Bahama-Florida region the subumbrella warts arearranged as in L. aquila of the Pacific. It thus appears that thePacific form is at best only a variety of the Atlantic species, and bothshould be called Linuche unguiculata.Prof. E. G. Conklin in 1908 1 described the swarming habits, struc-ture of the egg, and the segmentation in this medusa. The eggs arelaid at 8 o'clock in the morning, after which the medusa sinks andsoon dies. The segmentation is nearly equal and synchronous. Theperipheral layer of ooplasm of the egg becomes the peripheral layerof the gastrula and gives rise to the cilia of the ectoderm. The inter-mediate layer gives rise to the principal part of all the cells of thegastrula, while the central part of the egg is the precursor of thecleavage cavity and serves as a kind of a fluid yolk for the nourish-ment of the surrounding cells.Genus ATOLLA Haeckel, 1880, sensu Fewkes.Atolla Haeckel, 1880, Syst. der Medusen, p. 488.
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Fewkes, 1886, ReportCommissioner of Fish and Fisheries of U. S. for 1884, p. 934.
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Mayer,1910, Medusae of the World, vol. 3, p. 561.?Bkowne, 1910, NationalAntarctic Expedition, Nat. Hist., vol. 5, Coelenterata, V. Medusae, p. 47.Generic characters.?Coronatae with numerous (nine or more) ten-tacles and equally numerous marginal sense organs. Twice as manymarginal lappets as sense organs. Eight adradial gonads and fourinterradial subgenital ostia. Four lips. The tentacles and marginalsense organs alternate regularly, but the insertions of the tentaclesand their pedalia are higher up on the sides of the exumbrella thanare the insertions of the pedalia of the sense organs.The Albatross collection serves to show that A. wyvillei and A.bairdii are closely related if not mere extremes of an intergradingseries of one and the same species. For example, two specimensfrom station D. 5652 in the Gulf of Boni, depth of 525 fathoms, havethe margin of the central lens distinctly notched with radial fur-rows as in the typical A. wyvillei; but there is an annular ridgeon the outer side of the ring-furrow with a plain peripheral marginas in A. bairdii. Also several other specimens show such very slightnotches in the margin of the central lens that if one were not look-ing carefully for this feature it would surely pass unobserved andthe medusa would be called A. alexandri. A large specimen of A.giyantea, from a depth of 519 fathoms in Buton Strait, shows affini-
1 Papers from the Tortugas Laboratory of the Carnegie Institution of Washington,vol. 2, p. 155.
196 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.
ties with A. wyvillei, A. bairdii, and A. verrillii. Thus the marginof its central lens is irregularly notched as in A. wyvillei, but withoutradial furrows. There is an annular ridge upon the outer side ofthe ring-furrow, and the outer edge of this ridge is simple and entire,as in A. bairdii, in about two-thirds of its circumference, and notchedas in the typical A. giantea in the remaining one-third. The centrallens is more than half as wide as the medusa, as in A. bairdii, A.verrilli, and A. valdiviae.It is evident that intergrading conditions prevail to a hopelessdegree among many of the so-called "species" of Atolla. In fact,I think there are but two well-distinguished species: A. bairdii withsmooth exumbrella and A. chuni with well-developed and quite regu-larly arranged papillae upon the exumbrella sides of the lappets.As a matter of convenience, however, we may distinguish A. bairdiivar. wyvillei by the notched margin of its central lens, and the ab-sence of a well-marked annular ridge on the outer side of its coronalfurrow; for while there is often an annularridge on the outer side of the ring-furrow,the margin of the central lens usualty pro-jects over it, overarching and concealing itfrom view. A. bairdii is a case where thisridge is so well developed that it projectsvar. wyvillei from the beyond the margin of the central lens.Philippines Gulf op g 35 specimens f Atolla Were foundBon 1 ; depth, 492 fath- roms. by the Albatross among the PhilippineIslands; and of these 18 are more or less typical A. vjyvillei, 4 areintermediate in condition between A. wyvillei and A. alexandri, 3are intermediate between A wyvillei and A. bairdii, 1 combines thecharacters of A. bairdii with the forma valdiviae.The specimen of Atolla gigantea, Cat. No. 28690, U.S.N.M., whichshows affinities with A. bairdii and A. verrillii, was dredged at Sta-tion D. 5647, from Buton Strait, about 11.6 miles off North Island,depth 519 fathoms, bottom green mud. This medusa is a large one,being 130 mm. wide and with 29 tentacles. The diameter of the cen-tral lens is 94 mm. and the thickness of its gelatinous substance 21mm. The margin of the central lens is irregularly notched, but iswithout distinct radial furrows. Its margin overarches the ring-furrow, which is 15 mm. deep. The outer edge of the ring-furrowexhibits an annular ridge, which is, however, overarched and hiddenunder the projecting margin of the central lens. Two-thirds of thecircumference of the peripheral edge of this annular ridge is entireand plain, as in A. bairdii, but about one-third of it is notched, thenotches tending to lie in the radii of the tentacular pedalia. as inthe typical A. gigantea.
SCYPHOMEDUSAE COLLECTED BY STEAMER "ALBATROSS." 197Other dimensions of this medusa (given in millimeters) are asfollows: Tentacular pedalia, 11.5 long, 10.5 wide; rhopalar pedalia,11.5 long, 8.5 wide; diameter across subumbrella to outer edge ofring-muscle, about 126; width of ring-muscle, 77.5; diameter acrosszone of gonads, about 96 (each gonad is circular, disk shaped, andabout 15 in diameter, the medusa being a female and apparentlynearly mature); diameter across central stomach, about 66; lengthof manubrium, 46.It thus appears that in this specimen the central lens is morethan half as wide as the medusa, as in A. bairdii and A. verrilUi,its outer margin being slightly notched as in A. bairdii and A. val-divae. The annular ridge is probably concealed under the over-arching edge of the central lens, as in A. wyvillei and A. verrillii.Part of the outer edge of the annular ridge is entire and even, as inA. bairdii and A. valdiviae, and part of it is notched and furrowed,as in the typical A. gigantea.It is probable that most of the so-called specific distinctions be-tween the various Atollas are mere individual peculiarities of nogreater specific value than the difference between blue eyes andbrown in man.
List of stations among the Philippines from which specimens of Atollas alliedto A. wyvillei were obtained.D5201, April 10, 1908, from Sogod Bay, Southern Leyte Island, depth 554fathoms, bottom gray sand and mud. One typical A. wyvillei with 23 ten-tacles. Dimensions given in table. Cat. No. 28231, U.S.N.M.D 5285, July 20, 1908, about 17.5 miles off Malavatuan Island in the China Sea,depth 272 fathoms, bottom soft mud. Eleven specimens of medium size, notwell preserved, but all are probably A. wyvillei. Cat. No. 28685, U.S.N.M.D5348, December 27, 1908, in Palawan Passage, about 33.5 miles from PointTabonan, depth 375 fathoms, bottom coarse sand. One specimen too imper-fect for accurate specific determination. Cat. No. 28681, U.S.N.M.D 5471, June 19, 1909, about 15 miles off Sialat Point Light on the east coastof Luzon, depth 568 fathoms. Two specimens closely allied to A. alexandri,one with 29 and the other with 26 tentacles. The dimensions of both aregiven in the table. Cat. No. 28686, U.S.N.M.D5486, July 31, 1909, about 6 miles off Batobolo Point, between Leyte andMindanao, depth 585 fathoms. Four badly preserved specimens, too imperfectfor specific determination. Cat. No. 28708, U.S.N.M.D 5493, August 2, 1909, about 5.5 miles off Diuata Point, between Leyte andMindanao, depth 47S fathoms, bottom green mud. Four specimens of A.wyvillei. The largest was 60 mm. wide, central lens 44 mm. wide with in-dented margin, 22 tentacles each 21 mm. long. Another medusa was 50 mm. indiameter, central lens 29.5, with indented margin and 23 tentacles. Anothermedusa was 42 mm. in diameter, central lens 33 mm. wide with distinctradial furrows and notched margin, 24 tentacles. The smallest medusawas 31 mm. wide, central lens 21 mm. wide with radial furrows and notchedmargin, 23 tentacles. Cat. No. 28689, U.S.N.M.
198 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.D 5533, August 19, 1909, between Cebu and Siquijor, about 9.5 miles off Balica-sag Island, depth 432 fathoms, bottom green mud and sand. One specimenof A. wyvillei. Cat. No. 286S4, U.S.N.M.D 5631, December 2, 1909, south of Patiente Strait, about 4.5 miles off DoworraIsland, depth 809 fathoms, bottom green mud. One specimen approachingcondition of A. alexandri in having no distinct indentations or furrows atthe margin of the central lens. The dimensions of this specimen are givenin the table. Cat. No. 28682, U.S.N.M.D5650, December 17, 1909, in the Gulf of Boni, about 12.5 miles off LamuluPoint, depth 540 fathoms, bottom green mud. One specimen distorted bypressing upon the bottom of the bottle in which it was preserved, so that itsspecific affinities can not be determined with accuracy. The medusa was 60mm. wide, central lens 42 mm. wide, with a few faint indentations in itsmargin. Twenty-four tentacles. Cat. No. 28683, U.S.N.M.D5652, December 17, 1909, in the Gulf of Boni about 7.5 miles off Lamulu,depth 525 fathoms, bottom green mud. Four specimens, one of which is 89mm. wide with 31 tentacles and is an A. wyvillei approaching the conditionof A. alexandri. Its dimensions are stated in the table. Another specimenhas five furrows in the margin of its central lens, but there is a plain-edged,projecting annular ridge on outer side of ring furrow, as in A. bairdii. Itthus combines characters of A. loyvillei with those of A. bairdii. Thismedusa is 51 mm. wide and has 29 tentacles. Two other specimens, one33 and the 37 mm. in diameter, have each 20 tentacles, with the marginsof their central lenses notched as in A. loyvillei, while they have well-developed annular ridges which project beyond the margin of the ringfurrow, resembling A. bairdii in this respect. Cat. No. 28687, U.S.N.M.D 5657, December 19, 1909, in the Gulf of Boni, about 15.5 miles off Oland Point,depth 492 fathoms, bottom gray mud. One specimen of A. loyvillei 54 mm.wide, central lens 41 mm. wide, 25 tentacles. Cat. No. 28688, U.S.N.M.Dimensions (in mm.) of some specimens of nontypical and typical A. wyvillei, collectedby the "Albatross" in the Philippine Islands.
Diameter of entire medusaThickness of bellNumber of tentaclesDiameter of central lens of exumbrellaCondition of margin of central lens of exumbrellaDepth of coronal furrowLength of tentacular pedaliaWidth of tentacular pedaliaLength of rhopalar pedaliaWidth of rhopalar pedaliaLength of marginal lappetsLength of tentaclesDiameter to outer edge of ring-muscle of subumbrella
.
Width of subumbrella ring-muscleDiameter across zone of gonadsDimensions of each gonad (circumferential Xradial)...Diameter of central stomachLength of manubrium
FormaA . wyvilleiapproach-ing theconditionof .4.alexandri.^
(?)
19735.547
I A. wyvilleiTypical A . approach-wyvillei? ' ing A
.
