2172 A new hydrozoan,Geomackiea zephlrolata gen. nov., sp. nov. (Anthomedusae: ),from inland marine waters of British Columbia and WashingtonState

Clnupn E. Mn-ls Friday Harbor Laboratories, University of Washington, Friday Harbor, WA, U.5.A.98250 Received December 14, 1984

Mrls, C. E. 1985.A new hydrozoan,Geomackieazephyrolatagen. nov., sp. nov. (Anthomedusae:Pandeidae), from inland marine watersof British Columbia and WashingtonState. Can. J. Zoo1.63:2172-2175. The hydromedusa Geomackiea zephyrolata gen. nov., sp. nov. is described and illustrated. It has been placed in the subfamily Protiarinae of the family Pandeidaeon the basis of its stomachand gonad morphology and the four large perradial tentacleswith conical basal bulbs. A new has been erectedbecause the medusais supplied with four broad interradial bulbs, eachrimmed by up to eight closelypacked solid tentaculae.A total of 29 specimenswith bell heightsof 0.9-3.5 mm have been collected in the plankton of Saanich Inlet, British Columbia, and Friday Harbor, Washington, between 1978 and 1981.Juvenile as well as adult specimensare describedand the cnidome,gametes, seasonal and vertical distributions,and taxonomic affinities within the family Pandeidaeare discussed.

Mu-t-s,C. E. 1985.A new hydrozoan,Geomackiea zephyrolata gen. nov., sp. nov. (Anthomedusae:Pandeidae), from inland marine watersof British Columbia and WashingtonState. Can. l. Zool.63 2172-2175. On trouvera ici la description illustr6e de I'hydrom6duse Geomackieazephyrolata gen. nov., sp. nov. La morphologie de I'estomac et des gonadeset la pr6sencedes quatre grands tentaculesradiaux i bulbes basaux coniques placent cette meduse dansla sous-famille des Protiarinae, dans la famille des Pandeidae.Un nouveaugenre est propos6parce que la m6dusepossdde quatrebulbes interradiaux larges, chacunbord6 de tentaculescompacts et sen6sles uns sur les autres(usqu'd huit). Vingt-neuf sp6cimensde ce nouveau taxon, dont la hauteur de l'ombrelle varie de 0,9 e 3,5 mm, ont 6t6 r6colt6s dans le plancton de SaanichInlet, Colombie-Britannique,et de Friday Harbor, Washington,de 1978e 1981.Les jeuneset les adultessont d6crits ici; le cnidome, les gamdtes, la r6partition saisonnidreet la r6partition verticale de m€me que les affinit6s taxonomiques au sein de la famille des Pandeidaefont I'obiet d'une discussion. [Traduit par le journal]

Introduction This was the only Geomackieazephyrolata gen. nov. , sp. nov. speci- men seen during 7 years of extensive day- and night-time inspection During a2-year investigation of the medusa, siphonophore, of in Friday Harbor from 1977 to 1984 (Miller 1979; Mills and ctenophore fauna of Saanich Inlet, British Columbia, l98l; C. E. Mills, unpublished data). The specimenwas collected in Canada, 28 specimensof a new anthomedusawere collected, a glass bowl and was maintained in the laboratory for several days and an additional specimenwas later taken nearbyin the waters before its preservation. of Friday Harbor, Washington, U.S.A. The speciesis placed in Nematocystswere examinedin fou specimensby using differential the family Pandeidaebecause of its quadratemouth lacking oral interference contrast optics and were identified according to the tentacles,its four adradial gonadsattached to both the stomach scheme of Mariscal (1974). At least 20 unfired nematocystsof each and the radial canals, along which they extend nearly half the type were measured at 1000x magnification; the numbers are pre- distanceto the bell margin in large specimens,and its four large sented here as size ranges, and were obtained from specimensmea- perradial tentacleswith typically pandeid conical basal bulbs. suring 1.3-3.0 mm in bell height. It is placed in the subfamily Protiarinaebecause it has not more than four tentacles.A new gents, Geomackiea,is established Results on the basis of its four distinctive interradial marginal bulbs, Geornackica gen. nov. each of which is rimmed by up to eight closely packed ten- taculae. This