NOAA Technical Report NMFS Circular 397
Marine Flora and Fauna of the Northeastern United States. Cnidaria: Scyphozoa
Ronald J. Larson
August 1976
U.S. DEPARTMENT OF COMMERCE Elliot l. Richardson, Secretary National Oceanic and Atmospheric Administration Rob\~rt M. White, Administrator National Marine Fisheries Service Robert W. Schoning, Director
t'ot Sale by the Superintendent of Documents. U.S. GO'o'crnment Printing Office Wo,hington. D.C. 20~02 - Stock No. 003.017-00387 ,'., NOAA FOAM 25-13 U. S. DEPARTMENT OF COMMERCE 12-73) BIBLIOGRAPHIC DATA SHEET NATIONAL OCEANIC AtJD Al'MOSPHERIC ADMINISTRATION 1. NOAA ACCE:;SION NUMBER 2 3 RECIPIENT'S ACCESSION NUMBER NOAA-7610l8l5 1 • 1 • 4. TITLE AND SL,'ST:TLE S. REPO~T DATE Marine Flora and Fauna of the North~astern United States. Au!! 1976 Cnidaria: Scyphozoa 6.
7. AUTHOR!SI 8. REPORT NO. Ronald J. Larson CU.S. National Museum, Washington. DC) NOAA-TR-NMFS Circular ~~7 9. PERFORMING ORGANIZATION NAME AND ADDRESS 10. PROJECT/TASK NO. NOM, National Marine Fisheries Service, Seattle, WA, 181 CJ S' 11. CONTRACT/GRANT NO. Scientific Publications Staff
12, SPONSORING ORGANIZATION NAME AND ADDRESS 13. TYPE OF REPORT AND PERIOD Same COVERED 1'~~h Rent' 14.
15. PUBLICATION REFERENCE NOAA Technical Report NMFS Circular 397, August 1976. 21 p, 28 fig, 24 ref.
16. ABSTRACT This manual is an introduction to the scyphomedusae found in coastal waters from
Maine to the Chesapeake Bay. It includes a discussion of their identification, ~- collection, rearing, preservation, and nematocysts. Also included is an introduction to the natural history of the scyphopolyps and medusa,e, a discussion of stinging scyphomedusae, a glossary of terms, an illustrated synopsis of ephyrae, an illustrating key to the scyphomedusae (including the Stauromedusae), an annotated systematic list, a bibliography of major refer~nces, and an index. (Author)
17. KEY WORDS AND DOCUMENT ANALYSIS
17A. DESCRIPTORS
Coelenterata, Jellyfish, MaTlua1s, Taxonomy, Invertebrates, Descriptions
11e. IDENTIFIERS/OPEN-SNOED TERMS
Cnidaria, Scyphozoa, Stauromedusae, Scyphistomae, Ephyrae, Scypr.omedusae, Sea net'tles, Sea wasps, Keys, U.S. Northeast Coast, Systematic lists
!7C. COSATI ~IE:::LD/GROUP
6C
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Thj~ isslle of the "Circulars" is part of a subseries entitled "Marine Flora and Fauna uf the Northeastern United States." This subseries will consist of original, illustrated, modern manuals on the identification, classification. and general biology of the estuarine and ~'Oastal marfine plants and animals of the Northeastern United States. Mcnuals will be published tlt irregular intervals on as many taxa of the 'egion as there are spp-.:ialists available to collaborate in their preparation. The:nanuals are an outgrowth of t:.e widely used "KeyS to Marine Invet1ebrates ofthe Woods Hole Region," edited by R. I. Smith, published in 1964, p.nd produced und'er the auspices of the Sys tematics-Ecolo~ Program, ~,hrine Biological Laboratory, Woods Hole, Mass. Instead of revising the "Woods Hole Keys," the slaff of the Systematics-Ecology Program decided to expand the geographic coverage and bathymetric ra"ge p.nd produce the keYf in 'In entirely new set of expanded pub. lications. The "Marine Flora and Fauna orthe Northeastern United States" is being prpareu in, :ollaborati':m with systematic specialists in the United States dnd abroad. Each manual will be based primarily on recent and ongoing revisionary systematic ,'eser,rch and a fresh examination of tht plant~ and animals. Each major taxon, treated in a separst,; manual, will include an introduction, :Ilustrated glossary. uniform originally i'lustrated keys, annotated check list with information when available on distribution. habitat, life history. and re!at'ld biology, references to the major literature oithe group, Bnd a systematic index. These manuals are intended for use by biology students. hiologists, biological oceanographers, in fermed l~ym~n, and others wishing to identify coastal organisms for this region, In many instances the malluals will serve as a guide to additional information about the species or the group. r.eogra»h.c coverage of the "Marine Flora and Fauna of the Northeastern United States" is plan. p"d to ir.c1ude organisms from the headwaters ofestuaries seaward to approximately the 200·m depth .;., the c:mtinental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the inter~sts of collaborators. Whenever possible representative specimens dealt with in the manuals will be deposited in the reference collections of major museums in the region. After a sufficient number Df manuals of related taxonomic groups have been published, the manuals will be revised. grouped, and issued as special volumes. These volumes will thus consist of compilations of individual manuals within phyla such as th:l Cnidaria, Arthropoc\a, and Moiiuse.., or of groups of phyla. CONTENTS
Introduction . .. . . Description of orders of 3cyphozoa . Methods Collecting .. .j Rearing ... ·l Pre3crvation ~ Nematocysts ... .j Nalural historv Stauromedusae ~ Scyphistomac . 4 Ephyrae .... 4 Scyphomedusae ~ Stinging medu!;ae . i) Glossary .... i) introduction to the keys to tho.! Scyphozoa £, Kev tn the Stauromedusae of the northeast coast of the United States fi Sy~opsis of ephyrae of Chrysaora, Cyanea. and Rhopi/ema . H Key to the pelagic; scvphomedusae of Ihe northeast coasl of the United States .10 Annotated systematic list . 14 Selected bibliography .. .W Index to scientific names .17 Acknowledgments .... IR Coordinating eJitor's comments .IR
The National Marine Fisheries Service (NMFS) does not approve, rec ommend or endorse any proprietary product or proprietary materiai mentioned in this publication. No reference shall be made to NMFS, or to thb publication furnished by ~MFS, in any advertising or sales pro motion which would indicate or imply that NlIIFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS pul>1ication. Marine Flora and Fauna ofthe Northeastern United States. Cnidaria: Scyphozoa
RONALD J. LARSON'
ABS'l"RACT
This manual is an introduction to the sC}llhomedusae found in coastal walers rrom Maine to the Chesapeake Bay. It includes a discussion oCtheir identifi~ation., collection, rearing, preservation. and nematocysts. Also included is an introduction to the natural history of th" scyphopolyps and medus.e, s discussion ofstinging 8cYDhomedusac, n glo8~ary or terms, an illustrated synopsis 01 ephyrBe, 1\'D'il· lustrating key to the scyphomoousae (including the Stauromedusael, an annotated systematic list, a bibliography of major references, and finally an index.
