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" ;"""',''.1.( '.....,'..'" : .." ',' ."'..,-'.;; .'vi;;,", Nc,:.':Y',' .' j",;")c';.:;i; .,.'.." ':, " '\i'." '".,' .; . ,''i;\.' :, '. ,"""',),;:.',. ..',......., "'. 'iC.':,::"",/;',.., . ';"i';.. ':', ",/j:;",.,,;,(:r. ';:iY';:'i,/;'," . ,~:". , ':,:,:~, .;..': '.',?' ,";:,\; ..\/\' .,~.,::;.?,(:: "';"":"";'; ;.1. ,it, i\,;;' ,'i'. "i'i~:"çl.: .1' l.I, ·. '','""/''::;'(';, :a:~':S:c"i""S;;;;' d,;"""',':;,,;'/L '.', "'\,¡ "'.\t",")ç;,,.:/;,' ........';;:;\!'.,/:.:~:.~;,.; " Y:':/,':: \:::' ..;;'f:-" rfJ . I:,'.~ ~;'i~; '..i)).........'..:,'.., d . .'-C'\".:,"" ',i,' :,); , ,:',,:, ':';;",/,:;)' '! ....,...."'.:,. "'!'.'.,,:- ' ".,' .'-'., ",(., '." ,ii-",.,;",:, ,;t,:;: ','i,c"i'L,D ¡SlrJ.li,lJt¡Qr(,dân,~;Taxa;n.øta)'~"'.Óf.__.¡ZO'øJ.tl,a,nldOI1 ,,¡ n.thf;,l,l IiÇJr~ff;:':,~~l ,'; , ':". ,L"',f . "',iV, '.i/":""" i,"',,' ;":,,iL " d,"','.',':,:(;' '; : d '.....",".:,,:":';.,",: ,);". ?: ;2)":,: '. ,., ':, "','x:, ...,.... ")':E~é, ""';):, ~" 'y/,s!?;;"n : '::;!D:f¡~;¿L', .,¡e:,;,. ',' ," ';' .,', .' ..'".' "');ti:'.\~~;fîi.'i¡\ M:dil~~r'~~0~~\;I;;' . ..'N ;'..d;'i'" ;);':",,;'..'i,:',i ..,,:,;;''~;:(~!;~~~~'~),Ä':f~J~~.~rn. ',:"'., ,'" . ... ,.:;;/',. " , .. .' """'. . . ...'..... , ...:::ê..' ,( ..'. .' ".:':"'...;"',...../.."'."......;,:..,,..".:.,.,...:..~'c:':...,..'.,':.';:',.:.,,'....d:,.'...:d '''.''.'','..'';;'''''''"::::' ':';.- ,,'.' ",'.,. .':.' ". ," ..: ."'''', ':,Y:..',:'. ',' ::"\':':c: ':.'.,.,......: ,,'_.., -'..,;:..~:.;_-,",.",:,.,._,~:.":.~::--.-....:..,:.._':;,',',.~:,:-''.',', '..-,_-:':~ "., . \",l.. ,.' ., ':'_'_'::~::-';'~.'_' ":':' .......' ":'-" J, ,"., ~ . WHOI-91-26 Distribution and Taxonomy of Zooplankton in the Alboran Sea and Adjacent Western Mediterranean A Literature Survey and Field Guide. by Laurence P. Madin Woods Hole Oceanographic Institution Woods Hole, Massachusetts 02543 September 1991 Techncal Report This report is submitted in fulfillment of a subcontract from the Harbor Branch Oceanographic Institution, Ft. Pierce, Florida, to the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts. Funding was provided by Grant No. N00014-91-C6007 from the Naval Oceanographic and Atmospheric Research Laboratory to the Harbor Branch Oceanographic Institution. -~----~ ~rr -----~== :: I" I' õ= .J , :c= tr m:¡~C1 .. _rn Submitted to Harbor Branch Oceanographic Institution on March 25, 1991. Reproduction in whole or in part is permitted for any purpose of United States Government. This report should be cited as: Woods Hole Oceanog. Inst. Tech. Rept., WHOI-91-26. Approved for publication; distribution unlimited. Approved for Distribution: ;:_ CI ::_ 0 CI _C1 Peter H. Wiebe, Chairan Department of Biology the 2 Abstract. This is a survey of literature records for occurrence and taxonomy of zooplankton in the Western Mediterranean, with particular emphasis on the Alboran Sea. It is intended to give a general background on the fauna, and facilitate identification of specimens collected or observed. A description of the hydrography of the Alboran Sea is followed by a general account of zooplankton biomass distribution, and more detailed lists of the occurrence of 361 species of medusae, siphonophores, ctenophores, worms, tunicates and crustaceans in 7 regions of the Western Mediterranean. Bioluminescent properties of the organisms are indicated where known. An ilustrated taxonomic guide provides capsule descriptions and ilustrations of 254 of the listed species. Key Words. zooplankton, Alboran Sea, bioluminescence 3 Table of Contents Abstract 2 Introduction. 4 General distribution patterns. 6 1. General hydrography 6 2. Distribution of zooplankton biomass 7 Occurrence of zooplankton groups 1. Colonial radiolaria and acantharia 2. Hydromedusae and scyphomedusae 3. Siphonophores 4. Ctenophores 5. Polychaetes and nudibranchs 6. Pelagic tunicates 7. Crustaceans Illustrated systematic guide Hydromedusae and Scyphomedusae Siphonophores Ctenophores Polychaetes and Nudibranchs Pelagic Tunicates Crustaceans 9 9 11 18 22 25 27 31 36 37 71 91 101 105 121 Acknowledgments 139 References 139 4 Introduction. This document is a literature-based survey of the occurrence and taxonomy of zooplankton in the Alboran Sea and adjacent regions of the western Mediterranean. It's purpose is to provide background on the kinds of plankton that one would expect to encounter in this area, and a convenient reference for shipboard identification of collected or photographed specimens. Because it is intended to support in-situ investigations, by submersible and SCUBA diving, of luminescent organisms, the taxonomic