THE \V:\SM:\XX JOUR NAL OF B J O L OGY

\'oL ;4, Xo. I SPRII'C, 1976

A N ew Species of Aglauropsis (: Limnomedusae) from the Northeastern Pacific, with Notes on Aglauropsis conantii and Eperetmus typus

CLAU DIA MTLLS'. JOHX T REES, and CADET H:\1\D, Bodrga Marine L aboratory aud Depart mcul of Zoology, l.'uit·ersil y of Calijomia, Bndcga Bay, California 9-1913.

l l'TRODUCTION ince 1960, an undescribed species of the Umnomedusan . lJ{Iauropsis has been collected from Lime to time in 01e plankton of Bodega Bay and Tomales Bay, central California. imi larities to Epcrctmus typus Bigelow {1915) h ave led us to a re-examination and reassessment of specimens from J apan reported to be E. t·ypus. We here describe this new Cali fo rnian Al(lauropsis, a genus pre­ viously recogni?.<'d only from the south Atlantic, and present ob­ servations on its behavior and ecology. We have adopted the spelling Olindiasidae for this family of Limnomedusae, as emended by 1\Ioreira and Yamashita {1972).

~ IETHODS exually mature medusae of Aglauropsis acora new species were collected in J une and July of 1972 and 1973 in Bodega Bay by means of surface plankton tows with a meter net. Live. healthy anima ls, including some ju\'eniles were also collected in beach wash on alrnon reck Beach which lies just north of Bodega Bay. ~I edusac were maintained in the laboratory in I 500 ml beakers of filtered sea water and gently aerated by means of ~-t l ass pipettes

I l'rcsc•n( uc lc l c·c· >~: 70·1•1 50th Avenue N.E., Seattle. w :.

[23] 24 THE \\'ASMAX:\" JOUR.'\AL OF BIOLOGY, \'!1L.• 1-1 , Xo. I, 1976

fitted to sur~ica l tubi ng. TI1e beakers were placed in a shallow water table and maintained a t a temperature of about li"C. The water in the beakers was chan)!e

and from the pre. erved material listed above. ize ranj!es were de­ termined by meas urin~ at least 20 of each kind of nernatocyst, except in cases of uncommon varieties, where 5 I 0 were measured. Some fired nematocysts were stained with methylene blue to observe fine structure more clearly. Cnidoms were studied from seven specimens of :1 . acora new species, two specimens of . 1. conantii, two specimens of E. typus, and three specimens of " E. typus" from japan.

Aglnuropsis ncora l\Iills, Rees & Hanel, new species. ( FJCUIIL:S 1-7)

. l glaurop.!i; l>(>., R"-LS and II.\1\:D ( 19 i5:77, 8.1). Dt,\GNOsrs. Adult medusae with gonads hanging as a wavy cur­ tain with simple edges from the entire length of the radial canals. Lips of the manubrium bordered by a row of spherical nematocyst batteries. P dunde absent. D ESCR IPTION. Planulae are oval in shape, from 75 to 100 11m long and about 50 11m wide (fig. 1). Planulae settled from 4 to 12 days after being removed to clean fi nger bowls. The planulae were slightly adhesive and difficult to remove with a pipette from the bottom of a container. They did not swim extensively, but moved slowly aJo n~ the bottom, usually settling in the ~cn c ra l area where they were inilially placed in the bowl. Jn some cases, the planulae were observed to undergo "amoeboid" shape changes ( fi g. 2) . This occurred when they had crawled into the thin space between glass slides and the bottom of a finger bowl. ·o obvious net change of position occurred during this activity and its function is unknown. Settled planulae fla ttened into discs 75 p.m in diameter. Polyps, appearin~ as protuberances arising from the basal discs, were seen about 5 to 10 days after planular seulement (fig. I ). The discs were in many cases obscured by debris adhering to the bases of the polyps. About 1 month after settlement, 12 to 20 vertically oriented nematocysts were seen on the tips of the polyps, now 0.2 mm in height. All of the nematocysts were microbasic euryteles and measured 6.5-8.5 X 4.0-4.5 p.m in living polyps. Pieces of brine shrimp larvae were observed to be caught at the nematocyst-laden tips of several polyps, but no evidence of ingestion was seen. So me of the polyps contracted lo about l, ~ their height when touched 26 THE \\"ASMA'\X.. JOUR..\1:\L OF BIOLOGY, \ "oL .•1 4, Xo. 1, 1976