INTRODUCTION Scyphomedusae are generally regarded as pests because of their irritating stings, but many are The class Scyphozoa of the phylum Cnidaria or ecologically important. Chrysaora, well known as the sea Coelenterata as it once was called, are commonly known nettle, consumes large numbers of ctenophores which as "jellyfish," "sea nettlt-s," and "sea wasps." might otherwise be very detrimental to oyster and clam Scyphomedusae, as these jellyfish are termed, are the populations by feeding on their larvae. Other most conspicuom of the jellylike animals. Some species scyphomedusae prey on jellyfish which have few reach 1 m or more in diameter and most are very color predators and feed on commercially important invl'r ful. All have stinging organs, hence the names "sea tebra(~ and fish larvae. nettle" anct "sea wasp." 8cyphomedusae are seasonal; FiVl' ;,,'ders of Scyphozoa are recognized. The most species are seen during the warmer months. They Stauromedut·.:. or "stalked jellyfish," are not well represent the sexual stage of most scyphozoan species. know,' even ~hough they may be abundant. They are in The asexual stage is a small benthic (bottom-living) active and cryptically pigmented and are difficult to see po:yp (scyphopolyp) wh;,:h is perennial. The when attached to algae. The Cubomedusae, or sea wasps, scyphopolyp generally buds larval scyphomedusae are also infrequently seen but arc infamous because of (ephyrae) during the spring. The medusae are large by their potent sting. The Coronatae, or "crown medusae," midsummer. are mostly deepwater (bathypelagic). The SCYP!:lOmedusae have a muscular saucer-shaped or Semaeostomeae, commonly known us sea nettles, are the hemispherical bell or umbrella which propels the most fam.iliar order, and most shallow-water medusae through the ','!ilter by contracting and expelling scyphozoans belong to this group. The Rhizostomeae, water behind. Long threadlike tentacles often hang from which lack tentacles, are mostly tropical and are un the umbrella and are covered by stinging organs common in New England .....aters. (nematocysts) which are us"d to capture prey. Sur roul1.~ing the mouth are ribbol'.like, curtainlike, or gelatinous arms which transport prey to the mouth. DESCRIPTION OF 8cyphopolyps are minute !lnd rarely seen. They are ORDERS OF SCYPHOZOA saclike with a circle of tentaclr;s around the oral end and Order Stauromedusae-Small benthic scyphozo'lns are attached to the bottom by a stalklike peduncle. which lack a free-swimming medusa stage. Stauromedu Seyphozoans usually have a scyphomedusa and a sae are urn- or funnel-shaped and are attached to the scypht)polyp stage in their life history but one group substrate by a Dtalklike peduncle. In some forms, the remains as a'polyp only. The 8tauromedusae or stalked margin is divided into eight arms, the knobbed tentacles medusae lack a medusa stage. 80me scyphomedusae project from the arm tips; in others the tentacles project lack a polyp stage; Pelagia, an oceanic medusa, has eggs from the margin. Modified tentacles, which may act as which, when fertilized, transform directly into a juvenile anchors or sense organs, occur on the margin hetwpA'1 the medusa and bypass the normal polyp stage. groups of tentacles in some species. The pnads ;,re paired, leaflike, or folded structures extending along t.:~ length of the calyx. A wl)rmlike planula larva develops from the zygote and creeps ever the substrate, even 'Department of Marine Sdence,. University or Puerto Rico. Mayal!Uez, Puerto Rico 00708; present address: Division of Echinoderms, U.S. tually attaches itself and becomes a juvenile National Museum, Washir,gton, D.C. 20560. stauromedusa. Stauromedusae occur mainly in cold, shallow marine waters. Adult Stauromedusae usually oc brella near the margin. The mouth is at the end of a cur only during the summer, and the wintering juveniles flask-shaped stomach which hangs within the umbrella. may be difficult to find because of their small size. The life cycle of the Cubomedusae is unlike that of other Tropical and deepwater forms are rare. scyphozoans. The cubopolyp has several features in com Order Cubomedusae (Fig. I)-Small- to medium-sized mon with hydrozoan polyps, e.g., solid tentacles, steno medusae with & somewhat cuboidal or rectangular tele nematocysts. Cubopolyps metamorphose directly shaped transparent umbrella. A velarium, or shelflike and totally into a single medusa, other scyphopolyps projection, encircles the inner margin of the umbrella. undergo transverse fission producing one to many ephy Tentacles vary from four to many and are on bladelike rae. Werner (1973) has placed the Cubomedusae in ~,;,·~ir projections of the bell, termed pedalia. Four rhopalia are own class, Cubozoa, because of this group's unique char located about midway between the corners of the um- acteristics. Cubomedusae are tropical or subtropical me-
/b.,,_~~...----gastric
1l!I'+----gonad------.H-!1fI
, /pedalia----..
velarium
2
gastric cirri
\------~rt_oralarm-----
3
~;gure l.-Cubomedusa. Figure 3.-Semaeoslome medus•• Figure 2.-Coronalemedu68. FilCW'O 4.-RWzostome med.us", 2
c-.:,',-. .j. dusae but are occasionally transported inti) temperate polyp stage is small and naked or has a thin cuticle regions by warm currents. Cubomedusac ure known as around its base and can encyst under certain conditions sea wasps because of their virulent sting. /Fig. 7). A typical ephyra (Fig. 5) is produced, usually by Order Coronatae (Fig. 2)-Small- or medium-sized polydisc strobilation (Fig. 9). Semaeostomes Ilrc the medusae with flattened or dome-shaped umbrella which most common scyphomedusae in temperate coastal is divided into two regions by a circular (coronal) furrow. waters. The tentacles occur between marginal lappets. Radial Order Rhizostomeae (Fig. 4)-Large medu~ae, usually thickenings of the margin, termed pedalia, I'ive rise to with a hemispherical umbrella. Marginal tentacles are the tentacles, lappets, and rhopalia. The gonads are on ab~ent, but eight oral arms serve both to capture and four septa which project into the coelenteron. The transport food to the ge.stric cavity by Nay of a canal Coronatae are mostly bathypelagic and darkly pig· system. A central mouth is usually absent; instead the mented. The coronate polyp, all known species being 11lllut h is formed by a large number of bifurcating grooves placed in the genus Stephanoscyphus, is covered by a which lie on the ventral and lateral sides of the oral arms. chitinous cuticle wpich has numerOlIS circular an The oral Elrms may have terminal appendages which nulations (Kramp 1959). have a high concentration of n£>matocysts at their tip. Order Semaeostomeae (Fig. 3)-Large medusae, The glmads e,re foids of the gastrodermis as in the usually saucer-shaped. The mouth has four long oral sel11ucostome~;. The polyp stage has a long stalklike arms. These may be folded once as in Pelagia, or folded a peduncle ',vhich is partially covered with a thin cuticle. number of times as in Cyanea. Tentacles occur between J<:phyrae ,Irf, usually produced by monodisc strobilation marj{inal lappets as in the Coronatae, or on the subum (Fig. 10). Rhizostomes are mostly tropical, with only a hrella. Gonads occur on folds of the gastrodermis. The few species extending into temperate waters.