! alexandri? A . wyvilleiapproachingA . alexandri.*
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1348328.57X521
682345.5(')5.756.756.755.251456.5438(9 )24
911929549.58.257.56.255.75
4482525( 8 )464.53.53
754.550.5
SOYPHOMEDUSAE COLLECTED BY STEAMER ' ' ALBATROSS." 199ATOLLA BAIRDII forma VALDIVIAE Vanhoffen.Atolla valdiviae Vanhoffen, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedi-tion, Dampfer Valdivia, vol. 3, Lief. 1, p. 13, pi. 1, fig. 3 ; pi. 6, figs. 41-46.?Maas, 1903, Schphomedusen der Siooga Expedition, Monog. 11, p. 17, pi. 1,figs. 3, 4 ; pi. 3, fig. 23 ; pi. 12, fig. 108.?Mayeb, 1910, Medusae of the World,vol. 3, p. 565, fig. 358.This form is very closely related to A. bairdii, but the central diskis only half as wide as the medusa, and the four septal nodes arewider than in A. bairdii.The Albatross obtained three specimens (Cat. No. 27927, U.S.N.M.)of this medusa in the Philippine Island on April 10, 1908, at dredgingstation No. 5202, in Gogod Bay, depth 502 fathoms, bottom greenmud. The characters and dimensions of these specimens are givenby Mayer.1 Genus PELAGIA Peron and Lesueur, 1809.Pelagia Pebon and Lesueub, 1809, Annal. du Mus. Hist. Nat. Paris, vol. 14,p. 349.?Mayeb, 1910, Medusae of the World, vol. 3, p. 570.Generic Characters.?Semaeostomata of the family Pelagidae, inwhich the central stomach gives rise to 16 completely separated radi-ating pouches, 8 in the radii of the tentacles and 8 in the radii of thesense-organs. There are 8 adradial tentacles, 8 rhopalia, and 16partially cleft marginal lappets. Each of the 16 stomach pouchesis cleft at its distal end, where it enters the marginal lappets. Thereis no ring-canal. PELAGIA PANOPYRA Peron and Lesueur.Medusa panopyra Peron and Lesueub 1807, Voyage aux terres Australes,pi. 31, fig. 2.Pelagia panopyra Mayer, 1910, Medusae of the World, vol. 3, p. 575.This widely distributed tropical Pacific species is distinguished byits small, low, rounded exumbrella warts, which are elliptical in out-line and have a longitudinal furrow with cross-foldings. The bellbecomes about 50 mm. wide, and the esophagus is about as long as thebell-diameter, the mouth-arms being somewhat longer. In commonwith other species of Pelagia, the color is highly variable, but the bellis usually rose-colored or violet, and the nettling-warts are violet.The gonads are usually purple and the mouth-arms violet.There are 21 specimens of this medusa in the collection of theAlbatross from the Philippines; 10 (Cat. No. 28719, U.S.N.M. arefrom Station 5422, March 30, 1909, from a tow made at a depth of15 feet below the surface, about 10 miles off Lusaran Point Light,between Panay and Guimaras; the largest specimen is 33 mm. wideand with large gonads, while another 28 mm. wide has only smallgonads. Eleven young specimens (Cat. No. 28716, U.S.N.M.) were
1 Medusae of the World, vol. 3, p. 565.
200 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.
obtained at Station D5220, April 24, 1908, on the surface betweenLuzon and Marinduque at 12.57 p. m. These are all small and about10 mm. in diameter, evidently being taken from a swarm of immatureindividuals. Genus CHRYSAORA Peron and Lesueur, 1809.Chrysaora Peron and Lesueur, 1809, Annal. du Mus. Hist. Nat., Paris, vol. 14,p. 364.?Mayer, 1910, Medusae of the World, vol. 3, p. 577.Generic Characters.?Pelagidae with 8 marginal sense-organs, 24(3X8) tentacles, and typically 32 (4X8) marginal lappets; althoughin C. malanaster the lobes may still further divide, giving 48 (6X8)marginal lappets. CHRYSAORA MELANASTER Brandt.Chrysaora melanaster Brandt, 1838, Mem. Acad. Sci. St. Petersburg, Sci.Nat., ser. 6, vol. 4, p. 385, pis. 16, 17?Maver, 1910, Medusae of theWorld, vol. 3, p. 582.There is a well-preserved specimen of this medusa from stationD5461, June 14, 1909, San Miguel Bay, east coast of Luzon, depth 11fathoms (Cat. No. 28712, U.S.N.M.).The bell is about 130 mm. in diam-eter and slightly flatter than ahemisphere. There are 16 radiatingspoke-like streaks of faint umbercolor extending from near the apexof the exumbrella to the bell-marginin the radii of the 16 cleft velarlobes. These 16 streaks occupy de-pressed radial areas sunken belowthe general level of the contour ofthe exumbrella, and they are be-sprinkled coarsely with wart-likenematocyst clusters of cinnamon-brown color.There are 8 rhopalia, 3X8 tentacles, and 6X8 marginal lobes. Thevelar lobes are cleft as in Brandt's figures and are nearly similar inshape and size to the ocular lappets. They are, however, not nar-rower at the base than outwardly, as in Brandt's figures, but are ovaland taper quite regularly from base to tip.The tentacles are short and slender, the longest being not over60 mm.The mouth-arms are long and slender, folded complexly, and about170 mm. long. The gonads are well developed, apparently mature,and protrude through the subgenital ostia, the subgenital ostia beingfully twice as wide as the perradial columns between them. Thus
Pig. 8.
?
Chrysaora melanaster.octant of the bell margin. AnSan-Miguel Bay, east coast of Luzon.
SCYPHOMEDUSAE COLLECTED BY STEAMER "ALBATROSS.' ' 201
each ostium is 28 mm. long (circumferentially) and 15 mm. wide(radially), while the perradial columns of the mouth-arms are only13 mm. wide.In formalin the general color of the medusa is milky custard-yellow, the gonads being lighter. The apex of the exumbrella isbesprinkled with cinnamon-colored nematocyst warts, and the 16radial streaks of light umber color are also besprinkled with brown-colored clusters of nematocysts. This medusa is widely distributedover the north Pacific from Kamtschatka to California, but thisPhilippine Island specimen is the first which has been obtained inthe tropics. Genus DACTYLOMETRA L. Agassiz, 1862.Dactylometra L. Agassiz, 1862, Contributions to Nat. Hist. U. S., vol. 4,p. 166.?Mayer, 1910, Medusae of the World, vol. 3, p. 583.Generic Characters.?Pelagidae with 8 rhopalia. 5X8 tentaclesand 6X8 marginal lappets.DACTYLOMETRA AFRICANA Vanhoffen.Dactylometra africana Vanhoffen, 1902, Wissen. Ergeb. deutsch. TiefseeExpedition, Dampfer Valdivia, vol. 3, Lief. 1, p. 40, pi. 4, fig. 20.
?
Mayer,1910, Medusae of the World, p. 588.Disk 100 to 130 mm. wide with exumbrella thickly covered withwart-like clusters of nettling cells. Six well-developed marginallappets and five long tentacles in each octant. Lappets and tentaclesred. Red radial streaks over exumbrella. Esophagus, palps, andgonads not highly colored. Colors of large specimens duller andmore brownish than those of small medusae and not unlike the color-ation of D. quinquecirrha. Distinguished by its lappets being deeplypigmented near the margin on the exumbrella side.Vanhoffen's specimens came from the Great Fish Bay, coast ofGerman southwest Africa, in October, 1898. Five specimens (Cat.No. 28679, U.S.N.M.), all imperfect, the largest about 105 mm. indiameter and with only 3X8 tentacles and 4X8 marginal lappets,were found by the Albatross at station D5461, June 14, 1909, at adepth of 12 fathoms, about 7.2 miles off Corregidor Light, ManilaBay, Luzon. The bells are pinkish in hue, and thickly and uniformlybesprinkled over the exumbrella with red-brown mematocyst warts.The lappets are edged on the exumbrella side with reddish brown.The tentacles have been lost and the mouth parts are imperfect.Another specimen (Cat. No. 28680, U.S.N.M.), 166 mm. in di-ameter and with mouth-arms 280 mm. long, was found at Kowloon,China, on August 14, 1908. It was 3X8 tentacles, and 6X8 marginallappets, the lappets being edged on their exumbrella margins withrusset brown.
. . Tzr sr??s y^r:Llr-i 1 records a D+. tylomiit a -from thf .iich hebehY - - ientical with P. ^MinyuorivrkM The bell is white, trans-lucent, and covered on the exnmbrella with minute white 5[ IsL _ amens were all in die Ckrytmorm stage ~ : M :fntaclesand 32 marginal lar els The sting which this medusa inflic:-apparently far more serene than ths: given by : 'ylometra of
; ur A-.rr::..:. : r
"itlrri- - rzimssber. Akatd. Wissea. Berlin. Jaarg. 1385.
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"
\ _ ?- '_" _ ? r-.rfsee KurdH iI>?r:pfer r?24rno. \vL 3. Iiet 1- p. 37.?SU~
z
me ;z Ba
?
- \mrmei*r^?Pelagidae with U : gnal sense-organs. 16
: I . :
. peripheral stomach-pooches in die radii of
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gfer TwtM t i; wi *. Lief. 1 : i. K : :tx : : - : -_- ?
- : "i :_-_-;_ "~_ r-r. n:
SCYPH03JEDUSAF. COLLECTED BY STKi :: ;
-
:
'
I \ I
A perfect specimen (Cat. Ha ." -~ T.S.X.M. . found bjATbatrozs on March *. 1908, at station D5175. in the Snlu Sea-southeast of Cagayanes Islands. Philippine Islands, had a bellmm. wide, palps 46 long, central stomach 35 wide, contracted tenta-cles 65 long, and w:: _ 30 finger-shaped projections upon each2 One large imperfect specimen about 97 mm. in diameter is fromtkm D5991, July 23, 1908. depth of 173 fathoms, amiles off Escarceo Light, southern Luzon.These have between 11 to
- Eight other specimens ofmedium size are from -tion D5386, March 9. 1909(Cat. y _--.-.u.s.y
:
Depth 2S7 fathoms, about
. : miles off Arena Point.Bagay Gulf. Luzon.One imperfect sp*
I N 287 :. UJSJOIis from station D5532, Au-
_ st 13. 1909, from be-tween Masbate and Leyte.Bell about 73 mm. wideand wit _ I _ finger-shaped processes on thegonads.One perfect specimen. . : 1 _ : ~ _ U.S.X.M - : - ;I "-' Jane 7. 1909. east coast of Luzon, about 6.7 miles off LegLig.it. G?ius DISCOMEDUSA Claas, lST"D'feowKdu4c 1 : 7T. DpnLwhriTL, Wien Acad, ^oL 3 -^Ulwtcris-T-Umbn&a Hazcsiz. 1880, Sjrsfc. der Metinsen. p '
-
D .brosa Ma Z? -Minos Antaictiqpe Fraueaise. Metfoses. p. 9.Z>i*coae#Ma Matis. 1910. Mnteir of the World. toL 3, p. 606.zraeterz.?Ulmaridae with _ I * tentacle s
- B lappets, and B sense-orgins. The toBtades arise from theclefts between the marginal lappets. There t: ile. unbranched.adradial canal. S branched. 1 perradiaL and 4 interradial canals, anda marginal ring-canaL
-
:-'--Fig. 10.
?
Discokzbcsa isnonii rant Conr*-3.3.T. Fhiupftsx Islaxrs, apxtl 20. 1906.
:r r: r >. ?HmprnoDi&xmedMM phQippima IEatxs. 1 Issse of die World toL S. p. 607.fig. 388.