guide focusses on the gelatinous macrozooplankton and the more common crustaceans. It emphasizes characteristics of intact, live animals, and indicates whether they are known or suspected to be luminescent. The western Mediterranean Basin is divided into several regional seas, as ilustrated in Figure 1. The present survey includes distributional records for zooplankton in the: a. Alboran Sea - extending from Gibraltar eastward to approximately 0° longitude; b. Strait of Gibraltar; c. Catalan (Balearic) Sea - between the southeast coast of Spain and the Balearic Islands; d. Gulf of Lyon - extending southeast into the central basin west of Corsica and Sardinia; e. Ligurian Sea - between the French Riviera and Corsica; 1. Tyrrhenian Sea - bounded by Corsica and Sardinia on the west, Italy on the east and Sicily at the south; g. Adriatic Sea - between Italy and the Dalmatian coast. ...... :':-'. :.... . .... ~5° .... . ................. ..................... . ...................................... ......... '.. .................................................. ................................................ ................................................. .............................................., ..............-.. ............................ .," .. ".' .' . . '. '. .. -.................. .":":::::"' : ,', ~ :,'::,:,:' Mer CotoloRe 4Ô WO :.. .......-." '. 30° o Figure 1. Regions Furnestin, 1968) of the 10 Western Mediterranean Basin (from 5 The extent to which the planktonic fauna of these regions has been studied depends partly on the geographic distribution of marine laboratories on the coasts of these seas. Upwellng regions near Messina, Naples and Nice in the Tyrrhenian and Ligurian Seas have been known since antiquity. Laboratories have been established in these regions for over a hundred years, and the fauna is quite well known. Other laboratories in France and Italy have supported surveys in the Gulf of Lyon, the Catalan Sea and the North African coast. In addition, several oceanographic cruises have been undertaken in the western Mediterranean, adding coverage of the regions further offshore. There is a fairly considerable classical literature on the planktonic fauna of the Mediterranean, based on work done in the mid to late 19th century at Messina, Naples, Villefranche, Trieste and a few other locations by pioneers like Brandt, Chun, Haeckel, Lohmann and others. A valuable and comprehensive systematic treatment of phytoplankton and zooplankton in the Mediterranean, the "Manuel du Planctonologie Mediterraneenne" was published by Gregoire Tregouboff and Maurice Rose in 1957. It is a quite inclusive work, summarizing the basic biology of each group and providing keys and illustrations for identification. It is somewhat cumbersome to use in the field however, because of the complex structure of the keys and the separation of the illustrations from accompanying text (including captions) in a separate volume. This work, and some of the old literature, has been used here as a source. For the most part, however, the present survey is based on more recent investigations that used modern techniques for sampling zooplankton from larger areas and depth ranges. These studies also have the advantage of using a taxonomic nomenclature fairly well settled by major revisions published in the last several decades. Another relevant source of information for this survey are the reports of observations made from other submersibles and bathyscaphes. French scientists made numerous dives in the Gulf of Lyon and Ligurian Sea during the 1950's and 1960's (Bernard, 1955, 1958; Tregouboff, 1956, 1957) and more recently (Laval and Carré, 1988; Laval et al. 1989, Mills and Goy, 1988; Biggs et aI., 1987). Although these reports provide mainly qualitative visual observations, the sightings have been included in the distributional lists and discussions where possible. This survey is organized into three main sections. The first considers general patterns of zooplankton distribution. This is intended as an overview of hydrography, zooplankton biomass distribution, seasonal abundances and vertical zonation in the Alboran sea specifically, and in the adjoining regions. The second section considers the occurrence and abundance in the western Mediterranean of the major groups of zooplankton with emphasis on gelatinous forms and bioluminescent species: colonial radiolaria, hydromedusae, scyphomedusae, siphonophores, ctenophores, some polychaetes, some molluscs, pelagic tunicates and some crustaceans. Groups with no known bioluminescent species, notably the pteropods, heteropods, and