type of marginal bulb structure has not been DrAcNosIS:Pandeidae with four penadial tentacles having describedfor any other hydromedusa.The new species,Geo- tapering conical basesand with four broad, intenadial bulbs, mackiea zephyrolata, describedhere is the type speciesof the each rimmed by a cluster of short tentaculae. new genus. TypEspEcIES: Geomackiea zephyrolata sp. nov. ETyMoLocy: The genus is named in honor of Professor Materials and methods George O. Mackie of the University of Victoria, British Col- The SaanichInlet specimens'werecollected in 1978 and 1979 umbia. Professor Mackie has made numerous insightful con- duringa 2-yearseries of approximatelybimonthly stratified replicate tributions to our understandingof the biology ofjellyfishes and planktontows (Mills 1982)at Universityof Victoria Station"E' a wide variety of other marine organisms. (48"37.5'N, 123'30'W). (Onespecimen was collected in a plankton tow accompanyingthe aboveseries near Brentwood Bay in Saanich Geomackiea zephyrolata sp. nov. at 48'35'N, 123'30'W.) Verticalplankton tows using a 0.75m Inlet Undescribed anthomedusa,Mills 1982, pp. 58, 326, 336 diameter,333-pm meshclosing net weretaken over the following Hot-orypE: U.S. National Museum of Natural History intervals:0- 25,25 - 50,50 -7 5, 75 - 130,and 130- 180m. Saanich (Smithsonian Inlet is approximately200 m deepat the samplinglocation. Institution) USNM No. 72381. Collected by the The Friday Harbor specimenwas collectedfrom the Friday Harbor author in Saanich Inlet on June 14, 1978, at approximately Laboratoriesfloating docks (48'32.7' N, 123o0.7'W) at night, using 1530 in a vertical plankton tow which sampled between 130 anunderwater 100-W light bulbthat provided dark-field illumination. and 75 m deep. This specimenwas immediately preservedin 21'13

5Pt c a @ $@€ Frc. 2. Nematocystsof Geomackieazephyrolata. (a) Microbasic mastigophore(?) or eurytele(?), seenonly in the unfiredcondition; c occurringon both the perradialand interradial marginal bulbs, on the surfaceof the manubrium,and on the exumbrella.(b) Microbasic mastigophores,fired andunfired; occurring on the perradialtentacles andon the interradialtentaculae. (c) An undeterminedelongate type w of nematocyst,seen only in theunfired condition; occurring on or near the interradialbulbs and tentaculae, but their exactposition could not be determined.

without apical projection. Without a peduncle;stomach small, extending about one-third of the depth of the subumbrella; mouth quadrate, with four simple lips that are not armed with nematocysts (see below, Nematocysts). With four narrow radial canals. With four interradial gonads attached to and ringing the stomach wall and attached to the radial canals, along which they extend nearly half the distance to the bell margin, forming a pendant cruciform structure with lobed ends. With four large hollow perradial tentacleshaving scat- tered nematocystsand with tapering,conical basal bulbs. At each of the four interradii is a broad, flat bulb rimmed by five to eight closelypacked, solid, shorttentaculae which are cov- ered with scatterednematocysts (Fig. I c). Thesetentaculae are graduatedin size, with the longestoccupying the centerposi- tion, and decreasinglyshorter tentaculae adjacent to them. The tentaculaeare generallyheld curved up and outward againstthe bulb. Both the penadial and interradial marginal bulbs are Ftc. l. Geomackieazephyrolata. (a) Preservedjuvenile specimen pigmenteda dark gray-brown color. The velum is nalrow. of undetermined sex, collected April 21, 1978, in Saanich Inlet, The medusa is without statocystsand without ocelli. British Columbia, measuring l 3 mm in bell height and bell diameter. Description of juvenile specimens(Fig. Ia) (b) Living male specimencollected May 18, 1981,at Friday Harbor, were approximately I mm Washington, measuring 3.5 mm in bell height and 3.0 mm in bell The smallestspecimens collected diameter. (c) Enlarged view of an interradial marginal bulb with