- medusa bears a close resemblance to Pannmbrosa poiylobat*Kisiiinouye 1 1 , J urnal Collt r I S ence. Tokyo. Jarar
204 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.
art. 9, p. 19, pi. 4, figs. 20-23) ; but in Kishinouye's medusa the mar-ginal lobes are still further divided, being 64 instead of 32 as inD. philippina. Moreover, the canal system appears to be less com-plex in its branching in P. polylobata, so that it seems probable thatthe two are distinct species, not mere growth-stages one of the other.Should this view prove erroneous, however, the medusa should becalled Parumbrosa polylobata Kishinouye.Six specimens were collected by Fisheries steamer Albatross, sta-tion 5213, Masbate Island, Philippine Islands, April 20, 1908. (Cat.No. 27948, U.S.N.M. Type.)Genus AURELLIA Peron and Lesueur, 1809.Aurellia Peron and Lesueur, 1809, Annal. du Mus. Hist. Nat., Paris, vol. 14,p. 357.Aurelia Lamarck, 1817, Syst. Aniin. sans Vertebres, vol. 2. p. 512.Aurellia Mayer, 1910, Medusae of the World, vol. 3, p. 619.Generic Characters.?Semaeostomata with a simple central mouth-opening, which is surrounded by four well-developed, perradiallysituated, unbranched mouth-arms or palps. Eight marginal sense-organs. The tentacles are small and alternate with an equal numberof short lappets. Both tentacles and lappets arise from the sides ofthe exumbrella a short distance above bell-margin. The bell-marginis divided into 8 or 16 broad velar lobes. The central stomach givesrise to a number of branched, radiating canals which anastomose andare connected by a marginal ring-canal. There are four interradialgonads and four well-developed subgenital pits.AURELLIA AURITA (Linnaeus).Medusa aurita Linnaeus, 1758, Systema Naturae, ed. 10, vol. 1, p. 660.Aurellia flavidula Peron and Lesueur, 1809, Annal. du Mus. Hist. Nat.,Paris, vol. 14, p. 369.Aurellia aurita Lamarck, 1S17, Hist. Anim. sans Vert., vol. 2, p. 513.
?
Mayer, 1910, Medusae of the World, vol. 3, p. 623.Some 13 specimens of this universally distributed medusa werefound by the Albatross among the Philippine Islands. Of these, 11half-grown specimens (Cat. No. 28718, U.S.N.M.) are from stationD 5663, December 28, 1909, in Macassar Strait, depth 11 fathoms,about 1.7 miles off Kapoposang Island, 7h. 20m. p. m., while 2 largerbut still immature are from Station D 5662, December 21, 1909, FloresSea, near Tana Keke Island, 5h. 40m. to 6h. 12m. a. m. There areonly 8 notches in the bell margin corresponding to the 8 rhopalia,instead of 16 notches, 8 rhopalar and 8 inter-rhopalar as in Aurellialabiata.Aurellia aurita is found in all seas from the Polar regions to theTropics. In the Tropics it lives very close to its heat death-temper-
>?'"''"??,
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS." 205
ature, and thus it is barely able to survive in the surface waters ofthe warmer seas in summer. Romanes found that specimens of thismedusa from the British seas can withstand being frozen solidlyinto ice, and I find this to be true also of this medusa from Halifax,Nova Scotia. At Halifax, on the other hand, the medusa ceases topulsate at 29.4? C.,at which temperature it is most active atTortugas,Florida. On the other hand, the Florida medusa is killed by beingfrozen into the ice. Thus the medusa becomes somewhat acclimatedto the temperature of the waters in which it lives, and if accustomedto warm water it loses its resistance to cold and the opposite.AURELLIA LABIATA Chamisso and Eysenhardt.Aurelia labiata Chamisso and Eysenhardt, 1820, Nova Acta Phys. med.Leop. Car., vol. 10, p. 358, pi. 28, figs. 1 A. B.Aurellia labiata Mayek, 1910, Medusae of the World, vol. 3, p. 628, fig. 398.
?
Light, 1914, Philippine Journal of Science, vol. 9, p. 200.Light (1914) rec-ords a specimen fromthe Philippines, thebell of which was 225mm. in diameter.Aurellia labiata dif-fers fromA . aurita byhaving 16 notches inits bell-margin, by itspeculiar velum - like,interrhopalar, subum-brella membranes rep-resenting the true bell-margin, and by thevery small size of itssubgenitalostia. Themouth-arms are alsoshorter than one com-monly observes themto be in J., aurita, andthe terminal branchesof the radial-canals,anastomose to agreater degree than inA. aurita.The dimensions of three specimens, Cat. No. 27923, U.S.N.M., ob-tained by the United States Fisheries Bureau steamer Albatross at
Fig. 11.
?
Aurellia labiata fkom the Philippines. A,Oral view showing the 16 notches in the bellmagrin. b, aboral view of tentacles and lappets.C, Diagrammatic cross section of bell margin.c, eing canal ; ex, exumbrella ; I, lappets ; re, racialcanal ; sw, subombp.ella ; t, tentacle ; v, velarium.
206 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.Masbate Anchorage, Philippine Islands, on April 21, 1908, are asfollows
:
mm. mm. mm.Diameter of umbrella 174 1S9 128Diameter across zone of gonads 57 53 42Length of each mouth-arm 74 75 52Four other specimens (Cat. No. 27979, U.S.N.M.) were caught atnight upon the surface under the electric light at Jolo Anchorage, onFebruary 8, 1908; while seven more (Cat. No. 27924, U.S.N.M.) weretaken at Albatross station D 5230, between Bohol and Leyte, May 7,1908.An abnormal but perfect specimen of Aurellia aurita was foundat Tortugas, Florida, on July 27, 1914. This aberration closely re-sembled the normal A. labiata of the Pacific. There were 16 deepnotches in the bell-margin, 8 perradial and 8 interradial, and thevelumlike marginal membrane was like that of A. labiata. Thesmall subgenital ostia and simple mouth-arms also recalled A. labi-ata; but the terminal branches of the adradial-canal system did notanastomose and in this resembled A. aurita rather than A. labiata.The case is interesting, as it leads one to suspect that A. labiata ofthe Pacific has been derived as a mutation from the universally dis-tributed A. aurita. The bell of this Tortugas medusa was 270 mm.in diameter; diameter of genital cross, 78 mm. Palps simple andeach about one-eighth longer than the bell-radius. Eight senseorgans, 8 interradial notches in the bell-margin. Eight straight, sim-ple, non-pigmented perradial-canals, 8 straight pink-colored inter-radial-canals. The sparingly branched adradial-canals reach thebell-margin without anastomosing, and in this respect resemble thoseof A. aurita. The male gonads were pink, the tentacles rich purple,the velarium creamy white, the palps purple-pink, and the gelatinoussubstance pink. Order RHIZOSTOMAE.Scyphomedusae without marginal tentacles and with numerousmouths borne upon four dichotonously branched (eight) mouth-arms.Most of these forms are tropical and none are found in Arctic seas.They are the conspicuous large jellyfishes of harbors and coastalwaters in the East Indies, although usually rare and represented bybut few species in the West Indies.Genus CASSIOPEA Peron and Lesueur, 1809.Cassiopea Peron and Lesueur, 1S09, Annal. du Mus. Hist. Nat. Paris, vol.14, genre 24, p. 356.?Mayer, 1910, Medusae of the World, vol. 3, p. 636.Generic Characters.?Rhizostomata pinnata with eight (four pairsof) adradial, complexly branched mouth-arms, the lower or ventral
SCYPHOMEDUSAE COLLECTED BY STEAMER "ALBATROSS." 207
surfaces of which bear numerous mouth-openings and vesicles.There are four gonads and four separate subgenital cavities. Thereare more than eight marginal sense-organs and twice as many radial-canals as sense-organs. The radial-canals are placed in communica-tion one with another by means of an anastomosing network of ves-sels. A well-defined ring-canal may or may not be present, but is'commonly absent.CASSIOPEA ANDROMEDA, var. BADUENSIS Mayer.Medusa andromeda Forskal, 1775, Descript. que in Itinere Orientali Ob-servait, Hauniae, p. 107, pi. 31.Cassiopea andromeda Eschscholtz, 1829, Syst. der Acalephen, p. 43.Cassiopea andromeda, var. oaduensis Mayer, 1915, Publication No. 212,Carnegie Institution of Washington, p. 183.A specimen of this medusa (Cat. No. 28730, U.S.N.M.) is fromEndeavour Strait, between Australia and New Guinea, and wasfound by the Albatross on December 23,1908. The bell is 101 mm. in diameter,flat without an aboral depression, and oS^^?^0^ {' '' - V-with 18 rhopalia. There are four to eight, X^-^^ipw^S'^l^m 5 n i in t n ^v^'-' r,>*I ; lm#2WPPusually six, lappets between successive -S^^^.^S0^?M: :iirhopalia. The arm-disk is octagonal, 36mm. wide, and the eight mouth-arms are f^eeach 34 mm. long and definitely bifur- V;cated, the forks being 16 mm. long, thus ^nearly half as long as the total length ofthe mouth-arms. There are no append- fig. 12.
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cassiopea andbomeda,ages among the mouth-arms, but these ^^lZ7iIZV?Zlmay have been lost. The color has wholly view on the right ; aboealfaded in formalin. VIEW 0F BELL ON THE LEFT-Another specimen of this medusa was found at Badu Island, TorresStraits, Australia, within a few miles of Endeavour Strait, by theexpedition of the Carnegie Institution of Washington, on November5, 1913, and was studied alive, and is herewith figured.The bell is 61 mm. in diameter. There are 22 marginal sense-organs and (5X22) 110 bluntly rounded, barely perceptible, evenlyspaced marginal lappets all similar each to each. The arm-disk isabout one-third as wide as the bell-diameter and the mouth-arms arecompressed dorso-ventrally and when fully expanded extend slightlybeyond the bell-margin, but in their ordinary state of contractionthey do not quite reach the bell-margin. The side branches of thesemouth-arms are short, but each arm is bifurcated at its outer end, theforked part being about one-third as long as the entire arm. Thereare about 20 slender, flat, tapering, central arm-disk appendages ofvarious lengths, the longest of which is at the center of the oral side
208 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.and is about as long as the bell-radius. There is also a spatula-shaped appendage at the crotch of bifurcation of each mouth-arm.These are somewhat stouter than the central appendages, about halfas long as the bell-radius, and their entoderm is bluish. There arenumerous, minute, spatulate appendages among the mouth-arms.The general color of the bell of the medusa is olive-brown. Thereare 22 large, triangular white spots with forked outer ends near thebell-margin in the radii of the sense-organs, and also (3X22) 66short white streaks near the margin in the radii of the velar lappets.There are 22 interradial, dull bluish streaks in the subumbrella alter-nating with the rhopalia in position.This variety is distinguished by its bifurcated mouth-arms. Itsnearest ally appears to be Oassiopea andromeda, var. acyclobliaSchultze, from Amboina, but it differs in its color pattern, in the ab-sence of a central dome, and in its simple bifurcated mouth-arms,those of the Amboina medusa branching dichotomously.Oassiopea and<romeda is the common species of the Indian Ocean,Red Sea, and Malay Archipelago, and Kellar records its havingwandered into the Suez Canal. It gives rise to numerous localvarieties.There are evidently a number of other varieties of Cassiopea in thePhilippines, for Light x describes Cassiopea polypoides, var. culion-ensis, C. polypoides?, and C. medusa, new species. The last namedis distinguished by its very large mouth-arm appendages, which ina medusa whose bell is 260 mm. in diameter are 110 mm. long and7.5 mm. in diameter, being cylindrical near the base and flattened attheir outer ends. C. medusa is described from Culion Bay, Culion,Philippine Islands.Genus CEPHEA Peron and Lesueur, 1809.Cephea Peron and Lesueur, 1809, Annal. du Mus. Hist. Nat., Paris, vol. 14,p. 360.?Mayer, 1910, Medusae of the World, vol. 3, p. 651.Generic Characters.?Ehizostomata dichotoma in which the eightmouth-arms fork once dichotomously and each fork gives rise toshort dichotomous or dendritic branches. Solid, wart-shaped tuber-cles at the center of the exumbrella. The central stomach gives riseto eight rhopalar and numerous inter-rhopalar radial-canals, all ofwhich connect with a network of anastomsing vessels in a wide zonenear the margin. Rhopala without ocelli and without sensory pitson the exumbrella. There is no definite ring-canal. Developmentunknown.