rhopolium
cirri
mouth
5 6 7
gastric
8 9 10
Figure 5.-Ephyra, newly released. Figure 7.-Semaeostome polyp. Figure 9.-Semaeostome slrobila. Figure H.-SfepMnoscyphus. Figure 8.-Stauromedusa. Figure 10.-Rhi,oslorne slrobila. 3 METHODS with crustaceans being the major part of the food. Berrill (1962) noted that Lucernaria fed on the amphipc>d Collecting Amphithoe and the small gastrupod Lacuna. At Woods Hole, Mass., I found mostly harpacticoid copepods and To obtain medusae in the best possible condition they shadd be dipped from the water surface usinp.: a bucket gammarid amphipods, and a few chironomid fly larvae in or other large container. Deepwater species can be gastric cavities of Craterolophus. Prey, caught by the caught in plankton nets equipped with large sample tentarles, cause one or all arms to fold over the mouth uuckels. which then expands and engulfs the food. Scyphopolyps can be obtained by dredgin!:, collecting Stauromedusae, like scyphistomae, occur year-round. at low tide, or diving. Polyps commonly attach to shells, Berrill (1962) found that Stauromedusae, along the coast oysters being a favorite substrate. Polyps will also settle of Maine, spawn during the summer and then die. on "lass slides placed in areas where medusae spawn. .Juveniles appear during the fall and become sexually mature by summer. Rearing Apparently Stauromedusae are very sensitive to en· vironmental CO'lditions and have been difficlllt to raise in Scyphozoans can be raised from embryos to maturity aquaria. Berri11 (1962) remarked that they are becoming in the laboratory (Spangenberg 1965). Polyps can be rare in Massachusetts Bay due to pollution. reared in small dishes and fed newly hatched Artemia. Planktoni<: .. cages are more difficult to raise, requiring a Scyphistomae large volume of circulating water and suspended food. Medusae should be fed daily, preferllbly more often. Scyphistomae are the polyp stages of scyphomedusae; Artemia can be fed to such plan:
NATURAL HISTORY Ephyrne Stauromedusae Ephyrae (Fig. 5) are small, 1-4 mm in diameter, flat tened larv,,1 medusae which usually have eight arms. At Stauromedusae, contrary to what their name implies. the tip of each arm there is a pair of blunt or pointed are really polyplike. Some are permanently attached. lappets, and between the lappets, a rhopalium or sense Others can move in a somersaulting motion by adhering organ. The mouth is cruciform and a few gastric cirri to the substrat.e with the oral eO!d and releasing the pedal may occur in the coelenteron. Ephyrae feed on plankton disc, then reattaching the disc at a new location; but and grow rapidly. They soon appear like small scypho· none have been observed to swim. medllse.e, at which time they can usually be identified They attach to algae, sea grass (Zostera), and other with certainty. (See synopsis of Ephyrae.) substrates in shallow areas which have adequate water circulation. They may be abundant, but because of their Scyphomedusae inconspicuous coloration and inactivity, they are rarely seen. Most of the diet consists of other benthic animals, By being planktonic, scyphomedusae may find food, 4 which is unavaHable to the polyp, and can also di,'· small. Chrysaora reared by the author began r-pawning at tribute gametes over an extenilive area thereby in· the size of55 mm and spawned almost nightly thereafter. creasing the distribution of the species and also insuring Fertilization usually takes place within the gastric cavity tllat uninhabited substrates are utilized if conditiom are or on the surface of the ovary. In Cyanea and Aurelia the favorable for the polyp. embryos are retained on the oral arms. Aurelia has brood Scyphomedusae are usually short-lived. They gr'lW p<;('kets on the oral arms of the female where embryos are rapidly and later die due to changes in water tem r"!ained until the planula is released. perature or other factors, which for the most part are not The embryology is quite variable even within the same I ..lderstood. Chrysaora ephyrae from the Chesapeake speci~s. Segmentation is usually total but gastrulation Bay are released in early June. By July these medusae depends upon egg size, with ingression in small eggs and arc mature, and most have died by mid-September. invagination in large ones and by both methods in Aurelia and Cyanf!a may live somewhat longer. medium-sized ova. Usually a planula is produred, whkh The occurrence of some scyphomedusae is related to settles and forms a polyp, but very large <.l!;gs tr· ay hypass water temperature. Cyanea in the Gulfof Maine are first both the planula and polyp and form a m.,ciitied ephyra seen in April or May and are mature by early summer as they do in Pelagia (BerriIl1949). (Bigelow 1926). Cyanea is found in the Chesapeake Bay Symbiosis is widespread in the Cnidaria and occurs early in the winter and matures by February or March. frequently in the Scyphozoa. There are numerous reports Chrysaora occurs only at higher temperatures (about of juvenile fishes associated with scyphomedusae; Men ~O°C) in the Chesapeake Bay in early May, and slightly sueti (1963) discussed these associations. Associations later at Woods Hole, Mass. Because of this dependence between juvenile fish and medusae range from the fish on high temperatures, it apparently cannot strobilate in seeking refuge under the umbrella, or between the ten· colder water and is not found north of Cape Cod. Aurelia, tacles, to predation with the larger fish eating bits of the whose range overlaps' the two previous species. occurs tentacle or gonad. It is not fully understood how certain during the wP.rmer months in both the Gulf of Maine and species of fishes, 10'.1-(., Peprilus and Poronotus, are able to in the Chesapeake Bay, but appears earlier further feed on medusae without ill effects, but it is generally south. In the Gulfof Mexico and off Florida. both Aurelia thought that antibodies may be involved and that the and Chrysaora can occur at almost any time. Rhopilema secretion of murus by the fish may prevent the and Stomolophus also may occur nearly year-round. nematocysts [mm di,charging. Medusae feed on a variety of prey. These include Crustacea are also known to pllrasitize ctenophOl'p's, planktonic microcrustacea (i.e., copepods, scyph"medusne: hyperiid amphipods may both !1teal amphipods, and larvae), larval and small