high and I mm wide and have a nearly sphericalbell with thin tentaculae extended (they are normally held curved up against the jelly. The stomach is short, with a simple quadrate mouth bulb). hanging down about one-fourth the depth of the subumbrellar space.Even in the smallestspecimens, the gonadaltissue is the perradial arms extendingabout one-quarter 5Va formalin and has preserved dimensions of 2.0 mm in both prominent, with the radial canals. Mature sperm or pri- bell height and bell diameter. of the distance along gonadsof specimensas small as pARArypES: Five paratypes, USNM Nos. 72382-72386, mary oocyteswere seenin the (seebelow, Gametes). The four perradial were collectedon April 2'7, 1978, May 18, 1978, May 19, 1. 3 mm in bell height are present, as are the 1978, May 5, 1919, and May 14, 1979.Two additionalpara- tentacles with conical marginal bulbs having two to four tentaculaeeach. type specimenshave been deposited at the British Columbia four interradial bulbs, these is without statocystsand Provincial Museum, BCPM Nos. 984-354-l and 984-355-1. The velum is narrow. The medusa They were collected May 14, 1979 and May 5, 1979, re- without ocelli. spectively. Further specimens are on deposit at the British Nematocysts Columbia Provincial Museum, BCPM Nos. 984-356-l to Nematocysts(shown in Fig. 2) were examined in four pre- 984-358-l (three specimens),and the Friday Harbor Labora- servedspecimens which hadbell heightsof 1.3, 1.5, 1.5, and tories Synoptic Collection (four specimens). 3.0 mm. The nematocystsizes and locationsare similar in all Description of adult specimens(Fig. /b) four specimens. The large perradial tentacles and the small Medusa with a rounded, bell-shapedumbrella, slightly interradial tentaculae are supplied with microbasic mastigo- higher than wide; the largest specimencollected was 3'5 mm phoreswhich measure8.0-9.2 x 3.6-4.0 pm (all dimen- high and 3.0 mm in diameterin life. Jelly of mediumthickness, sions given are those of unfired capsules). These tentacular 2174 CAN. J. ZOOL,VOL 63.1985 nematocystsare not clusteredinto batteries,but rather are scat- TasLp l. Vertical locations of 27 specimensof tered over the surfaces of the tentacles and tentaculae. The Geomackiea zephyrolata collected in Saanich (towing tentaclebulbs (penadialand intenadial) are supplied with large Inlet in closing-net plankton tows inter- vals are shown on the vertical axis) nematocyststhat look similar in form to the tentacular micro- basic mastigophores.Although none of the large nematocysts Interval No. of specimens on the bulbs were seenin a fired condition, the shaft and thread depthof capsuleare proportionallysimilar to those of the inside the tows(m) Day Dusk Night Dawn tentacularnematocyst, so the large nematocystsare provision- ally also called microbasicmastigophores. (Microbasic eury- o-25 2 2- teles also occur commonly in pandeidmedusae and would be 2s-50 4 t2 difficult to differentiate from microbasic mastigophores in 50-'75 J 4- the unfired condition.)The large nematocystsmeasure 14.4- 75- 130 2 4- 17.6 x 6.4-8.0 pm. A third typeof nematocystis associated 130-180 with the interradialbulbs and tentaculae. These elongate nema- NoTE: Only two adult specimens were collected in tocysts occur in a loose cluster of about 50, although their tows, one at dusk (50-75 m) and the other at night (75- 130 m). The data suggest no preferenceby G. zephyrolatator preciseposition could not be determinedfrom the squashprep- any specific depth during either the day or the night, and a arations.These nematocysts measure 5.6-7 .2 x 1.6- 1.7p,m; lack of any vertical migration. The oxygen content of the name at present, since none were water column below 130 m was loo low to support aerobic they are given no descriptive zmplankton. seen in a fired condition. Microbasic mastigophoressimilar to those on the tentaclesand tentaculae,but measuring8.0- 9 .2 x 4 .O- 4 .4 p"m,are also sprinkledover the surfaceof the Saanich Inlet, British Columbia, and