1 1914, Philippine Journal of Science, vol. 9, p. 201.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 209CEPHEA OCTOSTYLA (Forsk&l).Medusa octostyla Forskal, 1775, Descript. Anim. Itin. Orient., p. 106, No.18, Icon., pi. 29.Cephea octostyla Mayer, 1910, Medusae of the world, vol. 3, p. 652, fig. 405.A number of well-preserved specimens of Cephea octostyla wereobtained by the following localities
:
Jolo, Philippine Islands, February 8, 1908, 1 specimen. Cat. No.27906, U.S.N.M.Jolo Anchorage, Philippine Islands, February 13, 1908, 3 speci-mens. Cat. No. 27908, U.S.N.M.Jolo Anchorage, Philippine Islands, March 5, 1908, 1 specimen.Cat. No. 27907, U.S.N.M.Station 5173, Jolo, Philippine Islands, March 5, 1908, 9 specimens.Cat. No. 27910, U.S.N.M.
"<Vi%M ?5~m^pi#>? i ? ? . i ii )>?&$
.'?x-s ;,:.;;.; v ,
Fig. 13.
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Cephea octostyla from Jolo Anchorage, Philippine Islands. A, B, andC, Variations in the development of the central warts of the exumbrella.D, Rhopaliom from the subdmbrella side.Bell 90 mm. wide, exumbrella flat, rim vertical, 20 mm. high.Near the center of the exumbrella there is a zone of numerous lowwart-like protuberances, leaving the exact center smooth. Eightrhopalia without ocelli and without sensory pits. Seven velar andtwo ocular lappets in each octant, all similar each to each and in-distinct rectangular in outline and separated by very slight indenta-tions spanned by a web. Fairly deep grooves extend up the verticalrim of the exumbrella surface of the bell between the lappets.Arm disk as wide as the bell radius. At its center 4 to 12 or moretapering, somewhat flattened, wart-covered filaments about one-fourth as long as the bell diameter. Each filament terminates in asimple, slender, pointed end. There are also numerous simple shortfilaments ranging from 15 to 5 mm. in length near the center of thearm disk, and in addition many still shorter arms between thenumerous frilled mouths of the mouth arms.Four small subgenital ostia and a unitary genital cavity. Ringmuscles of subumbrella entire, but weakly developed. Eight rhopa-101825??Bull. 100?17- -14
210 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.lar and (8X7) somewhat narrower radial canals all connected byanastomosing side branches. The eight rhopalar canals lead straightout to the rhopalia, but the 56 inter-rhopalar canals tend to losethemselves in the network of vessels. No distinct ring canal.Numerous small yellowish spots over the exumbrella and reddishbrown dots over the outer surface of the arm disk and reddish-brownstreaks around the warts of the exumbrella.CEPHEA CEPHEA, var. COEKULEA, atypical.Cephea coerulea Vanhoffen, 1902, Wissen. Ergeb. deutsch. Tiefsee Expedi-tion Valdivia, vol. 3. Lief. 1, p. 45, fig. 13, 14.
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Mayer, 1910, Medusae ofthe World, vol. 3, p. 657.One well-preserved specimen of this medusa was found by theAlbatross at Station D 5457, June 8, 1909, 5 miles off Legaspi Light,east coast of Luzon. Cat. No. 28715, U.S.N.M.The bell is flat, 109 mm. wide, with a low dome-like apex 34 mm.wide, which is completely covered with long, conical, wart-shapedprotuberances. There are two small ocular and eight completelyfused velar lappets in each octant, forming mere thickenings on theexumbrella side of the bell wall and bridged over on the subumbrellaside by a web of tissue. The arm disk is 50 mm. wide and themouth arms are laterally flattened, 29 mm. wide in the radial direc-tion, and 44 mm. long. There are three to six slender filamentsamong the mouths of each moutl> arm. These taper to pointed endsand the longest are only 13 mm. long. There are no filaments uponthe central parts of the arm disk, but there are some at the bases ofthe mouth arms. The filaments are thus much smaller and lessnumerous than in Vanhoffen's C. coerulea from the east coast ofAfrica.There are eight rhopalar radial canals which extend straight tothe sense organs, and in addition there are from five to seven inter-rhopalar canals in each actant, which anastomose and lose their iden-tity in a wide network of vessels which send branches to the rhopa-lar canals. The subgenital porticus is unitary. A color note statesthat the mouth arms were pale hyaline blue and raw umber.CEPHEA, species.Five specimens of Cephea, too poorly preserved to be determinedspecifically, were collected at the following stations
:
April 25, 1909, from Manila Bay, surface.April 29, 1909, from Manila Bay, behind the breakwater.D 5452, June 7, 1909, off Legaspi Island, east coast of Luzon.D 5453 June 7, 1909, off Legaspi Island, east coast of Luzon.D 5461, June 14, 1909, Carino Island, east coast of Luzon.Light (1914) records Cephea cephea from Manila Bay in Janu-ary, 1912.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 211Genus COTYLORHIZA L. Agassiz, 1862.Cotylorhiza Agassiz, L., 1862, Contr. Nat. Hist. U. S., vol. 4, p. 152.
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Mayeb,1910, Medusae of the World, vol. 3, p. 658.Generic Characters.?Rhizostomata dichotoma with eight simple,bifurcated mouth-arms, the ends of which branch pinnately. Thefour subgenital ostia are simple and funnel-shaped, and there is asingle subgenital porticus. The appendages upon the mouth-armsare mounted upon pedunculated filaments. There are eight marginalsense-organs and numerous radial-canals which anastomose laterallywithout any definite ring-canal in the adult. The sense-clubs have noocelli and no exumbrella sensory pit. There is a unitary peripheralzone of circular muscles and an inner zone of radial-muscles in thesubumbrella. The exumbrellais smooth and without anaboral " sucker-like " depres-sion, but with a prominent cen-tral dome without wart-shapedelevations upon it.COTYLORHIZA PACIFICA Mayer.Cotylorhiza pacifica Mayek,1915, Publication No. 212.Carnegie Institution of Wash-ington, p. 185.A single specimen of this in-teresting medusa was obtainedat the launch landing in ManilaBay, Luzon, Philippine Islands,on January 24, 1908 (Cat. No.28729, U.S.N.M. type). Un-fortunately it was cut into several pieces before being preserved,and this renders an attempt to study it unsatisfactory in many re-spects. The bell appears to have been about 200 mm. wide, ex-umbrella finely granular, with a central dome as in the Mediter-ranean Cotylorhiza tuberculata.There are 8 rhopalia without ocelli (in formalin), and without ex-umbrella pits, being similar in essential respects to those of Cotylor-hiza, tuoerculata. The rhopalar lappets are short and pointed.There are about eight irregularly spaced, bluntly pointed, large velarlappets in each octant, and deep furrows between them extend radi-ally inward over the exumbrella, as in C. tuberculata. The velarlappets vary in length, but the largest are about twice as long andtwice as wide as the ocular lappets.The circular muscles occupy the entire zone of the subumbrellabeyond the arm-disk. They are broken in the eight principal radii,and unlike C. tuberculata there are no radial muscles.
Fig. 14.
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Cotylorhiza pacifica from ManilaBay, Luzon. A, An octant of the bellMARGIN. B, ABAXIAL VIEW OF ONE OF THEMOUTH-ARMS.
212 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.The arm-disk is about 100 mm. wide, the perradial columns beingeach 36 mm. wide. The subgenital ostia are much larger than inC. tuberculata, but they were so mutilated that one can not state theirexact size, which appears, however, to be nearly that of the perradialcolumns themselves. The specimen having been cut into pieces, wecan make no statement concerning the condition of the subgenitalporticus.The eight mouth-arms resemble those of C. tuberculata, but thereare windowlike openings in the lateral membranes, as in Lobonemasmithii. The total length of each arm is 81 mm., the upper arm beingone-fourth as long as the lower arm. At their widest part the armsare about three-fourths as wide as they are long. The center of thearm disk is thickly covered with slender filamentous appendageswhich terminate in nematocyst-bearing, swollen, knoblike ends, as dothe appendages of the mouth-arms of C. tuberculata. In this Philip-pine Island medusa the appendages of the outer parts of the mouth-arms much less numerous and smaller than in C. tuberculata, but aresimilar in general form to those of the Mediterranean medusa. Thelongest are about 15 to 20 mm. long.The cruciform central stomach gives rise to about 140 radial-canals,the eight rhopalar canals being about twice as wide as the others, in-stead of being of the same caliber, as in C. tuberculata. All thesecanals anastomose in a network under the zone of the circular muscles.There is no distinct ring-canal. In formalin the specimen is dulluniform yellowish brown.It differs from Cotylorhiza tuberculata in having no radial-muscles,and in the circular muscles being interrupted in the eight principalradii. The subgenital ostia and arm-disk are larger and the ap-pendages of the mouth-arms smaller and fewer than in G. tuberculata.Moreover, the peculiar perforations in its mouth-arm membranes atonce distinguish this species.These distinctions are indeed of such nature that if one felt so in-clined a new genus could be established to receive this medusa. Ibelieve, however, that its relationships will be more clearly indicatedby placing it in the genus Cotylorhiza, within which it forms a well-marked species. Genus CATOSTYLUS L. Agassiz, 1862.Catostylus (part) Agassiz, L., 1862, Contr. Nat. Hist. U. S., vol. 4, pp. 152,153.Generic Characters.?Rhizostomata triptera, in which the mouth-arms bear neither clubs, filaments, nor other appendages. Sixteenradial-canals, 8 rhopalar, and 8 adradial. The rhopalar-canals ex-tend 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 anas-
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 213tomosing vessels, which may join with the radial-canals, but whichdo not connect directly with the central stomach. Among charactersof minor importance, the marginal zone of circular muscles in thesubumbrella is only partially interrupted in the eight principal radii.There is an exumbrella pit with radiating furrows above each sense-organ. CATOSTYLUS PURPURUS Mayer.Catostylus purpurus Mayek, 1910, Medusae of the World, vol. 3, p. 671, fig.412.
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Light, 1914, Philippine Journ. of Science, vol. 9, p. 207.This form is closely related to Catostylus stiphropeterus, fromTernate, but differs in the number and arrangement of its marginallappets and in its deep uniform purple-brown color. Light (1914)describes this medusa from life, whereas Mayer had only preservedmaterial. In life the bell is higher than a hemisphere, whereas in itscontracted state in preservative fluids, as in our figure, it is flatter
Fig. 15.
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Catosttlus purpurus, Manila Bay, Luzon. A, Oral view., six mouth-armscut off. B, Side view, bell contracted. In life it is nearly hemispherical.C, Genital ostium. D, Exumbrella view of rhopalium.than a hemisphere. When mature it is deep purplish brown, and thesense-organs have brillant silver ocellus-like spots which are largerin small than in full-grown medusae. When young the medusa maybe plum-colored, or even translucent white. Small cyclops-like Crus-tacea were found by Light to be commensal with this medusa, thecrustaceans lying upon the rhopalar canals close to the sense-organs.Light finds that this medusa is not a bottom form, but swims inshallow water near the surface.Seven specimens (Cat. No. 27934, U.S.N.M.) found in Manila Bayon December 9, 1907, are in the collection made by the United StatesFisheries Bureau steamer Albatross, and a larger one (Cat. No. 27980,U.S.N.M.) found on March 11, 1908. This largest specimen servesas the type of the species in the United States National Museum atWashington. Its dimensions in millimeters are as follows : Bell 115wide, evenly rounded, 35 high ; arm-disk 75 wide where it arises fromthe 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.