fishes, pelagic food from medusae and feed on medusa! tissue. Phillips polychaetes, siphonophores, and even other medusae. et aI. {I968) discussed the parasitic hahits of the two Fraser (1969) calculated that a Cyanea less than one-half species of brachyuran crabs on medusae in the Gulf of meter in diameter would eat over 1.5 million copepods of Mexien. the genus CalantL< in its lifetime. One can only guess at the amount of food eaten by a 2·m Cyanea. Chrysaora STINGING MEDUSAE feeds mostly on ctenophores and other medusae. Studies which I have made in Puerto Rico (unpublished) indicllte Cllry.'aora is popularly known as the sea nettle because that a Chrvsaora medusa would have to consume about its sting can be very painful. Cyanea also is a stinger: 15 kg of ~tenophore Mnemiopsis in order to attain a Russell (1970) noted that Cyanca was the caust; of a diameter of 150 mm. In Aurelia, food is trapped on the mysterious death in Conan Doyle's story, "Ad\'en1.ure of umbrella and by the short tentacles, and is transferred, the Lion's Mane." The majoritv of the Cubomedusae are not dire.::tly to the lips, as in most medusae, hut to a food stingers. as are coronate nll'dusae, some producing a \'ery groove formed by the small velarium. Cilia move the food severe stillg. Alire/ia, RllClpilema,· Stomolophus. which is mixed with mucus to the adradial canals where Phacellophora, and the Stauromedusae ar~ not known to it is then transferred to the lips (Southward 1955). be severe stingers. Halstead (1965) gave an account of Rhizostomes are filter feeders and many are active t he nature of the wounds and medical aspects of stings. swimmers. In forms such as Rhopilema, water is forced Cnrgo and Schultz (1967) found that when meat ten around the oral arms by the contracting urn brella during derizers containing proteolytic enzymes were mixed with swimming. In Stomolophus, water is forced through the water and applied t, the affected area, the pain was keellike appendages. Minute tentacles, which line these quickl~' relieved. structures, filter the suspended food from the water. The food is then carried to the stomach by the cilia-lined GLOSSARY canals. Stomolophus collected at Beaufort, N.C.. fed mostly on bivalve veligers and harpacticoid copepods ancllors (Fig. 81 adhesive organs between arms of some (author, unpublished). Stauromeduslle. The gonads of scyphozoans are always near the site of calyx flmne!'5haped umbrella of Stnuromedusae. digestion. In semaeostomes and rhizostomes the gonads I.'nidoblar,l c(,ll in which the nematocY3t forms. are infoldings of the sl1bumbrella which also bear the t'llidume complement of nematocyst types present. gastric cirri. Some scyphomedusae spawn while still very co"le'llcrull gastric cavity or stomnch. 5 .'. c~nal furrow (Fig. 2) circular groove which divides furrow orthe coronate medusae (Fig. 2); bladelike pro· ,I'.""'S' the exumbrelle. of the coronate medlJ~ae. jections of Cubomedusae to which the tentacles arl;) ,'- ",uciform cross-shaped. attached (Fig, 1). '. ,..,' phyra (Fig. 5) planktonic larva oi Sl'yphozoa except peduncle (Fig. 8) stalk of polyps, or Stauromedusae. i :: Gllbornedusae and Staurcrnedusse. Usually with 8 po(ydisc _.trobi/ntion (Fit!,. 9) several ephyrae produced 'f '4 Pairs of arms and 8 rhopaHa. from a polyp at the same time. C' e1Cumbrella aboral or upper surface of umbrella. rlwpalium (Figs. 1, 2) hollow, club-shaped sense organ -.. gastric cirri or phacelloe (Fig. 1) hollow fingerlike pro· l(eneTally located near the umbrella marl(in, usually jections in the coelenteron which aid in digestion. with a statocyst and, rarely, an ocellus or ocelli. (appet flaplike structure making up the scalloped mar- scyph,stoma polY!l of scyphozoan medusae excluding '., I"in of the umbrella. th", Stauromedusae and Cubomedusne. ,~'. ,_ odise strobilation (Fig. 10) a single ephyra produced sensor)' niche open structure partially enclosing rho- f. ii'". r strobila at one time. plllium in Cubomedusae. ~ ~:! arm armlike extensions of the corners of the mouth s(mbilo scyphistoma which is forming ephyrae. 1' ' which hang from the subumbrella of the Scyphozoa, str"bilntion process of producing ephyrae. to! ,1 in number in the semaeostomes (Fig. :l) and 8 in the -'llbL,mbrelia orai or underside of umbrella. ..~ rhizostomes (Fi~. 4). ,·..rurilHTI (Fig. 1) sheltlike structure at margin of um pl'dalia radial thickenings between lappets and coronal 1lrella in Cubomedusae ar.d Aurelin.
INTRODUCTION TO THE KEYS TO THE SCYPHOZOA ',. ~ An attempt has been made in constru~ting the keys to remain unchanged. Because of the delicate nature of include those features which 8re most obvious or which mp,dusae .hey are often injured and parts may be missing occur in both juvenile and adult specimen.~_ As medusae or abnormal. Therefore, in using the key some discretion !{I'ow, structures such as tentacles usually increase in must be used for each specimen to determine which number and oral arms become increasingly folded or ('haracterg are usable. complex. while other structures, e.g., rhapaHa, usually
Key to the StBuromedusae of the Northeast Coast of the United States
With anchors ...... :2 Without obvious anchor;;
2 (1) Anchors larj!r: no shieldlike covering: gonads extend into arm!' 3 :! (I) Anchor" small with shieldlike covering at base: arms shon: gonads do not extend into arms .. ,, ,,... .. , Thauma/Uscyphus at/anticus
c
·:;;:;:":t:, r-/'~/ .. b . . .. ". '. " '''''· . G'-~, . d
Fi(ure 11.- Th"umatoacyphWl nl/antic.a. R. sidevie",' b. \\\ at vif!w c. nnchor.liidevicw d. anchor. tront vie'\1:
(j 3 (2) Arms short; anchors bean-shaped; calyx as high as broad .Haliclystus auricula
.'.'; .~ . .:.' ,',". C (J:''':''. b :,:,t:·:: @, ,'..' d
Figure 12.-Hallelyohul auricula. a. lideview b. oralview c. ancbor. Iide view d. anchor, front view
3 (2) Arms long, anchors trumpet-shaped with short projection at center; calyx broader thail high ...... Halidystu.~ salpinx
b
Figure 13.-HaliclYBtU8 BcJpillX. 8. side view b. oral view c. anchor. side view d. anchor. front dew
7 4 (1) Peduncle and arms long; arms paired; calyx broader than high; largest New England Stauro- medusa Lucernaria quadricornis
i-.
b a Figure 14.-Lucerruvia qruufri<,Dm;'. a. side view ba orlll view
4 (l) Peduncle and arms short: arms not paired; calyx higher than brood ...... Craterolophus canuouulus