from Friday Harbor, exumbrella,being more abundantnear the bell margin than on Washington.The medusashows strong seasonality; all speci- the upper portions. mens except one were collected between late April and early Most hydromedusae have a dense aggregation of nema- June of 1978, 19'79,and 1981. The remainingspecimen was tocysts at the edge of the mouth. In contrast,the lips of the collectedin late Septemberof 1979. mouth of Geomackieazephyrolata are not provided with nema- The vertical distribution of most of the residentzooplankton tocysts,but consistof an 80 pm wide band of fusiform cells. of Saanichtrnlet is high stratified(Mackie and Mills 1983),and These cells have a finely granular internal structure and are most species of hydromedusae, siphonophores, and cteno- presumedto be adhesive-secretorycells, basedon their mor- phores assume characteristic vertical locations in the water phology and location. Similar glandularcells have been de- column (Mills 1982).The 28 specimensof Geomackieazephy- scribed in detail by Bouillon (1966) in the oral regions of a rolata were collected in a series of closing-net plankton tows variety of hydrozoan polyps and medusae.Above the lips of and are somewhat unusual in that they show no evidence of G. zephyrolafa, the surfaceof the manubrium is provided with maintaining a preferreddepth during either day or night. There sparsely scattered microbasic mastigophores that measure is no evidence that this speciesperforms diel vertical migra- 6.4-1 .2 x 3.2 pm and look similar to the tentacularnema- tions (Table l). Geomackiea zephyrolata is also unusual in tocysts. A few of thesenematocysts were also observedon the bearinggametes at sucha young age(indicated by small size). gonad surface above the manubrium. It is more typical of hydromedusaein cold temperateregions to grow to nearly their adult size before beginning to produce Gametes mature gonadal tissue. The gonads were examined microscopically on the four The family Pandeidaeis a conglomerationof hydromedusae specimens whose nematocystswere measured. Three of the whoseinterrelationships are problematical. Russell (1953) pro- specimenswere males. Theseanimals, having bell heightsof posed the division of the Pandeidaeinto four subfamilies, the 1.5, 1.5, and 3.0 mm, all had large numbersof morpho- Amphineminae, Protiarinae, Pandeinae,and Calycopsinae, logically mature spermatozoain the lobed terminal portions of based on the morphology and numbers of tentacles in adult the gonads (the cruciform central portion of the gonad appears medusae.Of Russell'sfour subfamilies,the last, the Calycop- to be largely composed of vacuolated rather than proper sper- sidae,has been raised to family status(see Kramp, 196l). Arai matogenic tissue). Many of these spermatozoabecame free and Brinckmann-Voss (1980) further removed the genusllali- from the surrounding tissue in the squash preparations' The medusa from the Pandeidae to its own family, the Hali- sperm headsare typically hydrozoan, acorn-shapedstructures, medusidae,and divided the remaining pandeid medusaeagain measuring3.2 p,min height and2.4 pm in diameter,with two into four subfamilies. Their subfamiliesStomotocinae, Amphi- spherical mitochondria in the basal portion. The fourth micro- neminae, Protiarinae, and Pandeinaeare basedon the presence scopicallyexamined specimen was a femalewith a bell height or absenceof a peduncleand, as in the schemeof Russell of 1.3 mm. This had primary oocytesin the gonads. (1953), on the numbersof tentacles. Pro- Hydroid Geomackiea zephyrolata belongs to the subfamily Arai and Brinckmann-Voss Unknown. tiarinae, which is defined by (1980)as includingthose pandeid medusae without a peduncle ETyMoLocy: The specific name zephyrolata means "borne and having four perradial marginal tentacles.As the subfamily by the west wind," and is derived from Zephyros, the ancient Protiarinae was not representedin either the British fauna Greek personification of the west wind and latus, past pat- (Russell 1953) or the British Columbia fauna (Arai and ticiple of the Latin verb fero, to bear. Brinckmann-Voss1980), affinities within this group have not been discussedby previous authors. A review of the literature Discussion yields eight generaof Pandeidaethat qualify for the subfamily Geomackiea zephyrolata has been collected only from Protiarinaeas defined by Arai and Brinckmann-Voss(1980) MILLS 2175