214 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.Another specimen (Cat. No. 28723, U.S.N.M.) of medium size,nearly mature, was found at Cavite Anchorage, Manila Bay, Luzon,on July 2, 1909.Light (1914) states that this is the commonest medusa in ManilaBay in November and December.CATOSTYLUS TOWNSENDI Mayer.Catostylus townsendi Mayer, 1915, Publication No. 212, Carnegie Institutionof Washington, p. 183.This species is named in honor of the author's friend, Dr. CharlesH. Townsend, the distinguished director of the New York Aquarium.Six specimens were found at Sta-tion D5594, September 30, 1909,about 6 miles off Mount Putri,Borneo, in 11 fathoms. Type, Cat.No. 28722, U.S.N.M.; paratypes28721, U.S.N.M. This medusa isclosely allied to Catostylus purpurusof Manila Bay, Philippine Islands,but in formalin its exumbrella ismilky in color and bespeckled irreg-ularly with numerous conspicuouspurple-brown spots. The mouth-arms are more pointed than inG. purpurus. In some octants ofthe bell-margin the velar lappetsare arranged as in G . purpurus,but they are usually more numerous and more irregularly arrangedthan in G. purpurus.The dimensions of the largest specimen of G. townsendi, stated inmillimeters, are as follows: Bell 97 wide, flatter than a hemisphere,exumbrella finely granular, gelatinous substance of horny rigidity.Shape and consistency of the bell as in G. purpurus. Eight rhopaliawithout ocelli, in formalin, and with a deep dark-colored, furrowed,exumbrella pit.The rhopalar lappets are small and oval, but the velar lappets areabout twice as wide as long. Deep clefts between the lappets extenda short distance up the sides of the exumbrella. The velar lappetsare very irregular in arrangement, although they tend to conform tothat seen in G. purpurus of Manila Bay, Luzon, yet in most of theoctants the subdivisions of the principal lappets are more pronouncedand irregular than in G. purpurus, so that there are usually 7 or 8main velar lappets with 10 to 14 marginal lobes in each octant.The arm-disk is similar in shape to that of G. purpurus. It is 61mm. in perradial and 45 mm. in interradial diameter. The perradial
Fig. 16.
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Catostylus townsendi fromoff Point Putri, Borneo.
SCYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS.' ' 215columns are 17 mm. and the subgenital ostia 20 mm. wide. The pro-jections and papillae of the subgenital ostia are similar to those ofC. purpurus. The subgenital cavity is unitary.The eight mouth-arms are each about 64 mm. long, the upper,naked outer part of each arm being 14 mm. and the 3-winged lowerpart 50 mm. long. The arms are widest at the proximal parts of the3-winged expansions which, when spread out, have a span of about31 mm. They taper to pointed distal ends and have no appendagesamong the mouth-frills.There is a powerful unbroken zone of circular muscles in the sub-umbrella, 27 mm. wide, from the outer edge of the arm-disk to thebell margin. There are also radial muscle fibers on the abaxial sidesof the four perradial columns of the arm disk, as in G. purpurus.Sixteen radial canals arise from the cruciform central stomach.The eight rhopalar canals extend straight to the rhopalia, but theeight adradial canals end in the ring canal, which is beneath anannular furrow or bend in the subumbrella 13 mm. inward from thebell margin. On its inner side the ring canal gives rise to from 4to 7, usually 5, centripetal canals betwen each successive pair ofradial canals. These centripetal vessels anastomose with one an-other and with the 16 radial canals, and end blindly before reachingthe stomach margin. On its outer side the ring canal gives off anetwork of vessels which ramify through the lappets. In formalinthe medusa is opaque milky white with irregularly clustered brownspots over the exumbrella.This medusa is closely related to Acromitus maculosus Light,1 butit has no filaments upon its mouth arms. It is possible, however,that these were lost, but this seems improbable among six well-preserved specimens.Moreover, in Catostylus townsendi the centripetal vessels anasto-mose with the 16 radial canals, whereas in Acromitus, according toLight, they join only with the eight rhopalar canals. It seems prob-able, therefore, that Catostylus townsendi is a distinct species, sepa-rate from Acromitus maculosus Light, and that Acromitus is a genusderived by mutation from Catostylus.CATOSTYLUS MOSAICUS (Quoy and Gaimard).Cephea mosaica Quoy and Gaimard, 1824, Voyage de VUranie, Zoologie.p. 569, pi. 85, fig. 3.Catostylus mosaicus Agassiz, 1862, Contr. Nat. Hist. U. S., vol. 4, p, 152.
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Mayee, 1910, Medusae of the World, vol. 3, p. 666.A single immature medusa (Cat. No. 28720, U.S.N.M.) which maypossibly be the young of C. mosaicus, was taken by the Albatrossin a seine off the beach near the mouth of Malampaya Eiver, Pala-wan Island, Philippine Islands, on December 26, 1908. It differs
1 1914, Philippine Journal of Science, vol. 9, No. 3, sec. D, p. 212, figs. 4-6.
216 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.from C. mosaicus in having only 10 marginal lappets in each octantinstead of about 16, as in G. mosaicus. Moreover, in G. mosaicus thelappets are all long, pointed, and similar in size and shape each toeach, whereas in the Philippine medusa there are eight long, pointedvelar and two much shorter, oval, occular lappets in each octant.The Philippine medusa is, however, quite small, being only 86 mm.in diameter, whereas C. mosaicus becomes fully 350 mm. wide. Thesedifferences may therefore be due to immaturity. In the Philippinemedusa the bell is 86 mm. wide, mouth arms 63 mm. long, the upperarms being 11 mm. and the lower 52 mm. The interradial subgenitalostia are 18 mm. wide, with a large oval or nearly spherical papillaon the subumbrella. The perradial columns of the arm disk are only12 mm. wide, thus the ostia are 1.5 times as wide as the columns.The perradial diameter of the arm disk is 52 mm. and its interradialdiameter 44 mm. The powerful ring muscles of the subumbrella areonly partially interrupted in the eight chief radii. The exumbrellais coarsely granular and besprinkled thickly with numerous minutecinnamon-brown flecks. Other parts of the medusa are pale milkypink. The gelatinous substance is tough and rigid.If this be not G. mosaicus it is certainly very closely related tothis well-known Australian medusa. G. mosaicus is abundant inbays and estuaries along the Australian coast from Melbourne tothe mouth of the Brisbane River in Queensland.In Sydney Harbor all specimens of this medusa are dull creamybrown or yellowish in color, but in Moreton Bay, Queensland, mostof them are cobalt blue. It is interesting to see that H. B. Bigelow xfinds that Stomolophus meleagris in San Diego Bay, California, isPrussian blue instead of being dull yellow, as in the Atlantic.Catostylus mosaicus appears to breed throughout the year inMoreton Bay, Queensland, but in the temperate regions of Australiait is said to become mature only in summer and autumn.Genus LYCHNORHIZA Haeckel, 1880.Lychnorkiza+Cramborhisa Haeckel, 1880, Syst. der Medusen, pp. 587,633.?Mayer, 1910, Medusae of the World, vol. 3, p. 672.Generic Characters.?Rhizostomata triptera with filaments, butwithout clubs, upon the 3-winged mouth-arms. No axial terminalclub at end of each arm, and no club-shaped appendages between themouths. The stomach gives rise to 16 radial-canals?8 rhopalar and8 adradial. The rhopalar-canals extend to the bell-margin, but theadradial ones end in the ring-canal. Blindly ending, centripetalvessels arise from the inner side of the ring-canal and may anasto-mose to some extent. On its outer side the ring-canal gives off a net-work of anastomosing vessels which extend into the lappets.
1 1914, University of California Publications in Zoology, vol. 13, p. 239.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 217LYCHNORHIZA BARTSCHI Mayer.Lychnorhisa hartschi Mayer, 1910, Medusae of the World, vol. 3, p. 674,figs. 413, 414.Named in honor of Dr. Paul Bartsch, of the United States NationalMuseum, to whom we owe the excellent preservation of all specimensof medusae whichpassed through hishands.Bell 84 mm. wide,natter than a hem-isphere and withsmooth exumbrellasurface. Gelatin-ous substance thickbut not very rigid.Eight rh op alia,each with an ocellus,I L 11 FlG- 1^ -?Lychxorhiza eartschi from Jolo Anchorage,ana an eXUmDreiia Philippine Islands. A, Oral view; n, notch in thesensory pit with perradial column of the arm disk ; sgo, sdbgenitali i ;.? ? _ OSTIUM. B, Rhopalium from the exumbrella side.dendritic iurrowsover its floor. About 96 (8X12) lappets, 10 bluntly pointed velarlappets between 2 somewhat smaller ocular lappets in each octant.Arm-disk 51 mm. wide where it arises from the subumbrella, butonly 47 mm. wide at the level of origin of the 8 mouth-arms. The4 subgenital ostia are crescent-shaped and each is covered above bya gelatinous flap. They are only half aswide as the perradial columns betweenthem. Each perradial column exhibits aniche on its outer side which bears a super-ficial resemblance to the subgenital ostia.The subgenital cavity is unitary.
' The 8 mouth-arms are laterally com-pressed and 36 mm. long, the lower 3-winged parts of the arms being 24 mm.long and 23 mm. wide. Numerous simple,laterally flattened, tapering filaments arisefrom between the frilled mouths on allsides of the mouth-arms and from the arm-disk. The filaments upon the arm-disk areabout 30 mm. long, but those from the outer parts of the mouth-arms are shorter.The central stomach is cruciform and about 46 mm. wide. Sixteensimple radial-canals, 8 rhopalar and 8 adradial. These are all putinto intercommunication with a wide ring-canal which is at somedistance inward from the margin. The adradial-canals terminate
Fig. 18.?L tchnorhizabartschi. Side view. Let-tering as in figure 17.
218 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.in this ring-canal, but the rhopalar-canals extend outward to thesense-organs. On its inner side the ring-canal gives rise to 16 blindlyending networks of vessels which do not connect either with thestomach or with the radial-canals. On its outer side a fine-meshednetwork of vessels arises from the ring-canal and fuses with therhopalar vessels. Around the margin at the bases of the lappets isa marginal ring-canal of fine caliber. There is a unitary uninter-rupted system of ring-muscles in the marginal zone of the sub-umbrella, but no radial-muscles.The gelatinous substance is translucent and milky in formalin,and the gonads, mouth-frills, and canal-system are milky yellow.Doctor Bartsch states that these colors in the living animal werenearly as they appear in the specimen preserved in formalin.The type-specimen, Cat. No. 27942, U.S.N.M., was found by theUnited States Fisheries Bureau steamer Albatross at Jolo Anchor-age, Philippine Islands, on February 13, 1908, and three others (Cat.no. 28737, U.S.N.M.) were taken in Limbe Strait, Celebes, in Decem-ber, 1909. The dimensions given above are those of the largestspecimen from the Celebes, this being somewhat larger than the type-species previous^ described.1LYCHNORHIZA BORNENSIS Mayer.Lychnorhiza bomensis Mayer, 1915, Publication No. 212, Carnegie Institu-tion of Washington, p. 191, fig. 6.A single specimen of this medusa was found at Tawao, Borneo, onSeptember 30, 1909, at 9h. 30m. a. m. Cat. No. 28736, U.S.N.M.Type.Bell 89 mm. wide, exumbrella smooth, somewhat flatter than ahemisphere. Eight rhopalia, each with an ocellus and a furrowedexumbrella pit. The rhopalar lappets are very short and lanceolate.In each octant there are usually seven, occasionally eight, velar lap-pets. The lappets adjacent to the rhopalar lappets are about twiceas wide as the remaining velar lappets. All are oval and bluntlyrounded.The arm disk is half as wide as the bell, and the subgenital porticusis unitary. The external faces of the perradial columns have each aslight concavity, not a deep niche, as in Lychnorhiza bartschi. Thesubgenital ostia are twice as wide as the perradial columns and eachis arched over by a flap-like projection. There are also four inter-radial papillae upon the subumbrella, one opposite the opening ofeach subgenital ostium.The eight mouth-arms are not quite two-thirds as long as the bell-diameter, the upper arms being not quite half as long as the lowerarms. Each mouth-arm terminates in a single, slender, tapering,
1 Medusae of the World, p. 674.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 219thread-like filament about 50 mm. long. There are also a few shorter,more slender filaments which arise from the sides of the mouth-arms between the mouths. There are no filaments upon the arm-disk.There is a wide zone of circular muscles in the subumbrella, andthese are only partially interrupted in the eight rhopalar radii.The central stomach gives rise to 16 radial canals, 8 of whichextend to the rhopalia and the 8 others end in the ring canal. Onits inner side the ring canalgives rise to 16X4 radial vessels,which do not reach the marginof the stomach but anastomoseone with another and with the16 chief radial canals. On itsouter side the ring canal givesrise to about 100 radiating ves-sels, all of which anastomose byside branches, forming a reticu-lum in the outer zone of thesubumbrella.In formalin the gelatinous sub-stance is translucent and milky,and the gonads and mouth frillsare milky yellow.The following table will serve to indicate the distinctions betweenLychnorhiza bomensis and Lychnorhiza bartschi of the Philippines
:
Fig. 19.