0 b

OJ mm

F1CUR£ I. Oc\"clopmenl of the primary polyp of .l glaurop. with ncmatocysts 20 dnys after planular settlement. All arc drawn to scale. J2 THE \\"ASMA<\1\ JOUR!\AL OF BIOLOGY, VoL. 34, Xo. 1, l 9i 6

0 b

0

d e

0

F l l.UUI:: i Cnidom of Aglauropsit acora, from li\ 111~ material. Drawn ap­ pro:\imatch to calc. From primary polyp. a, microh:1sic cur) lclc. From adult medu53 tent:.ck's: b. microbasic eurytele , c. holotrichous isorhi.za. Lips of the manubrium: d. lar~e microbasic curytele , ,. small microbasic cury­ telc. r holotrichou5 isorbiza. for " fishing" or swimmjng. The adaptive significance of these pos­ tures appears to be that with full tentacle expansion the presents a maxi mal area for fi shing as it hangs molionJess or sinks through the water column, and while swimming the tentacles are shortened which reduces drag through the wnter. Medusae also LIMNOMEDUSAE-MlLLS, REES & HAND 27

OJ mm

FIC:URE 2. ilglauropsis aeom planula undc r~ro in l! "amoeboid " chanJ.(CS in shape. A 2-minutc sequence is depicted. with a probe. o further development was evident and the polyps disappeared after about 5 weeks. The mature polyp and young medusa stages of AKlauropsis acora are unknow n. l\Iature adult medusae may reach 20 mm in diameter and 15 mm in height, with a thick, solid jelly (fig. 3). In life, the umbrella is bowl-shaped. The manubrium hangs down about ~:! the length of the bell cavity. The mouth has four large, flared, crenulated lips, bordered by a row of closely packed, spherical nematocyst batteries (fig. 4). The fo ur radial canals and the ring canal are broad. Curtainlike gonads extend the entire length of the radial canals, on mature specimens folding 4 or 5 times, separated by a short space from the manubrium and ring canal (fig.S) . About 200

FIGVI\E .3. Aglauropsis aeora, adult medusa. Dmwn from a photograph of a li ving animal 20 mm in bell diameter. 28 THE \VASMAI'\K JOUR:\AL OF BIOLOGY, VoL . .l4, :\o. I , 1976

TABLI:; I. Cnidom of A!!lauropsis aeora. All measurrme111s are of unfired cap.wles in p.m; from lh•ing material.

;. Iicrobasi c Hulotrichou; curytel

Primary polyp 6.5-8.5 X .J.o-5.0 nol prcscnl

:\dull medusa (20 mm diameter)

tentacles 14- IS X 8-9 6-9 X 4 (many) (few)

lips o f Lhc Size ] 13.2- 1 ~ .0 X 7.S-S.2 6.2- 7.2 x ;u-.u manubrium (few) ( icw)

Size ll 6.5-10.0 X 3.5-6.2 (many) hollow tentacles, all of one kind, arise from the bell margin. T he older tentacles are turned upward from their bases at the bell margin in fu rrows on the exumbrella and extend outward from the exumbrella a short distance above the margin. T he tentacles have rings of nematocysts which begin a short distance from the ten­ tacle bases and ex tend their entire length. :\lature tentacles have over 200 of these rings. The tentacles terminate in a round knob of nematocysts and p roximally arise from a pad of nematocysts on tl1e bell margin that, on the larger tentacles, is cleft (fig. 6). There are from 60 to 100 internal statocysts on an adult medusa, located between the bases of the tentacles just overlying the ring canal, each with one spherical otolith. Live animals are transparent, with the lips and tentacles a pale pink-brown, and the gonads and manu­ brium a light, translucent blue-green. \Yhen preserved in 10'7a iormalin, the medusae become slightly opaque and the gonads appear either a pale blue-green or pink-brown. The cnidom of A. acora is presented in table l and figure 7.