chaetognaths, are not included in 6 this survey; neither are adult or larval fishes. Cephalopods, although luminescent have not been included for lack of time and space, and because they are thought unlikely to contribute significantly to luminescence observed from the submersible (E. Widder, pers. comm.). Occurrence in the western Mediterranean of a total of 361 species is summarized in 7 tables. Species are listed alphabetically within Class, Order or Suborder, as appropriate. Abundance and vertical distribution of the most common species are discussed in more detaiL. The tables also indicate whether the species is bioluminescent. The letter "a" in the "Lum" column means the genus is considered "definite" in the list of Herring (1987). The letter "b" indicates a genus is considered "uncertain" and the letter "c" indicates that the particular genus is not known to be luminescent, but one or more other genera in the same family is. A blank in the "Lum" column indicates no mention in Herring (1987). The third section is a taxonomic guide ,designed to facilitate rapid field identification of animals collected by divers or a submersible, or photographed or videotaped in situ. Instead of keys, brief descriptions accompanied by line drawings are arranged in the same order as they appear in the tables of distribution. The illustrated guide includes 254 (70%) of the species listed in the tables. For each species, two higher taxa (Family, Suborder, Order, Subclass or Class) are listed to place species in context of their classification. It is hoped that acccurate identifications can be made fairly quickly by flipping through the pictures. Because the majority of Mediterranean species also occur in the Atlantic and elsewhere, this part of the survey should prove useful in other oceans as welL. General distribution patterns 1. General hydrography The Alboran basin is relatively shallow, exceeding 1000 m only at the east and northeast. On the south it is bounded by a plateau stretching between Oran (Algeria) and Cabo Tres Forcas (Morocco). On the north, banks exist southeast of Malaga and southwest of Almeria (Spain). As the entry point for Atlantic waters into the Mediterranean, the Alboran Sea is strongly influenced by incurrent water masses. Circulation in the Alboran and western Mediterranean is discussed by Furnestin (1960) and Allain (1960); this brief outline is taken largely from the latter source. The principal Atlantic surface current entering through the strait of Gibraltar bears east-northeast, but soon curves to the right, taking a more easterly direction (see Fig. 2). Water in the lower edge of this current comes completely around, forming an anticyclonic eddy to the west of Cabo Tres Forcas. Currents in this gyre attain about 1.2 knots on the westerly side. The main current accelerates in passing over the ridge beneath the Isla Alboran, changes direction toward the north. A second anticyclonal eddy is spun off in the bight east of Cabo Tres Forcas; it circulates more slowly, at about 0.2 knots. Turning southerly again, the main 7 -+ ¥it.... lfi"ilN (0. 2 " I ~ ",ite.. ...i.. 1... n. H¥. 999999 Ii!... d. diy.rg.... Ii T T T T T T lig".. d. covergence i I 40 i -i-i I i S. i-o, (wJ i-,.i:, (G.J S. 10. Figure 2. ,Surface currents in the Alboran Sea and Western Mediterranean (from Allain, 1960). current passes close to the coast at Oran, then bears northeast, over deeper water, toward the Balearic Islands. A branch of the current continues to follow the north past Tunisia, and a large cyclonic eddy is produced on the north side of the main stream, within the bight bounded by Cabo de Gata and Cabo Palos in Spain. African coast The general pattern of surface circulation remains the same to a depth of about 200 m, though velocities are lower. Below 200 m, the water is mainly of Mediterranean origin, and a westerly current carrying Mediterranean water towards the strait of Gibraltar becomes established in the northeast part of the Alboran Sea. Below about 400 m, the circulation is reduced to almost nothing, with only the large cyclonic gyre east of Cabo de Gata and Cabo Palos still moving slowly. 