a b
Figure )1),- Craterolophu.a CORlIovulus. 8. side "'Iiew b. ofal view
8 Synopsis ofEphyrae of ChryBaora. Cyanea. Aurelia, and Rhopilema. (See Russell 1970.)
Newly released ephyrae ca. 2.5 mm in diameter (Fig. 5) Postephyrae Shape of lappets; Morphology of 5 mm in diameter position of rhopalia ephyral arms Gastric cirri Morphology Chrysaorc. lappets slender a pair ofnematocYbt lack oral arms split; lmlg pointed tips; clusters on each arm; ribbonlike; (Fig. 16a, b) rhopaliar cleft deep radial canals in arms marginal tentacles; ca. 1/2 arm length short, consists of 1 pair no ring canal Cyanea lappets slender radial canals in arms usually 4 oral arms not split, pointed tips; long, consists of 2 pairs wide and flaring; (Fig. 16c, d) rhopaliar cleft deep subumbrellar ca. 1/2 arm length tentacles; no ring canal AUr2lia lappets broael, tips radial canal to with 4 or more oral arms split; blunt, rounded; rhopalium only short; marginal (Fig. 16e, f) rhopaliar cleft short tentacles; ring 1/2 arm length canal present
Rhopilema lappets slender, tips radial canals in arms with 4 or more oral arm split several p;..\nted; rhopaliar long, consists of 1 pair times; no marginal (Fig. 16g, h) cleft deep ca. 1/2 al'm or subumbrellar length tentacles
a b c d
e 9 h
Figure 16.-Epllyrae (a, c, e, g); postepbyrae (b, d, f, h). e, b Clu-ysaora qu./nquecirrha c, d Cya"ea capUlata e, r Aurelia aur/ta g. h Rhopi/cma verrilli (from I).. Calder 1972, unpuhlished; and Calder 1973) 9 Key to the Pelagic Scyphomedul'ae of the Northeast Coast of the United States 1 Umbrella, saucer·shaped, dome-shaped, or hemispheIical; with 8 or more tentacles, or no tentacles;
with 8 or more rhopalia ..••...... 0 • 0 0 •• 0 0 0 0 0 0 0 ••••• " 2 1 Umbrella tall rectangular; 4 tentacles; 4 rhopalia; sensory niche with covering scale above and below; stomach supported by 4 mesenteries Order Cubomedusae Tamaya haplanema
....\.{- ~~
b
Figure 17.-Tamoyahap/onem 2 (1) Umbrella divided into two regions by a deep circular groove; stomach darkly pigmented; bathy· pelagic, rarely at surface or coastal 00000••0•0•00•000••• Order Coronatae _ 3 2 (1) Umbrella not divided by circular groove, not darkly pigmented; mostly coastc.l or at surface 0 .4 3 (2) Umbrella flat, disc-shaped; tentacles and rhopalia equal in number, usually 22; gonads button· like 0000•00000•0•0 ••• 0 0 ••••• 0 0 • 0 • 0 0 •••••• 0 0 0 0 0 •• 0 0 • 0 0 .Atalla wyvillei Figure 18o-Atolla W)'lJilIei••ide view. 10 3 (2) Umbrelll' conical or dome-shaped; usually 3 tentacles between r.hopalia; 4 rhopalia; 12 ten- tacles: gonads horseshoe-shaped , PeriphyUa periphyl/a FIl(1a'e 19.-Perlph:.llo perlJ)hyllo. aide vie... 4 (2) With marginal or subumbrella tentacles; 4 oral arms; distinct mouth; saucer-shaped umbrella ...... Order Semaeostomeae 5 4 (2) Without tentacles on umbrella; 8 tough gelatinous oral arms; mouth smali or absent; umbrella usually hemispherical ...... Order Rhizostomeae 9 S (4) With numerous narrow radial canals extending from stomach to margin; ring canal near mar gin; tentacles either on margin. short and numerous or in 16 linear groups on subumbrella ...... Family Ulmaridae 8 Figure 20.-Famlly Ulmaridae, radial and ring canala in block. Figure 21.- C)lan{!Q and family Pelagirlae. stomach pouches in block. 5 (4) With broad stomach pouches ext.ending to margin; without ring canal; telltacles long 01' 8 U- shaped subumbrellar groups . . 6 11 6 (5) Oral arms ribbonlike; tenta_~les on margin Family Pelagidae ...... 7 6 (5) Oral arms wide, curtainlike; tentacles in 8 V-shaped groups on subumbrella Cyan,'u capillata Q Figure 22.- Cyane"""pUlata. a. olde"lew b. oral ,"jew, oral armsremoved and rour groupo orlolnlacleo removed 7 (6) With 8 tentacles; exumbrella with prominent wartlike projections; umbrella nearly hemis- pherical in shape, usually lees than 100 mm in diameter; oceanic .. Pclngia noctilum Figure 23.-Pelagia noctU...,,,. a. sici.eview b. oralview, orsl a.rms remov~ 7 (6) With more than 8 tentacles; exumbrella smooth; umbrella saucer-shaped, up to 200 mm in diameter; estuarine or coastal Chr)'saora quinquecirrha a Figure 24.-Chrysaora quinqueclrrha. a. side view b. oral view, oral arms romo"ed 8 (;') Marginal ter. a Figure 25.-AurclialUU'ila. a••Idevlew b. orlll view, oral arms relnoved 3 (7) Tent.acles long, in 16 subumhrellar linear groups; oral arms broad. curtainlike; 16 rhopalia ...... Phacellophora camtschatica .' ~. r-:~ . • 7:'" 0;.,., r~ 1\ I lb, ~~ 1 'rIClMtz; :1 ~ \ ~ I< ,\ , \ \ II•• J()I \ I ~I; I : I I I a b Figure 26.-Phae.lloph.ro camt.hallea. a. aideview b. oralvle'w, oralarmssndone- halfortenlacle. remov<'li 13 9 (4) Oral arms free along most of their length; spindlelike '1ppendages hang from oral arms; um- brella hemispherical or flattened ...... Rhopilema uerrilli /.~ . ',- f' . • • , , 'J. . .' 'J':'. - . ·· .:~,..."', l Figure '1:1.- Rhop/lema "errilli, aide view. i \::i :,1 ,. :t. 9 (4) Oral arms fused; no spindlelike appf:ndages; umbrella higher than a hemisphere (globular)...... Stomcilll!?hus meleagris .:.c . . . {,. . (' Figure 28.-Slomolopllue meleaaria, aide view. ANNOTATED SYSTEMATIC LIST HaLiclystus auricula (Rathke 1806) Massachusetts northward, Northeast, Atlantic, The following list is arranged according to the North Pacific, lower intertidal and subtidal on algae classificat:on of Kramp (1961). Distribution notes are and Zostera, at Woods Hole most common on brown from published records and from museum specimens. algae Scytosiphon. Juveniles pasily confused with Most of the species listed are represented by specimens H. salpinx, but arms are very' short. Coloration: in the U.S. National Museum, Washington, D.C., and highly variable, brown, greer" red. Berrill (1962); the Gray Museum, Marine Biological Laboratory, Woods Mayer (1910). Hole, Mass. Notes on seasonal occurrence, habitat and life history, and coloration are included. References to Haliclystw; salpinx Clark 1863 important papers are cited for both the families and for Massachusetts northward, lower intertidal and slib each species. tidal on algae and Zostera. May be confused with juvenile H. auricula but with long arms. Coloration: Q..ASS SCYPHOZOA variable, green, brown. Berrill (1962, 1963). Lucernaria quadricornis 1776 ORDER STAUROMEDUSAE O. F. Muller Massachusetts northward, Northeast Atlantic, Arc FAMILY ELEUTHEROCARPIDAE tic, lower intertidal and subtidal mostly on References: Berrill (1963); Kramp (1961). Laminaria. Largest New England stauromedusa 14 reaching over 50 mm in length. Coloration: highly At least four other coronate medusae may also occni in variable, brown, green, red. Berrill (1962, 1963). the northwestern Atlantic: Linuche unguiculata (Schwartz), a specimen at the U.S. National Museum, FAMILY CLEISTOCARPIDAE collected off Cape Hatteras, lat. 35~N, long. References: Herrill (1963); Kramp (1961). 