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Lychnorhiza boenensisTawao, Borneo.
L. bartschi. L. bomensis.
Velar lappets
Interradial papillae on the subumbrella.Perradial niches in the arm-disk.Mouth-armsFilaments
Circular musclesCanal system
Central stomach
(8X12) all oval and similar eachto each.NoneFour deep cleftsLess than half as long as bell-diameter.Numerous. Lateral filamentsas long as those at outer tipsof mouth-arms. Longest fila-ments are upon arm-disk.Entire
.
Network on inner side of ring-canal does not fuse with 16radial-canals.Narrow and cruciform
(8X7) those adjacent to rhopalariappets being twice as wide asthe others.4.Four wide, shallow grooves.About two-thirds as long as bell-diameter.A single slender filament at outerend of each arm. Other arm-filaments are very short andslender. No filaments upon arm-disk.Almost interrupted in 8 principaradii.Network on inner side of ring-canafuses with 16 radial-canals.Wide and cruciform.Genus MASTIGIAS L. Agassiz, 1862.Mastigias L. Agassiz, 1862, Contr. Nat. Hist. U. S., vol. 4, p.1910, Medusae of the World, vol. 3, p. 677. 152.
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Mayer,Generic Characters.?Ehizostomata triptera with 3-winged mouth-arms, which terminate in a naked, club-shaped extremity. Thereare also smaller clubs and filaments between the frilled mouths.
220 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.The mouths are developed not only along the edges of the three leaf-like wings of the lower parts of the mouth-arms, but also over partsof their flat, expanded sides. The central stomach gives rise to eightrhopalar-canals and numerous, interocular radial-canals, all ofwhich anastomose and finally connect with the ring-canal. Therhopalar-canals extend straight to the sense-clubs, but the 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-zoneand fuse with the outer ends of the rhopalar-canals. The ring-muscles of the subumbrella are interrupted in the eight rhopalarradii. There is a unitary subgenital porticus.MASTIGIAS PAPUA (Lesson).Cephea papua Lesson, 1829, Voyage de la Coquille, Zooph., p. 122, pi. 11,figs. 2, 3.Mastigias papua Agassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 162.
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Mayek, 1910, Medusae of the World, vol. 3, p. 678, fig. 415.?Light, 1914.Philippine Journ. of Science, vol. 9, p. 209.This common medusa is widely distributed over the Malay Archi-pelago, Indian Ocean, and China Sea to Japan, and outward overthe Pacific to the Fiji Islands. It gives rise to a number of varieties.Eight specimens, the largest 59 mm. in diameter, were found by theAlbatross in the Philippines. Two specimens are from SantiagoEiver, Pagapas Bay, Luzon, February 20, 1909; two, Cat. No. 28735,U.S.N.M., are from Pandanon Island, between Cebu and Bohol,March 24, 1909, and four, Cat. No. 28732, U.S.N.M., were taken onthe surface off Cebu on September 5, 1909. It was abundant in shal-low water among the mangroves in Port Moresby, Papua, in No-vember, 1913. MASTIGIAS OCELLATA (Modeer).Medusa ocellata Modeer, 1791, Nova. Acta. Phys. Med., N. C, vol. 8, Ap-pend., p. 27.Cephea ocellata Peron and Lesueur, 1809, Annal. du Mus. Hist. Nat., Paris,vol. 14, p. 361.Mastigias ocellata Haeckel, 1880, Syst. der Medusen, p. 623.
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Mayer, 1910,Medusae of the World, vol. 3, p. 680.The Albatross found small medusae of this variety in March andApril, and a mature one, Cat. No. 27919, U.S.N.M., in January inthe Philippines, in 1908. It is distributed over the eastern parts ofthe Indian Ocean and in the China Sea.Genus PHYLLORHIZA L. Agassiz, 1862.Phyllorhiza Agassiz, L., 1862, Cont. Nat. Hist. U. S., vol. 4, p. 158.
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Mayer,1910, Medusae of the World, vol. 3, p. 684.Generic Characters.?Similar to the closely allied Lychnorhiza,but the centripetal vessels which arise from the inner side of the
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 221
ring-canal join with the central stomach, as in Mastigias, instead ofending blindly as in Lychnorhiza. The canal-system resembles thatof Mastigitis, but the mouth-arms have no terminal clubs.PHYLLORHIZA LUZONI Mayer.Phyllorhiza luzoni Mayee, 1915, Publication No. 212, Carnegie Institutionof Washington, p. 194, fig. 7.Two specimens of this medusa were captured at a depth of 150feet in Varadero Bay, Southern Luzon, Philippine Islands, on July23, 1908. Cat. No. 28728, U.S.N.M. Unfortunately both becamesomewhat macerated in the preservative fluid.The bell of the larger and more perfect specimen is 60 mm. wide,flat, and with finely granular exumbrella. Eight rhopalia withoutocelli in formalin and without exumbrella pits. Sixteen scimeter-shaped rhopalar lappets, and (9X8)72 rounded velar lappets which aresomewhat wider but not longer thanthe rhopalar lappets, so that the gen-eral contour of the bell-margin iscircular, without deep niches in therhopalar radii.The arm-disk is cruciform, 32 mm.in perradial and 20 mm. in interradialdiameter. The subgenital ostia are14 mm. and the perradial columns8 mm. wide, and the subgenital por-ticus is wide and unitary.The eight mouth-arms are slenderand strongly compressed laterally, their lower parts being 3-wingedwith deeply incised lateral membranes. The naked outer part ofeach upper arm is 13 mm. and the 3-winged lower part only 11 mm.long. Any appendages which may have existed among the mouth-frills have disappeared in the preservative fluid, owing to the macera-tion of the specimens.The muscular system of the subumbrella consists of a broad, unin-terrupted zone of weakly developed circular muscles. In otherhitherto known species of Phyllorhiza the muscles are interruptedin the eight principal radii.The central stomach is cruciform, 32 mm. in perradial and 16 mm.in interradial diameter. Eight radial-canals arise from the stomachand extend straight to the eight rhopalia ; these main canals areconnected one with another by a ring-canal which is 6 mm. inwardfrom the margin of the bell. In each octant six to nine radiatingvessels arise from the cruciform stomach anastomosing in an irregu-lar network with one another and with the eight main canals. On
Fig. 20.
?
Phyllorhiza luzoxi fromVaradero Bat, Luzon.
222 BULLETIN 100, UNITED STATES NATIONAL MUSEUMits outer side the ring-canal gives rise to a fine-meshed network ofvessels which ramify through the lappets.A color note states that the medusa was light green with grayishwhite spots. Genus VERSURA Haeckel, 1880.Crossostoma preoccupied for rnollusks by Nokbis and Lycett, 1850.Crossostoma Agassiz, L., 1862, Contr. Nat. Hist. U. S., vol. 4, p. 155.Versura+Crossostoma Haeckel, 1880, Syst. der Medusen, pp. 606, 607.Versura Mayek, 1910, Medusae of the World, vol. 3, p. 685.Generic Characters.?Rhizostomata triptera with clubs and fila-ments upon the mouth-arms. The 4 perradial canals arise directlyfrom the stomach, but the 4 interradial canals result from the fusionof a number of anastomosing vessels which arise from the interradialsides of the stomach. There is no definite ring-canal, but merely amarginal network of vessels. There are no radial-muscles in the sub-umbrella, but the ring-muscles are well developed. Among the char-acters of minor importance, the subgenital ostia are wide openings,wider than the columns between them, and the sense-organs havea simple, exumbrella pit without radiating furrows. At the centerof the arm-disk is a prominent, raised cluster of frilled mouths hav-ing filaments between them.VERSURA MAASI Mayer.Versura maasi Mayer, 1910, Medusae of the World, vol. 3, p. 687, fig. 416.Named in honor of Prof. Dr. Otto Maas, in recognition of hisnotable researches upon Medusae. A single perfect specimen (Cat.No. 27943, U.S.N.M. Type) was obtained by the United States Fish-eries Bureau steamer Albatross on April 8, 1908, along the shore atMantacao Island, west coast of Bohol, Philippine Islands.Bell 90 mm. wide, flatter than a hemisphere and evenly rounded.Exumbrella finely granular without furrows. Gelatinous substancefairly thick but not very rigid. Eight rhopalia each with a pigmentspot and an exumbrella pit with smooth floor. Usually 12 marginallappets in each octant, with outer edges bluntly rounded. The 16rhopalar lappets are slightly narrower than the velar lappets. Armdisks five-ninths as wide as bell-diameter at the level of the origin ofthe eight mouth-arms. The four interradial subgenital ostia are twotimes as wide as the perradial columns of the arm-disk. Eight mouth-arms each one-third as wide as the bell-diameter. The three-wingedlower part of each arm is somewhat more than two times as long asthe unbranched upper part of the arm. The two lateral outer wingsof each arm are deeply cleft. (See fig. 21, C.) There are a largenumber of laterally flattened clubs at the center of the arm-disk.These are besprinkled with small nettle-warts which are most promi-nent at the broad outer end of the club. In addition there are a few
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 223
very small appendages among the mouths of the mouth-arms. Themouth-arms are strongly compressed laterally. A single duct extendsdown each mouth-arm and gives rise to four terminal branches, twoto the lateral wings and two to the triangular extremity of the arm.There are eight wide rhopalar radial canals, of which the fourperradial ones rise directly from the stomach ; but each of the 4 inter-radial arise from the confluence of a Y-shaped fork. Seven to ninenarrow radial vessels arise in each inter rhopalar octant and extendoutward anastomosing profusely with each other and with the eightrhopalar canals. There is no definite ring-canal.A wide, entire, annulus of circular muscles is found in the sub-umbrella, but there are no radial muscle strands. This muscular zone
v-Q^lCn-
Fig. 21.
?
Veesura maasi from Mantacao Island, west coast of Bohol. A, OralVIEW. B, A CLUB FROM THE MOUTH-ARMS, MUCH MAGNIFIED. C, A MOUTH-ARM SEENFROM THE OUTER (ABAXIAL) SIDE. D, EXUMBRELLA VIEW OF ONE OF THE MARGINALSENSE ORGANS.is wider in the interradii than in the perradii, and is somewhatthinned but still unbroken in the eight rhopalar radii.In formalin the rhopalar canals, and all vessels near the stomach,are bluish purple. The mouth-frills are brownish to brownish purple.The bell is milky and the gonads dull brownish yellow.Genus LOBONEMA Mayer, 1910; sensu Light, 1914.Lobonema Mayeb, 1910, Medusae of the World, vol. 3. p. 638.