2. Distribution of zooplankton biomass Biomass and diversity of zooplankton are generally higher than in the eastern parts, due largely to the influence of ,Atlantic waters. The surface waters (to about 8 200 m) of the Alboran Sea therefore have the greatest abundances and the most similarity in species composition to the Atlantic. Species composition is in most respects identical to that found outside the strait of Gibraltar. Both abundance and Atlantic character of the fauna are diluted as the suriace currents move east and northeast, so that the Ligurian and northern Tyrrhenian seas are poorer, and of a more Mediterranean character (Furnestin, 1968). Within the Alboran Sea, a divergence zone south of the Spanish coast was found by Rodriguez et al. (1982) to have a zooplankton community distinct from that of neritic waters to the north of it. They did not provide any data, however, on biomass distribution within these communities. Bracconot et al. (1983) provide some rather sketchy data from October and November, 1981, on total zooplankton biomass in the 0-200 m layer from stations both within the Sea and in the strait of Gibraltar. Lowest values, around 150 mg d.w. per m2, were found in the axis of the strait. Values of 500 mg/m2 for the 200 m water column were found in the northwest part of the Alboran. In the divergence zone south of the Spanish coast and in the southeast part of the basin biomass ranged from 200 to 500 mg/m2. Much of the zooplankton biomass in the east and southeast parts of the Sea was due to numerous Salpa maxima. \ , Sampling by Greze et al. (1983) on the Alboran (270 m deep) and Tofinio (90 m deep) banks in the southern part of the Alboran Sea indicated that zooplankton abundance (mainly copepods) was similar to that found in adjacent areas of open water. Numbers of individuals ranged from about 500 to 4600 per m3, and biomass from 22 to 100 mg (d.w.) m3 over the two banks. Zooplankton distribution along the Catalan coast near Barcelona was investigated by Sabates et al. (1989) between April and July, and September through October, 1983. They found greatest abundances in April and May, when biomass values were as high as 60 mg/m3 in the top 200 m that were sampled. Biomass decreased to about 12 mg/m3 by June and July, and reached a seasonal minimum of 4.5 mg/m3 in September, increasing slightly in October. Values were higher further from shore. Gelatinous forms were a major part of this biomass in the spring. Salps peaked in April and May, and doliolids in July. Medusae and siphonophores were present throughout the sampling period at about the same abundance. Euphausiids were most abundant in April and June, but copepods dominated the abundances in April, June and July. 9 Occurrence and distribution of zooplankton groups in the Alboran Sea and adjacent areas. 1. Colonial Radiolaria and Acantharia Radiolaria, both solitary and colonial forms, are widely distributed in all the world oceans. Colonial forms consist of hundreds of cells in a gelatinous matrix and can attain sizes of several cm. The Collozoum, Thalassicolla, Raphidozoum, Sphaerozoum, Acrosphaera, Collosphaera, Siphonosphaera and Cytocladus are bioluminescent (Herring, 1987). These organisms are readily recognized as radiolarians by their gelatinous or "fluffy" appearance, and some species have quite consistent appearances. The species listed in Table 1 are those reported from submersible obseNations. Bernard (1958) ranked the radiolarians, mainly colonial forms, third in abundance after copepods and other crustaceans in his visual census of the water column. They were found throughout the water column, to 900 m. i 10 TABLE i. RAIOLAIANS AN ACATHAIANS. Species Figu re Lu m Alb ora n Gibr aite r Geographic Occurrence Cat a ian Lyon X X X X a a Ligurian: X X X X a X References Lyon: X b streptacantha Bernard' 58, Franqueville ' 70 Tregouboff ' 56, , 58 Tyrr rian heni an acantharians Acanthometra sp. Arachnosphaera sp. Aulacantha scolymantha Aulosphaera spp. Collozoum spp. Myxosphaera coerulea Sphaerozoum spp. Spongosphaera Ligu X Adr iat ic 11 2. Hydromedusae and Scyphomedusae. There appears to be relatively little data on the distribution of hydromedusae or scyphomedusae within the Alboran Sea itself (Goy, 1983; Rodriguez, 1983), but there are several studies that consider seasonal and sometimes vertical occurrence of medusae from the Catalan Sea (Gil et aL, 1987, 1988), Gulf of Lyon (Casanova, 1970) Ligurian Sea (Goy, 1972; Goy et aL, 1989), Gulf of Naples (Vannucci, 1966; Brinckmann, 1970, 1987) and the Adriatic (Benovic, 1973a, 1973b, 1976, 1977; Vucetic, 1982). Probably many of these species are widely distributed throughout the Mediterranean, but simply haven't been as well sampled in the Alboran Sea as they have at Naples, Messina or Villefranche. Although Goy (1983) refers to the strait of Gibraltar as a "planktonic desert" and considers it a zoogeographic barrier for hydromedusae, most species known from the Mediterranean also occur in the Atlantic and elsewhere. Table 2 lists 104 species of hydromedusae and 9 species