TYPE LOCALITY. Bodega Bay. Sonoma County, California (38" 18' N lat., 123" 2' W long. ). Surface waters.

TYPE MATERIAL. The type specimen is an adult female, the bell of which is 12 mm high and I S mm in diameter. The holotype (collected 23 June 19 72, Bodega Bay, at the surface) has been de­ posited at the National M use um of Natural H istory (Smithsonian LIM:\OMEDUS:\E- MILLS. REES & HA."\"D 29

200 IJm

Fu;URf. 4. Manubrial cd~c of an adult mcdus:\ of .tglaurofJ•is arora.

Institution) , t: I'OI 54259. Two lots of paratypes have also been deposited at the .1'\:\I ·H. as follows: t:SX:\1 54258 (seven speci­ mens. collected Au!_rtiSt 1967) and t;SX:\1 54260 ( three specimens collected 19 July 1972). Additional specimens have been deposited at Bode!{a :\Iarine Laboratory, Bodega Bay; the alifornia Academy of ciences, an Francisco: and at the British :\luseum (X atural History), London.

ETYMOLOGY. The specific name is derived from the Creek acora, meaning oscill at ory or pulsating movement.

SEASONALITY J\ND DISTRIBUTIO~. Adult medusae have been collected in the surface plankton, occasionally in moderate numbers, from June th rou~h K"ovember. \Yhile po55ibly more widespread alonA the California Coast they are known only from the adjoining Bodega and Tomales bays, Salmon Creek Beach, and the open ocean connccti ng these l wo areas.

R"CORDS OF COLLrCTIO;-; IX CALIFORNJ,\ . I. June 18. 1960 by G. 0. :\fackie a nd \\". Renshaw. Several ma­ ture medusae hand-dipped from the surface waters of T omales Bay: also collected from fresh beach drift, Dillon Beach, Cali­ fornia. 2. August 196 7 by cl ass, Bodega i\1arine Laboratory. Fifteen ma­ tme medusae co ll ected in a surface pl ankton tow in Borl cga Bay. 30 THE WASMA~~ JOUR~AL OF BIOLOGY, VoL. 34, ~ o. I, 1976

b 0

10 mm

FrcURE 5. T wo stages of gonad de,·elopmcnt in Aglauropsis aeora: a. gonad forming. not ripe; b. mature gonad present.

3. September 10, 1971 by ]. T . Rees. One mature medusa coiJected in a surface plankton tow in the open ocean just beyond Bodega Bay. 4 . October 18, 1971 by ]. T . Rees. Two medusae, not fully mature, collected in a surface plankton tow in Bodega Bay. 5. November (?) 1971 by W . Gladfelter. About eight mature medusae collected in the surface plankton in Tomales Bay. 6. July 5 through July 20, 1972 by C. 1\Iills. About 15 medusae in various advanced stages of development collected from fresh beach drift on Salmon Creek Beach. 7. J uly 19, 1972 by J. Rutherford. About 15 medusae in various advanced stages of development collected in surface plankton tow in Bodega :Bay. 8. June 23 , 1973 by C. Mills. Six medusae in various advanced stages of development coiJected in fresh beach wash on Salmon Creek Beach.

B EHAVIOR OF THE MEDUSA. The adult medusae assumed four basic positions in the laboratory. Tn the " fi shing" posture ( fig. 3) the tentacles of the animal are well extended, the oldest being held up, nearly vertically, next to the bell. The tentacles of inter­ mediate size are held at various degrees of horizontality; the youn­ gest and shortest are held vertically downward. The animals may LIMNOMEDUSAE-MILLS, REES & HAND 31