(the last three generalisted possesssecondary marginal cirri or This work was supportedby a University of Victoria graduate tentaculaein addition to the four perradial tentacles): Cnido- fellowship and by National Science Foundation grant OCE tiara, Paratiara, Protiara, Urashimea, Zancleopsis, Hali- 8214058. tiara, Halitiarella, and Geomackiea (Kramp 1959, 1961, 1968; Bouillon 1980; the presentpaper). With the exceptionof Urashimea, these genera further have in common a simple AneI, M. N., andA. BnrNcrueNN-Voss.1980. Hydromedusae of mouth and smooth (not folded) gonads. Urashimea, in con- British Columbiaand PugetSound. Can. Bull. Fish. Aquat. Sci. trast, has a folded mouth, folded gonads,and an unusualcom- No. 204. plement of nematocyststhat may ultimately place it in its own BouILLoN,J. 1966.Les cellulesglandulaires des Hydroideset des family (S. Kubota, personal communication). Kramp (1959) Hydrom6duses.Leur structure et la naturede leurss6cr6tions. Cah. Biol. Mar.7: suggeststhat the genusProtiara is the most primitive of the 157-205. 1980.Hydrom6duses de la mer Bismarck(Papousie, Nou- Pandeidae. I would further suggest that all members of the velle-Guin6e).III. Anthom6dusaeFilifera (-). Protiarinaebe consideredrelatively primitive within the family Cah.Biol. Mar. 2l: 307-344. Pandeidae. KRAMI,P. L. 1959.The hydromedusaeof the Atlantic Oceanand Medusaein the remaining three subfamiliesof the Pandeidae adjacentwaters. Dana-Rep. Carlsberg Found. No. 46. may generally be characterizedas having large, highly folded 1961.Synopsis of themedusae of the world.J. Mar. Biol. mouths, and gonadswith prominent folds or reticulae on their Assoc.U.K. 40:l-469. surface. The small, simple mouth and smooth gonadsthat are 1968.The hydromedusaeof the Pacificand IndianOceans. typical of members of the Protiarinae are, hbwever, also SectionsII andIII. Dana-Rep.Carlsberg Found. No. 72. presentin the pandeid generaMerga (whose medusaehave 4, MACKIE,G. O., andC. E. Mrus. 1983.Use of thePrscas IV sub- mersiblefor zooplanktonstudies in coastal 8, or more tentacles).Pandeopsis (with 8-16 tentacles), and watersof British Col- umbia.Can. J. Fish.Aquat. Sci. 40:''763-776. (which has 12 tentacles). Whereas, on the basis of MARISCAL,R. N. 1974.Nematocysts. ftr Coelenteratebiology: Re- genera tentaclenumber, all three of these fall into the subfamily views and new perspectives.Edited by L. Muscatineand H. M. Pandeinaeas it hasbeen defined by Arai and Brinckmann-Voss Lenhoff. AcademicPress, New York. pp. 129-178. (1980), their mouth and gonad structures suggest that these MILLER,R. L. 1979. Spermchemotaxis in the Hydromedusae.I. species may be anatomically intermediate between the two Species-specificityand spermbehavior. Mar. Biol. (Berlin), 53: subfamilies Protiarinae and Pandeinae.It seemsmost prudent 99-114. at this time, however, to leave these specieswithin the sub- Mru-s, C. E. 1981.Seasonal occunence of planktonicmedusae and family Pandeinaeuntil further comparative work is done. ctenophoresin theSan Juan Archipelago (NE Pacific).Wasmann J. Biol. 39:6-29. 1982. Patterns Acknowledgments and mechanismsof vertical distributionof medusaeand ctenophores.Ph.D. dissertation,University of Vic- I thank Don Horn, Alex Hartley, and Bob Baden of the R.V. toria.Victoria. B.C. John Stickland for assistanceduring the more than 600 plank- RusssLL,F. S. 1953.The medusaeof the British Isles. Cambridee ton tows in Saanich Inlet. and Richard Miller. who collected UniversityPress, London. the specimenin Friday Harbor and brought it to my attention.