of scyphomedusae reported from the Western Mediterranean; of these 92 are described and ilustrated in Section D. The species are listed alphabetically within orders. The medusan species which appear to be most abundant in the Alboran Sea and adjacent regions are discussed here, with seasonal and vertical distributions, where known. Some hydromedusae noted as common in the Alboran area include Lizzia blondina, and Obelia spp., both abundant in March and April (Rodriguez, 1983). Goy (1983) reported 11 species in the Alboran Sea in autumn, of which Eucheilota paradoxica was most abundant, especially in the southwest part of the Sea. Numerous specimens of Pandea conica were collected in 1986 by divers in the Alboran (Harbison, pers. comm.). Persa incolorata was the only species found in any abundance in the strait of Gibraltar by Goy (1983). Along the Catalan coast, the commonest species collected in the upper 200 m during May and June were Podocoryne carnea, P. minuta, Lizzia blondina, Obelia spp. Eirene viridula, Aglaura hemistoma and Persa incolorata (Gili et aL, 1988). Spring and early summer appeared to be the times of peak abundance for the medusae in this area, with Lizzia and Aglaura occurring at densities of 10's m-3. Deeper collections were reported by Casanova (1970), who found a few species of trachymedusae and narcomedusae in tows as deep as 2000 m. Commonest was Solmissus albescens, a large, widely distributed and luminescent narcomedusa. This species occurs throughout the Mediterranean, and is a vertical migrator. In the Adriatic, populations of S. albescens migrate between about 600 m and the surface (Benovic, 1973). Mills and Goy (1988) characterize S. albescens as "the most numerous medusa in the mesopelagic western Mediterranean", and describe its vertical migration and swimming behavior as observed from a submersible diving near Villefranche. There the medusa moved from daytime depths between 400-700 m to the upper 100 m at night, swimming at about 100 m h-1. Solmissus has also been reported by other observers in submersibles as one of the commonest medusae seen (Tregouboff, 1956, 1957; Bernard, 1958). Laval et aL (1989) estimated densities of 15 to 208 Solmissus per 1000 m3. The abundance, 12 fairly large size (to 5 cm) and bright luminescence of a b this species make it likely to be an important source of midwater bioluminescence. c Sketches of its appearance insitu, as reported by Mils and Goy (1989) are reproduced in Fig. 3. Figure 3. In-situ appearance of Solmissus albescens The most (from Mils and Goy, 1989). abundant scyphomedusa from this area appears to be the ubiquitous and troublesome Pelagia noctiluca, a medium-size but strongly bioluminescent semaeostome. In recent years, populations of Pelagia have reached nuisance proportions in several parts of the Mediterranean. Gili et al. (1987) report maximium densities in the Catalan area of 30 m-3 in June. In the Gulf of Lyon and waters off Toulon, Franquevile (1971) found Pelagia migrated vertically between about 500 m and the surface. Individuals collected in April had bell diameters between 10 and 50 mm. Evidently, populations of Pelagia fluctuate on a cycle of approximately 12 years, going from almost none to very high densities (Goy et aI., 1989). Other scyphomedusae that appear fairly common in the western Mediterranean are Atolla wyvilei, and Periphylla periphylla, which do not migrate (Franqueville, 1971), but are found below about 500 m. 13 TABLE i. HYDRO- AND SCYPHOMEDUSA. Species Fig Lu m Alb ora n Geographic Occurrence Gibr alte r Cat a lan Lyon Liqu rian Tyrr heni Adr X X an iat ic HYDROMEDUSAE Anthomedusae Amhinema dinema M-l c rubrur M-2 c Amhinema rugosur M-3 c turrida M-4 c M-S c Amphinema Amphinema Bougainvillia ramosa Bythotiara murrayi radiatur tetrastyla Dipurena halterata Cytaeis X X X X M-6 Calycopsis 5 implex Calycopsis sp. Cirrholovenia tetranema Cladonema X X X X M-7 X X X , M-S M-9 X X X X X X M-IO X X Dipurena ophiogaster M-ll X X Ectopleura durortieri Ectopleura larynx Ectopleura M-12 X X X X X Eucodoniur brownei M-13 Euphysa aurata M-14 Halitiara formosa M-1S Hybocodon M-16 X Koellikerina M-17 X Leuckartiara nobilis Leuckartiara octona Lizzia blondina Li z z ia fulgurans M-1S a M-19 a M-20 b M-21 b X M-22 c X lifer fasciculata Merga tergestina Merga tregoubovii X X sacculifera Eleutheria dichotoma pro X X a X X X X X c X X X X X X X X X X X X X X 14 Fig Species Lu m Alb ora n Gibr alte r Cata Ian Lyon Ligu rian Tyrr heni an Merga violacea M-23 c Neoturris pileata M-24 c Niobia M-25 X Oceania armata M-26 X dendrotentaculata ic X X X X X Pandea conica M-27 