v mv np

FICURC 6. i'l:cmalocyst pads of Aglauropsis aeora at the bases of the ten­ tacles of a mature medusa· \", nlum; np, nematocysl pad ; mv, marginal \"esicle ; ex, ex umbrella. either maintain their position in the water column in this " motion­ less" position or slowly sink. I n the swimming posture the tentacles are fully contracted and are pushed in and out through the velar opening with each pulsation of the bell. There is an intermediate " prepa ratory" posture between these two positions in which the tentacles are held loosely around the base of the bell in preparation LIMKOMEDUSAE-M!LLS, REES & HA::\D 33 spent a considerable amount of time on the boltom of a beaker, with tentacles partially contracted. These behavioral stances are imilar to those described for Eperelmus t ypus C\laclJe and ~Iackie , 1963). Rate of pulsations for the adult medusae varied between 7 7 and II 0 beats per minute. The animal was propelled through the water from about 0.5 to 1.0 em with each pulsation. \\'hen several ani­ ma ls were kept together in a large beaker, they usually were all occupied with the sam~ activity (i.e. either swimming, " fishing," or resting on the bottom). Mechanisms by which activity sequences in individuals and aggregations of hydromcclusae are regulated arc unclear.

DISCUSSION .iglauropsis :\Hiller ( 1865) may now be recognized as containing five valid species: three from the south Atlantic Ocean (A. couantii Browne, 1902 ; A. jarli Kramp, 1955: A. kawari Moreira and Yama­ shita, 1972) and two from the north Pacific Ocean (...!. acora, this paper; and a misidentified species from Japan, " Eperelmus typus" of Japanese authors, see below). A sixth species, :l. agassizi :\1 tiller (1865) from Brazil, is too poorly described to permit recognition, and may be the same as A. kawari (l\loreira and Yamashita, 1972). Previous to our report, the genus has only been recognized from the south Atlantic (Kramp 195 7, 1959, 196 1). Uchida ( 1929) reported Eperetmus typus Bigelow (described from southern Alaska) from Japan, based upon three speci mens questionably collected at Hokkaido. While noting differences be­ tween the japanese material and Bil(elow's description, d1ida believed such distinctions were due to differences in developmental stages. t:chida ( 1940) and Xagao (1969a,b: 1973) subsequently reported E. typus from Akkeshi Bay near the Asamushi )farine Biological Station at Hokkaido. Kramp ( 1968: 141) discusses the amphi-Pacific distribution of E. typus. Epcrctmus Bigelow, a monotypic genus, is characterized by the possession of broad, flat, centripetal canals arising from the ring canal and projecting upward into the bell. ~ I ackie and ~ l ackie ( 1963) report small lateral pro­ jections, ' 1vestigial tentacle tips," on the tips of the more mature tentacles on a specimen of E. typus from San j uan Island. Ronald Larson (per~o n a l communication) has fo und similar stru ctures on 34 THE \\"ASMAJ'\l\ ]OURJ'\AL OF BIOLOGY, VoL. 3.J, )lo. 1, 19i6 specimens of E. t·ypus from Coos Bay, Oregon. However, we have not seen any " vestigi al tips" on the two specimens of E. typus we have examined (one adult specimen from San Juan Jsland, Wash­ ington, in the synoptic collection at Friday Harbor Laboratories, G. V. Mackie, collector; and an immature specimen from Yaquina Bay, Oregon, R. Larson, collector). After careful examination of material from Akkeshi Bay kindly sent to us by Dr. Zen ~agao, it is evident that the Japanese specimens do not possess the centrip­ etal canals characteristic of Eperetmus. In the J apanese medusae, exumbrellar furrows of the older tentacles have apparently been mistaken for centripetal canals. We have concluded that the Jap­ anese Epcretmus typus of Uchida and Nagao is an Aglauropsis and that the names Eperetm.us tyjms of Uchida ( 1929, 1940) and ~agao (1969a,b; 1973) are homonyms of Eperetmus typus Bigelow. Ko specific name for the Japanese Agtauropsis is available and we be­ lieve the designation of a name for this species is better left to a J apanese worker. In this study, we wi ll now reier to this med usa as Aglau.ropsis sp. The following reports appear to be valid records of E. t·ypus:

Port Clarence, Alaska (Bigelow, 1920 and Thiel1 1932a,b); Mary Island, southern Alaska (Bigelow, 1915); Vancouver Island, British Columbia (Foerster, 1923); San Juan Island, Washington (:\Lackie and Mackie, 1963 and i.Vlackie and l\lackie, 1966- personaJ com­ munication ); and Coos and Yaquina bays, Oregon (R. Larson, personal communication ). Aglauropsis aeora is compared with A. conantii, E. t')'pus, and Ag­ lauropsis sp. in tables 2 and 3 on the bases of nematocysts, character­ istics of lhe polyps and medusae, and habitats. The major differences can be summarized as fo llows. Of the medusae being considered Eperetmus t·ypus is the only one with centripetal canals. .4glau- 1'opsis aeora differs from 11 . co11antii primarily in the shape of mature gonads and structure of the edge of the manubrium ( table 3; figs. 4 and SA). Mature gonads of rl. conantii hang from the radial canals as a wavy curtain with the di stal edge profu sely lobed (See plates in Brown and Kramp, 1939). :\Iature gonads of .-1. aeora hang as a wavy curtain with a simple edge. The lips of the manubrium of ;J. conantii are bound by a swollen band of nemato­ cysts, whereas the li ps of 11. acora are bordered by a row of spherical nematocyst batteries. Aglauropsis acora di ff ers from ilglauropsis TABLr. 2. Comparison oj 11ematocy~t-~ i11 adult medusae (1.5-18 11111~ diameter) of tlm•r sprrit!s of :\Jdauropsis and of Epcrelmus typus . .-11/ spuime11s preser~ed in fnrmalill, all nu11atocysii mcasurrd wzcrplodl'd. All meoutrrmmts art rrprrssrd i11 p.m.

:o\ematoc>··t pad at T~ntadrs ~lonuL:-h:rn lit" Esumbrell• ubumbrdla Ia-• <•I t

(primary . ltltltllopliJ elltlfl.l pulyp} micr~ba.'ic 13.0--15.0 X ·u.o 14.0 x ? s.s- 9.s x no ?13.5- 15.0 X 6.5- 8.5 X •ur)"ttl• s.o- 9.5 7 5- 8.0 4.0-- 5.5 nematoc-y,a, i .5· b.5 4.0- 5.0 (J.S 10.0 1' 3.5- 6.0 l>ttrhlt.ns S.0-10.5 X 6.0 7.0 X ? 4.0- 5.5 X 7.5 9.5 X none 3.5 5.0 3.0 4 0 2.0 3.0 4.0 5.5 A. cmwntii microbnsic 1·1 .0- 16.0 X 11 .0- 14.0 X * 12 .5 14 .0 X no nuL polyp cur)'telc~ 7.0 9.5 6.5- 7.5 7 ..1 - 8.5 nemntocysls rxa mi1H'cl not knnwn i sorhiz u.~ • ? 5.5- 8.5 X *? (>.5 S.S X ? 8.5- II.SX 3.0- 5.0 3.0- 4.0 4.0- 5.0 .1. 'fl (Japnn) nticn,IJ:u:ic 13.0--15.0 X 9.0 10.0 X nc)ne no 11.0-13.5 X euryte1<" 6.5- 8.0 s.o 6.01 nC"rnatocyst~ 6.0- 7.0' i•·nrhiliS i.O-- 9.5 X nont s.s 10.5 X none l.O 5.0 4.0-- 5.0 £prrriHIU1 I7PNJ microba>ic 16.tHS.S X 11.5-H.SX ? 8.5 IJ.S X · : 16.0 19.0X nul polyp tur)"tr1<> i.O-- 9.0 6.5- 7 .0' 5.0- 6.5' i,$- 9.5 rumint n::mc no n ~ 6.0-- i .0 m:tnob., .. ic none nune • ?20.0--22.0 X none eur}"ttl,.. 10.5-11.0'

.. ~ot common 'Nagao. 1969 . "Snmr ns heterotrkh• ol :\Iackie and Madie, 1963. ? Kone lirt

Adull m<·dusa

Ratio nl ...., jrllr ;:t ( 16 mn1 :o.s)t•dntcn.') c:1 cog<· of lhicknt" Ccntril>eto l manubrium hll>l.'ll Ttntou.J~, !\tatocy3ts ;:..< Polr1• cana l ~ Gnnath l'cdunck Si