75°W. Nausithoe punctata Kolliksl', a specimen at the Craterolophus con'Jolvulus (Johnston 1835) USNM collected off Georges Bank, lat. 40oN, long. Massachusetts, Northeast Atlantic, intertidal on 68°W. Nauphantopsis diomedeae Fewkes, not reported Fucus where it attaches near the bladders, incon since the original description, off New England. (Kramp spicuous b~c6use of its shape and color. Colora 1961.) Atolla vanhoetfeni Russell, a specimen at the tion: olive green when attached to Fucu.~. Kramp USNM, collected off Virginia lat. 36°39'1'1. long. (1961). 74°39W. (Russell 1970.) Thaumatoscypus atlanticus Berrill 1962 Maine, subtidal on Laminaria. This species is un usual in that ocelli are located on the anchors. Coloration: not known. Harrill (1962, ORDER SEMAEOSTOMEAE 1963). FAMILY PELAGIDAE References: Kramp (1961); Russell (1970). ORDER CUBOMEDUSAE Chrysaora quinquecirrha (Desor 1848) Possibly cosmcpolitan, western Atlantic distribu FAMILY CARYBDEIDAE tion from Cape Cod to the Gulf of Mexito. Abun References: Bigelow (1938); Kramp (1961). dant during the summer months in estuaries, not Tamoya hap/onema·F. Muller 1859 common in the open ocean. Scyphistoma occurs on Tropical Atlantic, Gulf of Mexico to Massachusetts. oyster shells in upper estuary. Popularly called "sea Probably carried north in the Gulf Steam. Mayer's nettle" because of its stinghg ability. Coloration: (1910) report of this species in Long Island Sound is highly variable, pink, yellow, often with 16 yellow the only report of T. haplonema in the New England ocher or red radiating stripes. Cargo and Schultz coastal waters. Coloration: colorless except ten (1966, 1967). tacles, pink. Bigelow (1938); Phillips and Burke Pelagia noctiluca (Forskal 1775) (1970). Cosmopolitan open ocean species in warm and tem perate oceans. Carried north in the Gulf Stream, Three other Cubomedusae also occur in the sometimes seen in coastal New J£ngland northwestern Atlantic. Carybdea a/ata Reynaud. a waters. LackfJ scyphistoma stage. Coloration: specimen in the U.S. National Museum (USNM) was highly variable, blue, yellow, pink, tentacles collected off Cepe Hatteras at lat. 35°N, long. red. Russell (1970). 75°W. Carybdea marsupia/is (Linneaus) occurs in the Gulf Stream near Bermuda but has not been reported on FAMILY CYAl\iEIDAE our coast. Chirapsalmus quadrumanus (Muller) occurs References: Kramp (1961); Russell (1970). along U.S. coast south 0f Virginia. Bigelow (1938); Cyanea capillata (Linnaeus 1758) Phillips and Burke (1970). Cosmopolitan in colder oceans. Occurs in the Gulfof Mexico and Florida northward. Most abundant ORDER CORONATAE during winter through spring in the south, summer to fall in the north (New England waters). Popularly FAMILY ATOLLIDAE called "lion's mane" because of the long red-yellow References: Kramp (1961); Russell (1970). tentacles. Largest New England medusa reaching a Atalla wyvil/ei Haeckel 1880 meter or more in diameter. Coloration: highly Cosmopolitan in deep oceans. A deep-sea species, variable, pink, red, yellow, brown. Russell (1970). bathypelagic, seldom occuring at the surface or on the continental shelf. Commonly taken in midwater FAMILY ULMARIDAE samples in deep water usually below 500 References: Kramp (1961); Russell (1970). m. Coloration: stomach dark red-brown. Russell Aurelia aurita (Linnaeus 1758) (1970) . Cosmopolitan species in nearly all oceans except the Arctic. Popularly known as "moon jelly" because FAMILY PERIPHYLLIDAE of the whitish umbrella. Coloration: bell clear; References: Kramp (1961); Russell (1970). gonads, tentacles, pink, purple. Russell (1970). Periphylla periphylla (Peron and Lesueur 1809) Phacellophora camtschatica Brandt 1838 Cosmopolitan bathypelagic in deep oceans. Com Occurs north of Cape Cod in the western Atlantic, mon in midwater trawl samples from deep water. It probably the entire Pacific coast of the Americas has been taken near the surface off New and in the Mediterranean and off western England. Coloration: stomach dark red Africa. Rarely taken in New England brown. Russell (1970). waters. Coloration: yellowish. Mayer (1910). 15 ORDER RHIZOSTOMEAE guiculata. with a brief reexamination of scyphozoan nematocyst classification. Chesapeake Sci. 15:170-173. FAMiLY RHIZOSTOMATIDAE CARGO, D. G., and L. P. SHULTZ. 1966. Not... on the biology of the sea nettle, Chrysaora quinque References: Kramp (1961); Mayer (1910). cirrha, in Chesapeake BhY. Cbesapeake Sci. 7:95·100. Rhopilema vurilli (Fewkes 1887) 1967. Furtherobservati"ns on the biology ofthe sea nettle and jelly. Occurs in tlle northern Gulf of Mexico, North fishes in Chespeake Bay. Chesapeake Sci. 8:209-220. Carolina to Connecticut. An uncommon species, FRASER, J. H. found along ClJast and b mouths of es 1969. Experimental feeding of BOrne medusae snd Chaelognatha. J. ~-i.h. Res. Board -Can. 26:1743-1762. tuaries. Coloration: bell milky; oral arms HALSTEAD, B. W. brown. Mayer (1910); Calder (1973). 1965. Poi•.onous and venomous marine animals of the WNld. I. In venebrates. U.S. Gov. Print. Off.. Wash., D.C.. 994 p. (see FAMILY STOMOLOPHIDAE Phylum Coelenterata, p. 297·371). References: Kramp (1961). KRAMP. P.L.