Paragotoea bathybia M-28 c X X areolata X X X X X X carnea M-29 hartlaubi M-30 minima M-31 X minuta M-32 X Rathkea octopunctata Sarsia eximia Sarsia gerni fer a Sarsia prolifera Sarsia tubulosa Staurocladia iat X Octotiara violacea Podocoryne Podocoryne Podocoryne Podocoryne Podocoryne Adr M-33 X b X X X X X X X X M-34 X X , M-35 X X X M-36 X M-37 X X portmanni Steenstrupia nutans X M-38 X X Thamnostoma sp. X X X Tiaranna rot unda Tregoubovia M-39 Turritopsis nutricula M-40 Zanclea costata M-41 X f X atentaculata X X X X X X X X X X X Leptomedusae Aequorea aequorea M-42 Eirene viridula M-43 Eucheilota paradoxica M-44 a X X 15 Species Fig Lu m Alb ora n Eugyanthea inquilina Eutima gegenbauri Eutima gracilis Gibr alte r Cata lan Lyon Ligu rian X M-45 X Tyrr heni an X ic X M-46 X X Eutonina scintillans X Helgicirrha schulzei M-47 Krampella dubia Laodicea nept una Laodicea ocellata Laodicea undulata Lovenella cirrata M-48 X X X X b M-50 b M-5l b M-52 a X Mitrocoma annae M-53 c X Mitrocomella brownei M-54 c Obelia spp. M-55 a Octophialucium funerarium Orchistomella M-56 a Phialidium M-57 a Phialidium mccradyi M-58 a X X X X X X X X X X X X X X X X X X X X X X X Limnomedusae Gonionemus vertens M-60 Odessia maeotica M-61 Olindias phosphorica Proboscidactyla ornata Scolionema suvaensis X X a a X X graeffei M-59 X X M-49 Phialidium sp. Tima lucul1ana iat X Eutim sp. hemisphaericum Adr X X X X M-62 X X M-63 X X M-64 X X X 16 Species Fig Lu m Alb Gibr n r ora alte Cata lan Lyon rian Tyrr heni Adr X X X Ligu an iat ic Trachyredusae Aglantha digitale M-65 Aglaura hemistoma M-66 X X X X Amphogona pusilla X Arctapodema amplum Arctapodema australe M-67 Geryonia M-68 proboscidalis Haliscera bigelowi Haliscera conic a c X c X b X M-69 X M-71 Persa incolorata M-72 X b X X X X X X X X X X Ptychogastria asteroides X Ransonia krampi M-73 Rhopalonema M-74 c Rhopalonema velatum M-75 c Sminthea eurygaster M-76 c X M-77 a X funerarium X X M-70 Liriope tetraphylla X X X X X X X X X X X Narcomedusae Cunina glòbosa Cunina sp. Pegantha rubiginosa Solmaris flavescens Solmaris leucostyla Solmaris solmaris Solmissus albescens Solmundella bitentaculata a X M-78 X X M-79 X M-80 M-81 X X X X X X X M-82 a X X X X M-83 c X X X X X X 17 Species Fig Lu m Alb ora n Gibr aite r Cata ian Lyon Ligu rian Tyrr heni an Adr iat ic SCYFHOMEDUSAE Coronatae Atolla wyvillei M-84 Nausithoe punctata Nausithoe spp. M-85 Paraphyllina intennedia M-86 Periphylla periphylla M-87 a Chrysaora hysoscella M-88 c Discomedusa lobata M-89 Pelagia noctiluca M-90 Rhizostomae Rhizostoma pulmo M-9l a X X X X X X X X Semaeostomae eferences General: Kramp, , 59 Alboran: Gibralter: X X a X X X X X Goy , 83, Rodriguez ' 83, Harbison pers. comm. Goy , 83 Catalan: Gili et al, '87; '88 Lyon: Razouls & Thiriot ' 68, Casanova' 70, Franqueville Ligurian: Goy , 72, Goy et al ' 89, Tregouboff ' 56, , 58 Tyrrhenian: Brinckmann-Voss ' 87 Adriatic: X Benovic & Bender ' 87 , 70 ~, ~i. Î; :, F 18 3. Siphonophores. Siphonophores are diverse and widely distributed predators. Most Mediterranean species are also found in warm parts of the Atlantic or other oceans. Because of the complex life cycle and morphology of siphonophores, and their fragility, many species are known only from parts of the whole organism. Distribution of siphonophores in the Alboran Sea and adjacent areas has been reported by Alvarino (1957), Casanova (1970), Gili et al. (1987, 1988), and Patriti (1969). General distribution in the Mediterranean is discussed by Bigelow and Sears (1937), and worldwide distribution of most descnbed species is summanzed by Alvanno (1971). Table 3 lists 56 species of siphonophores reported from the Western Mediterranean. They are arranged alphabetically within suborders, and 49 of them are described and illustrated in Section D. The most abundant species in the western Mediterranean are discussed here. The small calycophorans are the most common siphonophores in surface waters. Of these, Abylopsis tetragona, Chelophyes appendiculata, Diphyes dispar, Muggiaea atlantica, Eudoxoides spiralis and Lensia conoidea are listed as common in the western Mediterranean. In the Catalan Sea, M. atlantica occurred in densities up to hundreds m-3 in May and June, and M. kochi was found in maximum densities of more than 4 m-3 in the Gulf of Gabes near Tripoli (Patriti, 1969). Franqueville (1971) found peak abundances of A. tetragona and Chelophyes appendiculata in the spring near Toulon, and no evidence for vertical migration. C. appendiculata was also the most abundant siphonophore seen during submersible dives near Vilefranche by Laval et al. (1989). They found