A -~8 THE \\"ASi\lA:\X JOURX:\L OF BIOLOGY, \'or .. J ~ , :\o. 1, 19i6 sp. in its lack of a peduncle and perhaps its primary polyp (the primary polyp of Aglauro psis sp. has not been described) . agao (1969a, 1973) has described small coloni es of .IKlauropsis sp. polyps, each with one filiform tentacle and bearing one medusa bud at a time. \\·hether further development of the primary polyps of !l. acora would reveal a simi larity with the polyps of AKlauropsis sp. is unknown. All four species of medusae are separnblc on thr basis of their cnidoms, particularly in a comparison of the nematocysts of the exumbrella (table 3). We found that the adult animals always had exumbrellar surfaces heavily covered wi th nematocysts. Si nce all exumbrellae examined were from formalin-preserved animals, no exploded nematocysts were seen ; the identification of nemato­ cyst types is based on comparisons with capsules located elsewhere in the . It is interestin!.{ to note the presence of what seem to be macrobasic euryteles in the exumbrella of a mature specimen of E. t·ypus (fig. 8B). This uncommon nematocyst type has previously been found in the family Olinrliasidae only on the cxurnbrellae of newly released medusae (\\'erner, 1965). They are apparently not replaced after their initial use. Since they are relatively uncommon on the mature E. typus, it is not known whether or not they are the remnants of the original complement of nematocysts. The subumbrella of E. typus, in the one juvenile we were able to examine, was also very sparsely provided with nematocysts. None of the observed subumbrcllar nematocysts were fired, so we can only tentatively call them microbasic euryteles. ·one of the other species appear to bear nematocysts on thdr subumbrellar surfaces. One specimen of A. acora had two clusters of exploded isorhizas on its subumbrella, but since no unexploded nemato­ cysts were seen. these are assumed to be souvenirs of an encounter with another cnidarian. \Ye could explain some exploded steno­ teles found in a localized area on the exumbrella of another specimen of .1. acora in the same manner. All four species of medusae also have very noticeable pads oi nematocysts at the bases of the tentacles. ln . 1. acora, E. typus. and !lglauropsis sp. the pads of the largest tentacles become cleft in the center (f i ~. 6). T hose of tl. ronantii never appear cleft and may be slighlly more swoll en ( this ma y be an nrtifact of the Lll\1:'1/0MEDUSAE- i\IILJ.S, REES & H:\::'110 39

preservation). imilar structures appear in other olindiasid me­ dusae: crescent-shaped nematocyst pads at the tentacle bases of .lglauropsis kawari C~I oreira and Yamashita, 1972), a marginal nematocyst rin~ in Craspcdacusta sowabii, a large cushion of nematocysts at the base of each maq.,rina l tentacle of Gossca cory­ nctcs ( Russell, 1953), marginal nematocyst pads in Gossca braclz-y­ mcra ( Kramp, 1961). and basal tentacle hulbs packed with ne­ matocysts a t the bell margin of Vallcnt inia aclhcrcns (Hyman, 194 7) a nd Vallcnl inia gabricllac ( Vannucci -l\lendes, 1948) . Swel­ lings at the bell margin of singu/aris (i\l ayer. 1910) and saclike swellings at the tentacle bases of Gonionrmus vcrtcns (Rus­ sell , 1953) are probably related structures. The primary or mature polyp is known for eight species of olin­ diasicl medusae. K agao ( 1973) lists M onobrachium parasilum, Gonioncmus vcrlcns, osltoro, sowcrbii, Olindias plrosplrorica, and "Epcretmus typus'' from J apan. Ad­ ditionally, the polyp of ratlenlinia f(Obricllac ( \'a nn ucci- ~l endes, 1948 : Fosler, 1971) and the primary polyp of A . acora arc known. The primary polyp of .-l. aeora most cloS<'ly resembles that of the freshwater jellyfish Craspedacusla sowerbii in its lack of a visible perisarc and tentacles and the placement of nematocysls al the distal end of the polyps. All known polyps in the family Olin­ diasiclae have only one kind of nematocysts, microbasic euryteles.