?
Light.1914, Philippine Journ. of Science, vol. 9, p. 216.Generic Characters.?Rhizostomata triptera in which the marginallappets are greatly extended, tapering to pointed ends. These lappetsare noncontractile and lack muscles. Mouth-arms with numerousfilaments. Mouth-arm membranes perforated by windowlike open-ings. Eight to sixteen rhopalia and twice as many radial-canals, anda ring-canal which gives off anastomosing vessels on both its innerand outer sides. The inner network does not connect with the stom-
224 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.
ach. All of the radial-canals extend beyond the ring-canal. Thesubumbrella exhibits a well-developed system of ring-muscles. Thereare numerous, prominent, tapering papillae upon the exumbrella.There is a sensory pit on the exumbrella side above each rhopalium,and the floor of the pit exhibits radiating, dendritic furrows.Lobonema smithii, the first-known species and type of the genus, wasobtained by the Albatross in Manila Bay, Philippine Islands, late inApril, 1908. It is named in honor of Dr. Hugh M. Smith, Commis-sioner of Fisheries of the United States. Another species from Pala-wan was described by Light * under the name Lobonema mayeri.Lobonema smithii has only eight marginal sense-organs, and thesubumbrella ring muscles are entire; while in L. mayeri there are12 to 16 marginal sense organs, and the circular muscles are com-pletely interrupted in the ocular radii. Light, who studied themedusae in life, states that the colors of Lobonema mayeri are anexquisite scheme of purple, violet, and rose pink. The gonads are, asa rule, pink, the general color is violet, and the fringe of taperingmarginal lappets purple. Doctor Light tells me that in Lobonemasmithii these colors are not so brilliant, and the bell of the medusamay be white. Light finds that the sting of this medusa is not verysevere, and that the cases of poisoning reported by Old were prob-ably due to Dactylometra and not to Lobonema.Light describes Lobonemoides gracilis, an immature medusa whichmay possibly be a young stage of Lobonema mayeri. The marginallappets are pointed but are not very long, and there are no window-like openings in the mouth-arm membranes.LOBONEMA SMITHII Mayer.Lobonema smithii Mayer, A. G., 1910, Medusae of the World, vol. 3, p. 689,figs. 417, 418.The Albatross found this medusa in Manila Bay, Luzon, at theship's anchorage on April 25, 1908, and again (Cat. No. 28725,U.S.N.M. Type) on the surface at station D. 5222, between Marin-duque and Luzon. 9 miles off San Andreas Island.This species is named in honor of Dr. Hugh M. Smith, nowUnited States Fish Commissioner, who found it in Manila Bay,Philippine Islands. The Albatross found a perfect specimen of thismedusa, and a quadrant of its disk and all of its mouth arms werepreserved. There were also two other imperfect specimens, so thatall three taken together afford data for a partial description of themedusa.Bell flatter than a hemisphere, 236 mm. across from each sense clubto the one 180? from it. Gelatinous substance thick, tough, and rigid.Exumbrella regularly besprinkled with erect, gelatinous papillae
1 1914, Philippine Journal of Science, vol. 0. p. 217, figs. 7-9.
SCYPHOMEDUSAE COLLECTED BY STEAMER ALBATROSS. 225
which are largest and most abundant at the center of the exumbrellabut disappear near the margin and are not seen over the lappets.Near the center of the exumbrella these papillae are about 6 to 10mm. apart and each is about 35 to 40 mm. long and 3 to 5 mm. wideat the base ; they are conical, usually more or less curved, and taperto pointed ends.Their surfaces arethickly covered withnematocysts, whichgive a bristling ap-pearance to the diskof the medusa.Eight rhopaliawhich lack ocelli inspecimens preservedin formalin or alco-hol. On the exum-brella side aboveeach sense club thereis a shallow, heart-shaped, sensory pitwith dendriticridges over its floor.The rhopalia areflanked by verysmall, oval, ocularlappets only 3 mm.long and 2.5 mm.wide. There are 32(4X8) velar lap-pets, which are mostextraordinary, eachbeing 90 to 100 mm.long and taperinggradually from baseto tip. They aremodified so as to re-semble superficiallytentacles of semaeostomous Scyphomedusse. They trail downwardfrom bell margin, waving flexibly to and fro, as do veritabletentacles. I can find no muscles in these lappets, however, andLight confirms the statement that they can neither contract norelongate. They are deep clefts in the exumbrella surface betweenthe lappets, but these clefts are bridged over by a thin subum-101825??Bull. 100?17 15
Fig. 22.
?
Lobonema smithii. A, Diagram showing the planOP STRUCTURE OP THE MOUTH-ARMS. B, SIDE VIEW OF ONEOF THE MOUTH-ARMS. C, RHOPALIUM AND ONE OF TBELONG, TAPERING MARGINAL LOBES.
226 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.brella membrane spanning between the lappets. The eight inter-rhopalar grooves are 35 mm., the 8 rhopalar 16 mm., and the 16intermediate clefts 31 mm. long. The eight rhopalar clefts areU-shaped and the exumbrella sensory pit is at the middle of thecrotch of the Y with the divided groove on either side of it (see fig.22, C) . The grooves between the velar lappets are simple, undivided,linear clefts.Sixteen radial canals, eight rhopalar and eight inter-rhopalarleave the central stomach, and all extend to the bell margin. There isa fairly distinct ring canal about 30 mm. inward from the senseclubs, and this ring canal gives rise on both its inner and outer sidesto an anastomosing network of vessels which connect with the 16radial canals but not directly with the stomach. This network of
FlQ. 23. LOBONEMA SMITHII FROM MANILA BAT. PART OF THE OUTER ZONE OF THESUBUMBRELLA, SHOWING CANAL SYSTEM AND MUSCLE STRANDS.
vessels extends downward throughout the length of the taperinglappets trending mainly longitudinally but with frequent anas-tomoses.The muscular system forms an annulus about 68 mm. wide in thesubumbrella from the margin of the arm disk to the zone of therhopalia. The circular muscles are powerfully developed and areonly thinned but not broken in the rhopalar radii. There are noradial muscles and no muscles in the lappets (fig. 23).The arm-disk is 100 mm. wide, but as it was cut off I can make nostatements in reference to the size or form of the subgenital ostia orof the gonads.The eight mouth-arms are separate, 150 mm. long, and each is 3-winged below. The upper shaft of each arm is 60 mm. and the3-winged lower part 90 mm. long. It is remarkable that each of thethree lateral membranes is perforated by three windows or openings(see diagram A, fig. 22). The axial duct of the arm extends down
SCYPHOMEDUSAE COLLECTED BY STEAMER " ALBATROSS." 227the center and gives off side branches in the tissue between the win-dows to the mouths. These side branches are joined one to anotherby longitudinal canals near the frilled mouth (see fig. 22, B).There are numerous appendages upon the mouth-arms arising be-tween 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 moreslender, and above these there are mere threadlike filaments. Theappendages are usually 70 to 100 mm. long, and the large ones con-tain an axial duct. The general color of the medusa in formalin ismilky-gray, the mouths and gonads being darker than other parts.Genus THYSANOSTOMA L. Agassiz, 1862.Thysanosioma Agassiz, 1862, Contr. Nat. Hist. U. S., vol. 4, p. 153.
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Mayer,1910, Medusae of the World, vol. 3, p. 691.Generic Characters.?Rhizostomata lorifera having mouth-armsbearing three rows of frilled mouths from base to lower end, withouta terminal club. Among characters of minor importance, the fourinterradial, subgenital ostia are wider than the perradial columnsbetween them. There are eight rhopalar canals and a ring canalwhich gives off a network of vessels on both its inner and outer sides.This network connects with all the radial canals and also at numerouspoints with the central stomach. The well-developed circular musclesare only partially interrupted in the eight principal radii. There isa small, shallow, exumbrella pit above each sense organ, and thereare no furrows in the floor of the pit. The only difference betweenthis genus and the closely allied Lorifera is that the frilled mouthsare developed even to the tips of the lower ends of the mouth-armand there is no terminal club, whereas Lorifera has a naked terminalclub. THYSANOSTOMA THYSANURA Haeckel.(?) Rhizostoma brachyura Lesson, R. P., 1829, Voyage de la Coquille,Zoophyt., vol. 2, p. 153 ; 1830, Centurie Zoologique, p. 227, pi. 80.Thysanostoma Agassiz, 1862, Contr. Nat. Hist. U. S., vol. 4, p. 153, figs.1-9.?Mayeb, 1910, Medusae of the World, vol. 3, p. 692, fig. 420.The dimensions, in millimeters, of a specimen (Cat. No. 27929,U.S.N.M.) obtained at Mindanao, Philippine Islands, by the UnitedStates Bureau of Fisheries steamer Albatross are as follows: Bell,100 wide; perradial diameter of arm-disk, 74; diameter of arm-diskat 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 thearm-disk, 10 to 15 long ; exumbrella finely granular.In another large medusa (Cat. No. 27930, U.S.N.M.) from Mansalay,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-
228 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.arms 190 mm. long. In a half-grown medusa (Cat. No. 27928,U.S.N.M.) obtained on the surface at the same time and place thebell is 59 mm. wide with finely granular exumbrella; mouth-arms,67 mm. long; arm-disk, 41 mm. wide at its origin from the subum-brella and 33 mm. wide at the level of the origins of the mouth-arms.Two specimens (Cat. No. 28710, U.S.N.M.) are from AtulayanBay, east coast of Luzon, June 17, 1909. Of these the larger one wascut into two pieces when viewed by me, but the bell appears to havebeen about 106 mm.in diameter, themouth - arms being167 mm. long, theupper arm 20 mm.,and the lower arm147 mm. long.A color note leadsone to infer that inlife the bell wastranslucent with aslightly brownishmargin, and withthe frilled mouthsdark burnt -umber.The smaller speci-men when alive hada translucent bellwith some whitishspots and with eightindigo streaks alongthe eight principalradii. The frilledmouths and bell-margin were pale brown. One specimen (Cat. No. 28711, U.S.N.M.)91 mm. in diameter with mouth-arms 155 mm. long, is from StationD2268, September 21, 1909, depth 13 fathoms off Singaan Islandnorth of Tawi Tawi. In this the arm-disk is 59 mm. in perradialdiameter, and the upper arms are 13 and the lower 142 mm. long.Genus LORIFERA Haeckel, 1880.Himanostoma preoccupied for Diptera by Loew, 1853.Himanastoma Agassiz, L., 1SG2, Contr. Nat. Hist. U. S., vol. 4, p. 152.Lorifera Haeckel, 18S0, Syst. der Medusen, p. 028.
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Mayer. 1910, Medusaeof the World, vol. 3, p. 693.Generic Characters.?Rhizostomata lorifera in which the eightmouth-arms bear rows of frilled mouths throughout their lengths but
Pig. 24.
?
Thysanostoma thysanura from PanabutanBay. A, Oral view of the 8 mouth-arms being re-moved. B, Cross section of mouth-arm near thebase. C, Cross section of mouth-arm near the outerEND OF THE ARM. D, ORAL VIEW OF RHOPALAR LAPPETS.E, Rhopalar lappets from the exumbrella side.