this species in the 100-250 m depth range, with evidence of a migration toward the surface at night. Densities of total diphyids (mostly C. appendiculata) ranged to over 200 per 1000 m3. They also noted that C. appendiculata could be distinguished in-situ from the similar Lensia conoidea because in the former both nectophores and stem hang vertically, while in the latter the nectophoreis horizontal and the stem hangs perpendicular to it. Other siphonophores reported by Laval et al. and earlier papers (Tregouboff 1956, 1957; Bernard, 1958) included Lensia subtilis, Muggiaea sp., Abylopsis tetragona, Hippopodius hippopus, Lilyopsis rosea, Agalma elegans, Nanomia bijuga, Halistemma rubrum and Forskalia edwardsi. Physonects are less commonly reported from plankton tows; they are harder to quantify because the colonies break apart in nets. Agalma elegans was quite abundant in May in the Catalan Sea (Gili et al. 1988) Submersible observations and collections elsewhere (Pugh and Harbison, 1986, 1987) indicate that large physonects and calycophorans are probably much more common in deep water than net tows suggest. 19 TABLE 3. SIPHONOPHORES. Geographic Occurrence Figu Species re Lu m Al Gibr ora alte n r Cata lan Lyon Ligu Tyrr rian heni an Adr iat ic Cystonect :':~ Rhizophysa filiforms 5-1 a Physonectae Agalm elegans Agalm okeni Agalm sp. 5-2 5-3 a X a X a Apolemia uvaria Athorybia rosacea Cordagalm cordiforms Forskalia edwards Forskalia spp. X i 5-4 5-5 5-6 a c X 5-7 a X a 5-8 Lychnagalma 5-9 c Marrus orthocanna c Physophora 5-10 5-11 5-12 5-13 Calycophorae Abyla haeckeli Abylopsis 5-14 5-15 a a X 5-16 a X a a X X Ceratocyma sagittata 5-17 5-18 5-19 a X X Chelophyes 5-20 a X X rum utricularia bijuga Nanomia cara Nanomia hydrostatiça eschscholtzi Abylopsis tetragona Arphicaryon acaule Bassia bassensis appendicula ta a X X X X X X X X X X X X X X X X X Halistemm Halistemra spp. rub X X X X a a X X X X X X X a X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 20 Species Figu re Chelophyes contorta Chuniphyes 5-21 5-22 Clausophyes ovata 5-23 5-24 5-25 5-26 5-27 5-28 5-29 5-30 5-31 5-32 5-33 multidentata Diphyes dispar Enneagonum hyalinum Eudoxoides spiralis Hippopodius hippopus Lensia camanella Lens ia conoidea Lensia fowleri Lensia meteori Lensia multicristata Lensia subtilis Lensia subtiloides Lilyopsis rosea atlantica Muggiaea Muggiaea kochi 5-34 5-35 5-36 5-37 Muggiaea sp. Rosacea Rosacea cyriformis plicata 5-38 5-39 5-40 5phaeronectes 5-41 irregularis m a Alb ora n Gibr alte r Cata Ian Lyon Adr iat ic X X a c c a X X X X X X X X X X X X X X X X X X X X X X c X c c c X c X X X X c c X X X X X X X X X X X X X X X X X X X X X X X X X X c X c X X X X c X X X X X a X a X X X X X X X X X kollikeri 5phaeronectes sp. quadri val vis Tyrr rian heni X 5phaeronectes 5ulculeolaria biloba 5ulculeolaria chuni 5ulculeolaria Ligu an c 5phaeronectes bougisi 5phaeronectes gracilis Lu 5-42 5-43 5-44 a X X a a X X X X X X X X X X X X X 21 Species Sulculeolaria turgida Vogtia glabra Vogtia pentacantha Vogtia spinosa Figu re Lu m Alb ora n Gibr alte r Cata Ian Lyon Ligu Tyrr rian heni an 5-45 5-46 5-47 a a X X X X a X X X X S-48 a X X Adr iat ic X X X References General: Alboran: Catalan: Lyon: Ligurian: Adriatic: Alvarino ' 71, Bigelow and Sears ' 37, Totton' 65 Harbison pers. comm. Gili et al ' 87, , 88, Rodriguez ' 83 Razouls & Thiriot '68, Casanova '70, Franqueville '70, Bernard '55,58 Biggs et al ' 86, . Laval et al ' 89, Tregouboff ' 56, , 58 Hure ' 55 22 4. Ctenophores Ctenophores are not easily collected in nets, and are rarely found in conventional zooplankton surveys. The only recent reports of ctenophoran fauna in the Alboran Sea found were unpublished dive logs (Harbison, pers. comm.) indicating the presence of P/eurobrachia sp., and unidentified cydippid ("redtentacle"), Bolinopsis vitrea, Leucothea mu/ticornis and Beroe spp. A diving survey made near Villefranche in 1986 also found Leucothea mu/ticornis, P/eurobrachia pileus, Callanira bia/ata, Cestum veneris and Beroe sp. in densities of ..1 per 1000m3 in the top 20 m (Biggs et al. 1987). Many ctenophore species were originally studied and described in the Meditteranean by Chun (1878, 1880, 1898), Fedele (1940) and others working in areas like Naples or Messina, where ctenophores were common at the surface and could be collected by dipping from a rowboat. Species found anywhere in the Mediterranean are likely to occur in the Alboran Sea. Most of the Mediterranean species also occur in the Atlantic, with the apparent exception of the genus Ocyropsis. Since this is known from the Canary Islands, it seems remarkable