SuMMARY l. A new limnomedusa in the famil y Olindiasidae, ..J glauropsis acora, from the plankton off cen tral California, is described. 2. Primary polyps of ..J. aeora l ackin~ tentacles were raised in the laboratory, but further attempts to culture tbe polyps were unsuccessf u I. 3. Comparisons are made between A. acora, A. COIIalltii, .-lglaurop­ sis sp., and Eperetmus typus. 4. Eperclmus typus of Uchida and "agao from Japan is found to be a different species tban described by Bigelow ( 1915) and should be placed in the genus Aglauropsis. 5. The cnidoms of . l. acora and :1. ronantii are described and those of EjJerctmus t ypus Bigelow and Aglauropsis sp. fr om Japan are amended. -10 THE \\":\SMA..'\::\ JOUR.X:\L OF BfOJ.OGY, \'or.. J -1 , :\o. 1, 1976

A cKNO\\'LEDG~tENTS The fu llowing colleagues kindly provided medusae to us for study: Dr. Paul Cornelius, British ~lu seum (Katural H istory): Dr. Dennis \\"illows, Friday Harbor Laboratories, University of \\"ashin~ton: Dr. Zen :\agao. Kushiro Branch, Hokkaido 'Lni ver­ sity of Education ; Dr. G. S. ~ f oreira, L"niversity of Sao Paulo; D r. William Gladfelter, Pacific ~Iarine Station of the University of the .l'ucific; Dr. ·ward Renshaw, California State University at Long Beach ; Ronald Larson, Smithsonian lnstitution. Dr. G. 0. :\Jackie was helpful in locating speci mens. james Carlton provided helpful suggestions and reviewed the manuscript. We are also grateful to members of the Classics Department of the t:niversity of \\'a hin~ton for help in choosin!{ an appropriate name.

LITERAT URE C TTED

BtCCLOW, H. B 1915. Eprrclmul, a new genus of Trachomrdusae Proceedings of the United Stales Xational Museum 49:.\99--10-1 19'0 l\fc>du ac and Ctenophorae. Reporl of Lbc Can:~di rm Arctic Expe­ dition, 1913-1918. Southern P:m~. 1913- 1916, 8 part R :23 pp.

BROWN! I E. T . 1902. A preliminary report on lhe hydromedusnc from the Falklnnd Islands. Anna ls and Magaziuc of Natural History, Series 7, 0:272-284.

B RO W '\'£, 8. T., and P. L. KRA:ill' 1 9 .~9. llydromcdusae from the F alkland lslands. "Di!Cot•rry" Reports 18: 26.5- Ul. f'OERSTI'R, R E l9lJ. The hydromedusae of the \\est coast ol ::\"orth America, with special retcrc-ncc to those oi the \ ancouHr I land restion . Contributions to C.uradian Biology and Fisheries, :\c-w Series. I. 2 19-281.

F OST!;R, X. R 1971. l'allcutmi.1, the bitcb-hikinl! jcll~fi,h Frontie-r' .\6(1):14-19. lint '~ • I. 11 19-17. Two new hydromcdusae from the Californi:1 con. l. T ransactions of the American Microscopical Society 66 (J) :262- 268.

Krt,\ ;\1 P, !'. 1. . 195 7. J'l ydl'()tncdusac from the "Di.I'Cotl('ry" colil·ctions. "/Ji,,CIJVI'I"J'' Reports 21): 128 pp. LIMNOMEDUSAE- MTLLS, REES & HAND 41

1959. The hydromcdusac of the Atlanti c Ocean and adjacent waters. Dana­ Heport Carlsberg Foundation 46:283 pp. 1961. Synopsis of the medusae of the world . Journal of the Marine Bio­ logical Association of the United Kingdom 40:429 pp. 1968. The hydromcdusae oi the Pacific and Indian Oceans. Sections II and liT. Dana-Report Carlsberg Foundation 72:200 pp.

M ,\ CKI E, G. 0., and G. V. MACKlE 196.1. Systematic and biological notes on living hyd romcdusac from Pugct Sound. National Museum of Canada l3u llctin ="Jo. 199, Contri butions to Zoology: 6,1-8-1 .

M A'I:J::R, :\. G. 1910. Medusae of the world. Volume IT. Hydromedusae. Carnegie Institute of Washington, Publication :-\o. 109:498 pp.

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