· 1959. Stephanoscyphus (Scyphozoa). Galathea Rep. 1:173-185. Stomolophus meleagris L. Agassiz 1862 1961. Synopsis of the medusae of the world. J. Mar. BioI. Aaaoc. Occurs in the western Atlantic from Brazil to Cape U.K. 40:1-469. Hatteras and from Panama to San Diego in the MANSUETI, R. eastern Pacific. Specimens occasionally carried 1963. Symbiotic behavior between small fishes and jellyfishes, with new data on that between the stromateid. Peprilus alepido~ north to NE'w England waters by the Gulf Stream. tus, and the Bcyphomedusa, ChrysQora quinqueci"ha. Copeiu Popularly known as the "cabbage head jelly5sh" 1963:40·80. because of its globular shape. Coloration: brown MAYER, A.G. band at margin. Mayer (1910). 1910. Medusue of the world. m. The Sc,)'phomedusse. CarllcF;ie Inst. Wash. PubI. 109:499-735. PHILLIPS. P. J., snd W. D. BURKE. 1970. The occurrence of sea wasps (Cubomedusse) in MiFosissippi Sound and the northern Gulf of Mexico. Bull. Mar. Sci. 20:853· SELECTED BIBLIOGRAPHY 859. PHILLIPS, P. J .• W. D. BURKE, and E. J. KEENER. BERRILL, M. 1969. Observations on the trophic significance af jellyfishes in Mis 1962. The biolollY of three New England Stauromeduaae. with a sissippi Sound with Quantitative data on the associative behavior dcscripti~n of a ncw spccics. Can. J. Zool. 40:1249-1262. of smaIl fishes with me 16 SYSTEMATIC INDEX Atolla salpinx . 7, 14 vanhoeffeni · ... 15 Linuche unguiculata · .15 wyvillei .. · .10, 15 Lucemaria quadri.:omis 8, 14 Atollidae ... · ...15 Nausithoe punctata ... · .15 Aurelia aurita 9, 13, 15 Nauphantopsis diomedeae · .15 Caa7a~~e~ Pelagia nocti/uca .. ... · 12 . .. · 1[; Pelagidae . · 11, 12, 15 marsupialis · 15 Periphylla periphylla · 11,15 Carybdeidae ...... · 15 Periphyllidae . · 15 Chiropsalmus quadrumanus · 15 Phacellophora camtschatica · .13, 15 Chrysaora quinquecirrha .. 9, 12, 15 Rhizostomatidae . · ...16 Cleistoc8rpldae . · ...15 Rhizostomeae ..... · .11, 16 Coronatae . · .10, 15 Rhopilema verrilli ... 9, 14, 16 Craterolophus COTlvovu!lts ·. B,15 Semaeostomeae .... .11. 15 Cubomedusae ...... · .10, 15 Stautomedusae . . 6,14 Cyanea capillata 9, 12, 15 Stomolophidae . ...16 Cyaneidae ...... 15 Stomolophus meleagris .14, 16 Eleutherocarpidae .. 14 Tamayo haplanema ...... 10, 15 Halicl;lstus Thaumatoscyphus atlanticus ... 6, 15 aurlcuUl 7, 14 Ulmaridae . ...11, 15 17 ACKNOWLEDGMENTS Preparation of the "Marine Flora and Fauna of the North to collaborate in i.he preparation of manuals, reviews eastern United States" is being coordinated by the following manuscripts, and advises the Scientific Editor of the National Board: Marine Fisheries Service. Coordinating Editor: Melbourne R. Carriker, Marine Studieg Thanks go to Marie B. Abbott, Curator of the Gray Museum, Center, College of Marine Studies, University ofDelaware, Marine Biological Laboratory, for the loan of specimens and for Lewea, DE 19958. assiatance during a short Visit there; S:laUf, Ruetzler, Curator of Editorial Advisers: the Division of Echinoderms, U,S. National Museum, Washing· Marie B. Abbott. Marine Biolr:gkal Laboratory. Woods ton. D.C.. for use of collections. dark room, and other facilities: Hole. Mass. Maureen E. Downey, also of the Division of Echinoderms, and Arthur G. Humes, Bosto" L'ni"ersity Marine Program, Charles E. Cutress of the University ofPuerto Rico, Mayaguez, Marine Biological Laboratory, Woods Hole, Mass. for encouragement, helpful suggestions, and assistance in proof· Wesley N. Tiffney, Department of Biology, Boston Univer· reading; Dale R. Calder, formerly of the Virginia Institute of sity, Boston, Ma58, Marine Sciences, Gloucester Point, and now at the South Ruth D. Turner, Museum of Comparative Zoology, Har Carolina WiidHfe and Marine Resources Department, Charles· vard University, Cambridge, Mass. ton, for critically reading the manuscript, for permission to ex· Roland L. Wigley, National Marine Fisheries Service Bio· amine the scypbozoan polyps maintained at the Virginia In logical Laboratory, NOAA, Woods Hole, Mass. stitute of Marine Sciences, and for the loan of unpublished data Robert T. Wilce, Department ofBotany, University ofMas· and figures; Cadet Hand, Kay Petersen, Roland Wigley, and sachusetts, Amherst, Mass. Ruth Turner for critically reading the manuscript; and typists The Board established the format for the "Marine Flora and A. Stonework. S. J. Larson, S. Khoury, K. S. Stemler, andJ. S. Fauna of the Northeastern United States." invites systematists Ballantine. COORDINATING EDITOR'S COMMENTS Publication of the "Marine FlO!a and Fauna of the North Scyphozoa. Currently he is involved with graduate studies at eastern United States" is most tim2ly in view of the growing un the University of Puerto Rico, Mayaguez. where he is inves iversal emphasis on environmental work and the urgent need for tigating the functional morphology of scyphomedusae and more precise and complete identification of coastal organisms aspects of growth and reproduction. than has been available. It is ma.,datory, wherever possible, Preparation of this manual was supported in part by a grant that organisms be identified accurlltely to species. Accura from the Environmental Protection Agency to the Editorial scientific names unlock the great quantities of biological infor Board of the "Marine Flora and Fauna of the Northeastern mation stored in libraries, obviate duplication of research al. United States." Work on the "Marine Flora and Fauna of the ready done, and often make poasible prediction of attributes of Northeastern United States" by the Coordinating Editor is sup organisms that heve been inadequately studied. ported by the College of Marine Studies, University of Ronald J. Larson started his study of medusae during the Delaware. summer of 1969 when he worked on the natural history and sys tematics of Hydromedusae from the Oregon coast. In 1972 he Manuals are available for purchase from the Superintendent began working at the U.S. National Museum where use of the of Documents, U.S. Government Printing Office, Washington, extensive collections and excellent library facilities furthered his D.C. 20402. The manuals so far published in the series are listed education and initiated an interest in (~Ie systematics of the below: COOK, DAVID G., and RALPH O. BRlNKHURST. Marine flora and fauna of the Northeastern United States. Annelida: OIigochaeta. BORROR, ARTHUR C. Marine flora and fauna of the northeastern United States. Protozoa: Ciliophora. MOUL, EDWIN T. Marine flora and fauna of the northeastern United States. Higher plants of the marine fringe. McCLOSKEY, LAWRENCE R. Marine flora and fauna of the northeastern United States. Pycnogonida. MANNING, RAYMOND B. Marine flora and fauna of the northeastern United States. Crustacea: Stomatopoda. WILLIAMS, AUSTIN B. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda. POLLACK, LELAND W. Marine flora and fauna of the northeastern United States. Tardigrada. LARSON, RONALD J. Marine flora and fauna of the ncrtheastern United States. Cnidaria: Scyphozoa. 18 Co u. S. GOV~RNHtNT PQINTING Of:~ICE: 1976-697.707/108 "EGlON 10 370. Collecting and proce5sing data on fish t"IiCKS and larvae in the· Calilcrnia 381. fi!lhery publlutions. ulendar year EJ67: Lists and indexrs. By I..ec C. Current re~on. By David Kramer, Mary J. Kalin. Eliubeth G. Stevens. Thorson and Mary Ellt"n Engett. July 1973, iv + 22 p.• 1 fig. For we by the Jamt.'s R. Thrailkill. and James It Zweifel. November 1912. iv + 38 p.• 38 Superintendent of Documenls. U.S. Government Printing Gffite. Woshington. figs .• 2: tablt.'s. FQr salt.· by the Superintendt'nt of D()("umenl.'l. U.s. Govern D.C. 20402. me-nt Prinlinl( Office. WashinK\on. D.C. 20402. 3H2. f'ishc:'fY publintions. calendar yrAr 19ti6: Lists and indexes. By Mary 371. Ocean fL-lht"ry mllnaKl"mt"nt; Discussions and rest"a~h. By Adam A. Elh'n EnKetl and Ll't! C. Thorson. July J973. i..· + 19 p.. 1 EJ{. For salr by SokolMki ,editor). 111 papeJ"!il, :?4 au' hors. I April 1973. "'i + 173 p.. 38 riKS.• the Supt'rinlendrnt of Documents. LJ ,S. Government Printing Office. Was.hinK :r.! table~. 7 appendix tables. ton. D.C. 20402, :.vn. Fishtor)' publications, calendar year l~:',;: Lists and indexes. By Ltol' C. l.i~t~ 372 "'ishery publicntion!o. calendar yur 1971: and indue!. By Thomu ThoNlR and Mary 1o~IIt'n EnKt'U. July 1973. j..,' + 12 p., 1 fiR'. For sale by the A. Manar. Odober 19T1. i ... + 2-1 p .• 1 fig. For !!oalti' uy the Superintendent of SuperintNlrlt'nt of Uocumt'nts, ll.S. Gov{'rnm"nL Printin~ OHirp. WuhinK' Documents. U.S. Governmt"nt PrintinJ,r Orrin'. WashinKlon, n.c. 20402. tun. h.C. ~,'O402. 374. Marine.' flora and (aunH of the.' northrnlt'rn Unitt'd States. Annelida: :*,.1. ~tarint· nora a.nd fauna uf tht> northt'8Stt'rn Unit~ States. HiKher p~nts OHgochaeu. By David G. Cook and Ralph O. Brinkhunt. May 1973. Hi + 23 tlf thl' mariru.' frin~t.'. By I::dwin T. Maul. SI'ptt'mbt'r 1973. iii + 60 p., lOS p., 82 fi~s. For ~ale by the.' Superintt>ndt'nt of Dnc·umenb. U.S. Go\'rrnment figs. 1o"or sale by the Supuintt'ndpnt of Documflnts. U.S. Government PrintillK Pri:"ltinlr\' Oflict.·. Washin..,.on, D.C. 2O~02. Offict·. W ol~hinh'1on. I).C. ~>()~02. 315. ~('w I·olyrharla from Bt'3ufort. with a ke-r to lSll spe::es ffforded from :~!i. 1o'istwry pUblications. c.ll.ll'ndar )"e.·ar 1972: Lisb and indt'Xf'5. B)' l.A'{" C. North Carolina. By John H, Day. July 1973. xiii + I.tO p., IB r:,i(!I .• 1 tahle. Thurson and Mary 1o:Ilt.'n En/(ett. !'in\'{'mlwr 1973, i\' + 23 .., .. 1 fiR'. For §.alt.· For ~Ic b)' tht· Superintt'ndent of llocumpnts. U.S. r;o..It.'r~In';.·:1l Printing hy thl' Supl'rintendl'nt nf Docur.lrnts. U.S. \io\'rrnment P:'intinK Offict" Offir('. Wa.... hin~on. 11.C. :'>Q40:? Wn..,hinKtlln. n.c. :"»O,1O:L :.176. Bottom·water tt.·mpl'raturt·s on tht, continental ~ht'lf. NOVA :;,('oti~ to :~i~~. B~al~~'~~~ R~~~CI~:k:;. t~:p~;~teeras:973. Yiri~d 1~t8~~S'1 Plt;~O~~r ~t·w Jt.·r"i.t'!·. Oy John R. Colton. Jr. ;snd Ruth It. Stoddard. JU:It,' !iH:t iii + 5:') p.• 15 fiKS., 12 a.ppendix tables. For !itnle b)' the Supt'rintendr~1 of :'0.114.' h)' thl' Supe.·rintcndt·nt of [)t)("uments, lJ.~. l"o\'{'rnmt'nt Prin~lnK Offict.'. Ouc·umt.·nt.... U.S, Gon·rnmt.'nt Printinj{ Office. WD.ShinKf,on. D.C. 20402. Wa."hin,lCton. D.C. ~>Ool":!_ :n7, Fho.her)' publications. cale.·ndar )'t'ar 1970: Li~h lind indcxt's. By Mar)' ;l.'ii, Marini' nura and fauna of thl' nurtht>uu·rn L~nitt'd Stah·!rI. rru~la{'t'A; Ellrn En!tetl and Lee C. Thonon. December 1972. iv + 3-S p.• 1 fig. For .'ialt' Stomlltopoda. By Raymond B. Manuin,lit:. Jo'ebruary 1914. iii + 6 p.• 10 fig.. For by the Supprintendrfit of Documl"nt.s. U.S, Go\'oemment PrintinK O(fiee, .."II' h)' tht' ~upt.·rintl'ndent of [)ocuml'nt.. , U.S. lio,,·('rnmt.'nt Printinlr\' Offil"t'". \\'"shinl{ton, n.c. 2O~02. \\'I\,\hin",'1nn, n.r, W-1O'.!. 37~. ~tarint' nor" llnd fauna of tht-· northe.·"st('rn L'nitt.'fI Statt."s. ProtolO": :IXH. I'rot'l'rdinl{s nr ttlt· first U.S.·Japan mt'l·tinK un aql..atultur," HI Tl·~.YO. Ciliophora. Hy Arthur C. Horror. ~i('ptt'mtwr 197:1. iii + 62 poo 5 fi~!'l. For salC' h,,'ln. l)rtoht'r H~ 19. 1971. WiIIi:tm ro-:. Shl\~' It"ditorJ. (1M pa~tt'r"". 14 auth~nd h)' the Supf'rint4'ndent of DO<'uml'nt:'l. V.S. Guvprnml'nt Printin/{ Offit"t,. Ft·hrlJar~.. 191·S. iii + 1:13 p. For salt· hy tht· SUpt.'rintl'f,ct:nt of n()('um,·M~. Washinl{lon. n.c. ~402, l'.S, GClwrnmt'nt Printinl{ OffiCI'. \\'ashin~clR. n.c. 2().l02 :J79. Fishl·ry publit'J,lion~. <"31l.'l"idar yrar 1969: Li~ts anfi indexe"i. By Lt't.· C. :\."l~l, ~1i\rint· n'lra l\nrl flluna or tht' nOrlht'a!it€'rn llnitl'd Stall'", CrustaC('!L Thor~on a.nd Mary Ellen EnRel'.. April 1973. iv + ~11 p.. 1 fiR. For !lalp by 1l1'4·;'!M"II!:t. By Austin B. William!o. April 1974. iii + ~O p.• 111 rii~S, 1"01' sale lhl' Su~>t.'rintt.·nrl[nt of nOtumt·nt!'i. V,S. l;un·rnme.'nt PrintinK OHi("l'. Washinl{ h.\ thl' Supt'rintl'ndl'nt of n,....ume.'nb. ll.:-;. GovI'rnmf'nt Printinf{ OUirt'. ~nn. n,e. :,>O-l02. W",hinKtnn. I t.e, :"~40~. :*,,0. rl~ht'r)' publicatiuns. ('alt-·nda.r )'('ar 1%H: l.i~t~ and ind{·xes. Ii)' ~tar)' :~~1. Fi.. lwr)' puhlir:ltion... rah'ndar )'I'ar 197:i: l.il>;ts nnd inde-xl·s. By ~!ar)' Elh'n ~:nKl'tt :and L4.'t" C. Thorson. ~hy t973. i..' + 2~ p.. 1 fiR. For ~lt· by ElIl'n EnKe.'lt and I.,t't." C. Thor~on, St'pterr.bl>r 191~. iv + 14 p., I £iK- f"or tht' Supt'rintt'ndcnt ()( {)ocume.·nts. l!.S. Govl'rnm.'nt rrinlinr-: O(fi("I'. \\':t!lhinJ{' ";1\1· ~l\' ttll' ~tJl)l'rin(t'ntlt'nl uf IhJ("ulnt·nt~. 1!.~. Go..... rnmt"nt Prinllnlr\' Orfict.'. t{ln. h.C. 20-102. W;l:\hln~ton. n.t". :">0402, /e;