SCYPHOMEDUSAE COLLECTED BY STEAMEK " ALBATROSS." 229terminate each in a naked knob. Among characters of minor im-portance the subgenital ostia are usually wider than the perradialdisk-columns. The circular muscles of the subumbrella are prac-tically entire. The sense clubs have each an occellus and a well-developed exumbrella pit with radiating furrows. This genus is dis-tinguished from the closely allied Thysanostoma only by the naked,club-shaped extremities of its mouth-arms.LORIFERA LORIFERA, var. PACIFICA (Schultze).Himanostoma loriferum, var. paciflca Schultze, L. S., 1897, Abhandlung,Senckenberg, Naturf. Gesell., vol. 24, Heft 2, p. 153, pi. 15, figs. 1, la, 6
;
1898, Denkschr. Med. Nat. Gesell. Jena, vol. 8, p. 446, pi. 34, fig. 9 (youngmedusa).Lorifera lorifera, var. paciflca Mayek, 1910, Medusae of the World, vol. 3,p. 695.A single half-grown specimen of this medusa, Cat. No. 28709,U.S.N.M., was found at Port Palapag, Luzon, by the United StatesFisheries Bureau steamer Albatross on June 2, 1909, being capturedthrough the use of dynamite. Its bell is 98 mm. wide, exumbrellafinely granular, flatter than a hemisphere. The bell margin is badlydamaged but there appear to be six to eight cleft or subdivided velarlappets in each octant. Their general contours are rounded, andinterlobular clefts extend radially a short distance up the exumbrellasurface.The eight rhopalia have each an ocellus, but no exumbrella pit.Perradial diameter of arm disk, 55 mm.; width of subgenial ostia,34 mm. ; width of perradial columns of arm disk, 12 mm. ; subgenitalporticus wide and unitary ; naked upper arm, 20 mm. long ; length ofthe abaxial side of the mouth-bearing part of each arm, 35 mm.;length of the terminal naked axial filament of each arm at least40 mm., but all are broken at their ends.There is a thick felting of short filaments upon the arm disk.The circular muscles are unitary, being only partially interruptedin the eight principal radii.The eight radial canals are each about 3.5 mm. wide, and in addi-tion there are about 8X10 slender radial canals, all of which anasto-mose with one another and with the eight chief radial canals, form-ing a wide network in the subumbrella.In formalin the eight chief radial canals are violet.Schultze describes this medusa from Ternate and from Amboina,Malay Archipelago.
230 BULLETIN" 100, UNITED STATES NATIONAL MUSEUM.TABULAR DESCRIPTIONS.Table showing the chief characteristics of Philippine Scypho-medusae. This table is designed to enable one to recognize eachspecies from its most conspicuous features, but it must be used withcaution, and is worthless for the medusae of regions other than thePhilippines. Family CARYBDEIDAE.Bell high, pale milky colored, more or less rectangular. Foursmall niches on the sides of the bell alternating with the tentacles.A minute club, bearing several eyes, is set within each niche. Thetapering flexible, pale pink sharply stinging tentacles arise from theends of stiff, spatula-shaped projections from the rim of the bell.1. Bell about 13 inches high, only slightly higher than it is wide. Four ten-tacles Carybdea rastonii.2. Bell about 9 inches high, about twice as high as wide. Four tentacles.Carybdea alata, var. grandis.3. About 28 tentacles arising from the ends of four stiff finger-shaped clustersof projections from the rim of the bell Chiropsalmus quadrigatus.Family CORONATE.These medusae have a deep constriction, or ring-furrow, cuttinginto the sides of the bell.1. A deep-sea medusa, purple-brown or claret colored. Usually about 2 to 6inches high. Sixteen marginal lobes, 12 tentacles. Bell conical or dome-like with a deep ring-furrow extending around about one-third the wayup the side of the bell Periphylla hyacinthina.2. Bell thimble-shaped, i an inch high, brown with darker colored warts on theconcave side. Swims very actively, with an incessant pulsation, near thesurface in great swarms Limiche unguiculata.3. A deep-sea medusa with flat, thick lenticular bell usually about 1$ to 4inches wide; the central part separated by a deep constriction from thenumerous lappets. Numerous short tentacles alternate with the equallynumerous cleft lappets. Deep brownish red lappets and mouth parts.Atolla bairdii, etc.Family SEMAEOSTOMAE.Usually large medusae, with marginal tentacles. Without a ring-furrow in the bell. With a single cross-shaped mouth at the middleof the concave side of the umbrella. Often seen in harbors.1. Animal blue-violet with purple streaks on the outer side of the bell, eightlong, tapering tentacles, and four long, flexible curtain-like lips. Bell about2 inches wide, and lips 3 inches long Pclagia panopyra.2. Sixteen areas of reddish-rosin colored dots radiate outward in a spokelikemanner from the center to the bell margin. Twenty-four tentacles and 48marginal lobes (16 small and 32 large). Bell about 5 inches wide.Chrysaora melanaster.
SCYPH0MEDU8AE COLLECTED BY STEAMER "ALBATEOSS." 231(3 and 4.) Resemble Chrysaora melanaster, but there are 40 tentacles.3. Marginal lobes not highly colored Dactylometra quinquecirrha.14. Marginal lobes red Dactylometra africana.5. Resembles Dactylometra (3) in general appearance, but the color is moreyellowish, and there are only 16 tentacles. Bell about 4 inches wide.Thirty-two cleft marginal lobes Sanderia malayensis.6. Bell only about \ inch wide, translucent, 24 tentacles, 32 marginal lobes, 8simple radial canals, alternating with 8 complexly branching canals on theconcave side of the bell Discomedusa philippina.7 and 8. Bell genatinous, about 10 inches wide or larger, hyaline or slightlymilky, with 4 whitish or pinkish horseshoe-shaped genital organs near thecenter of the bell. 4 long, fleshy lips. More than 100 short tentacles.8 simple and many branched and anastomosing canals all of which areconspicuous, due to their milky color.7. 8 clefts in the bell-margin Aurellia aurita.8. 16 clefts in the bell-margin Aurellia laoiata.Family RHIZOSTOMAE.Usually large medusae without marginal tentacles. With numer-ous minute mouths, the frilled tentacle-bearing lips which super-ficially resemble sea weed. Mouths born on 8 more or less branchedprojections (mouth-arms) arising from the center of the concaveside of the bell. Common in harbors in the East Indies. The gelatin-ous substance of the bell is often remarkably rigid. The branched,frilled, and tentacle-bearing mouth-arms are often greenish or red-dish in color, thus giving the appearance of some sort of vegetablegrowth arising from the center of the concavity of the bell.Living on the bottom, slowly pulsating, " lying on its back," withthe concave side of the bell, and its 8 mouth-arms, and mouths upper-most; remaining for hours, or days, fixed at one place and rarelyswimming through the water Cassiopea.1. Bell flat, about 4 inches wide, without a concavity where it lies upon theground. Olive-brown in color with about 18-22 triangular large whitishspots in the radii of the marginal sense organs. 3 small white streaks nearthe bell margin between the large triangular spots. 8 bifurcated mouth-arms Cassiopea andromeda, var. oaduensis.2. Bell olive green about 6 inches wide with a suckerlike concavity where ittouches the ground. Dull inconspicuous, white bands radiate outward inthe radii of the 16 or more marginal sense organs. The leaflike append-ages of the mouth-arms are opaque white.Cassiopea polypoides, var. culionenus. 13. Bell about 11 inches wide. Numerous large ribbonlike appendages on themouth-arms, 4? inches long Cassiopea medusa.'Large free swimming medusae a foot or more wide with a more orless conspicuous dome covered with wart-like projections at the center
1 See S. F. Light, 1914, Philippine Journal of Science, vol. 9, section D, p. 198.2 Idem, p. 201.3 Idem, p. 204.
232 BULLETIN 100, UNITED STATES NATIONAL MUSEUM.
of the convexity of the bell. The dome may be sky blue, rose-colored, or amber colored and the bell blue with reddish spots andmarkings. Mouth-arms fused forming an eight-sided prism, flaringoutward below. Gelatinous substance tough Cephea.1. Dome low, warts small Cephea octostyla.2. Dome high, warts conspicuous Cephea cephea.Bell amber colored, about 9 inches wide, with a smooth centraldome without wartlike projections. Clusters of grapelike append-ages on the eight leaflike mouth arms. Windowlike openings inthe membranes of the mouth arms. Marginal lappets of the bellshort and oval Cotylorhiza pacifica.Bell smooth, hemispherical, about 8 inches wide when full grown.Deep uniform, purple-brown. The commonest medusa in brackishwater canals in shallow water along shore in Manila Bay in No-vember and December Catostylus purpurus.Similar to the above but with irregular purple brown spots overthe convex side of the bell. No filaments on the mouth arms.Catostylus townsendi.Similar to Catostylus townsendi but with numerous long filamentson the mouth arms. The longest ones being near the outer endsof the mouth arms Acromitus maculosus. 1Bell high, domelike, 4J to possibly 12 or more inches wide, slightlygranular on the convex surface. Yellowish-white or cobalt blue.Eight tapering mouth arms without filaments or other appendages.Common in the mouth of Brisbane Eiver, Queensland, Australia, butwidely distributed over this general region Catostylus mosaicus.Bell about 3? inches wide, smooth, evenly rounded, flatter thana hemisphere, milky-yellow, with smooth convex surface. Slendertapering filaments in the eight mouth arms. No prominent spots inthe convex surface of the bell Lychnorhiza,1. A single long filament at the pointed end of each mouth arm. All otherfilaments very short Lychnorhiza bornensis.2. With numerous long filaments on the mouth arms Lychnorhiza bartschi.Bell smooth, about 2? to 3^ inches wide. With eight mouth arms,each tapering to end in a long naked more or less club-shaped fila-ment. This small medusa pulsates and swims rapidly, is usuallygreenish or reddish brown, and is common in harbors in the EastIndies.1. Bell blue, greenish, olive, or brown with solid white, brown or yellowish,spots. The eight terminal clubs on the mouth arms each as long as thediameter of the bell Mastigias papim.2. Bell reddish brown, with ringlike spots of white and brown. Eight terminalclubs only about half as long as the diameter of the bell?Mastigias ocellata.i See S. F. Light, 1914, Philippine Journal of Science, vol. 9, section D, p. 212, fig. 4.
SCYPHOMEDTJSAE COLLECTED BY STEAMER "ALBATROSS." 233Bell fiat, about 2^ inches wide, light green with grayish whitespots. Eight laterally flattened mouth arms, each ending in threeexpansions, eight simple radial canals in the radii of the eight mar-ginal sense organs and a network of vessels between these canalsand on both sides of the ring canal. No appendages on the moutharms. Phyllorhiza luzoni.Similar to Phyllorhiza luzoni in general shape, but there is noring canal and there are numerous small clubs upon the eight moutharms, especially at the center of the disk Versura maasi.Bell 15-20 inches wide, flat topped and thickly covered above withflexible tapering projections, the largest near the center. The mar-ginal lobes are greatly extended, forming tapering filaments. Themouth arms are perforated with windowlike openings and bearnumerous long filaments. Common in Harbors, Manila, and westcoast of Palawan in April and May-1. Twelve to 16 marginal sense organs and 70-80 long tapering marginal lobes.Purple, violet, and rose pink are the prevailing colors of the animal.Lobonema mayeri.12. Only eight marginal sense organs and 32 marginal lobes. Colors not so brightas in L. mayeri, and the bell may even be white Lobonema smithii.Bell 4 or 5 inches wide. Eight very long, " snakelike," mouth armswith frilled " seaweedlike " mouths all down their sides even to thetips of the arms Thysanostoma thysanura.Similar to Thysanostoma thysanura, but the eight arms end eachin a naked club Lorifera lorifera.
1 See S. F. Light, 1914, Philippine Journal of Science, section D, vol. 9, p. 217, figs. 7-9.o