that it has never entered the Mediterranean, and it is possible that it has simply been overlooked. Table 4 lists 25 species of ctenophores from the Mediterranean, of which 20 are described and illustrated. Some ctenophores occur elsewhere in periodically dense populations. These include species of P/eurobrachia, Mnemiopsis, Leucothea and Beroe. Large populations are more likely near the surface and near shore, where they may be partly caused by hydrographic aggregation. Most of the species listed here are known from surface waters, but a very rich mesopelagic ctenophore fauna has been discovered in recent years through the use of submersibles. Bathocyroe fosteri and Thalassocalyce inconstans (Madin and Harbison, 1978a,b), originally described from the Atlantic, have been reported in the Mediterranean (Laval et al. 1989, Carré, pers. comm.). A great many other new species have been reported from submersible dives in the western Atlantic (Larson et aI., 1988) and are in the process of being described (Harbison and Botkin, in prep.; Madin, unpubl.). Virtually all ctenophores studied to date are brightly luminescent, producing light in the meridional canals, or in Eurhamphaea vexiligera, releasing luminous secretions when disturbed. They are likely to be important luminous sources in midwater, but may also be difficult to collect and identify. Ctenophores have been reported from submersible dives by several authors. Laval et al. (1989) reported that Bathocyroe sp. was one of the most abundant species seen, occuring mostly between 200 and 750 m. Other species reported were Pleurobrachia rhodopis, Cestum veneris, Beroe ovata and Thalassocalyce inconstans. Tregouboff (1956, 1957) saw P/eurobrachia, Cestum, Bolinopsis and another lobate in bathyscaphe dives near Villefranche, and Bernard (1958) reported small cydippids between 50 and 1000 m off the coast of Toulon. r 23 TABLE 4. CTENOPHORES. Species Geographic Occurrence Figu re Lu m Alo Gibr Cata ran alte lan r Lyon Ligu Tyrr Ad an at ic rian heni ri Cydippida Callianira bialata Euplokams stationis Haeckelia bimculata C-l C-3 a Haeckelia rubra Hormphora hormphora Hormphora plumosa Hormphora spatulata Hormphora spp. C-4 a X a X C-S a X C-6 a Lampea pancerina C-7 a P leurobrachia pileus P leurobrachia rhodopis "red-tentacle cydippid" C-8 a a X X X C-2 X X a X X X a X X X X X X X Lobata Bathocyroe fosteri Bolinopsis spp. C-9 Bolinopsis vitrea C-IO a Deiopea kaloktenota C-LL a X Eurhamphaea vexilligera C-l2 a X Leucothea multicornis C-13 a Thalassocalycida Thalassocalyce inconstans C-l4 X a X X X X X X X Cestida Cestum veneris C-1S a Velamen parallelur C-16 a X X X X 24 Species Figu re Lu m AJo Gibr Cata ran alte r lan Lyon Ligu Tyrr Ad an at ic rian heni Beroida forskalii C-17 a Beroe mitrata C-18 a Beroe ovata C-19 a Be roe te:ferences X X X X General: Alboran: Chun '80, Fedele '40 Harbison pers. coro. Ligurian: Tyrrhenian: Tregouboff '56,'58, Laval et al '89, Biggs et al '86 Chun '80, Fedele '40 Lyon: Adriatic: Razouls & Thiriot ' 68 Fedele ' 40 X X ri 25 5. Polychaetes and Nudibranchs Records of pelagic polychaetes and nudibranchs are rather scattered; only Hure (1955) devotes much attention to the species found in the Adriatic. Most species however, have a fairly wide distribution and probably can be expected in the Alboran Sea as much as anywhere. The tomopterids are known to be bioluminescent (Herring, 1987), and the alciopids secrete a greenish-yellow ink when disturbed, which may be luminescent. The nudibranch Phyllrhoe is also luminescent. These zooplankters rarely seem abundant enough that their vertical or seasonal distributions have been analyzed. Bernard (1955) saw Tomopteris at 660 and 1085 m; Tregouboff (1956, 1957) noted that genus and other pelagic polychaetes at 200, 650 and 990 m. In the Caribbean, large (25 cm) tomopterids have been collected at about 900 m (Madin, unpubl.). Table 5 lists 14 species; 6 are ilustrated. 26 TABLE s. POLYCHATES and NUIBRACHS. Geographic Occurrence Tyrr Fig Lu Albo Gibr Cata Lyon Ligu heni Species rian ran alte Ian In ure an r Adr iat ic Polychaetes X Alciopa contrainii X Asterope candida Calizonella lepidota X P-1 X Lopadorhynchus brevis Lopadorhynchus uncinatus p-2 X X Sagitella X kowalevskii Tornopteris cavalii Tornopteris elegans Tornopteris helgolandica a X X X X a p-3 X X Tornopteris planktonis Tornopteris sp. Vanadis crystallina Vanadis formosa Nudibranchs phyllirhoe- sp. a X P-4 P-5 p-6 References Lyon: Bernard' 55, Franqueville ' 71 Adriatic: Hure ' 55 Ligurian: Tregouboff '56, '58 X 27 6. Pelagic Tunicates There don't seem to be any recent reports of pelagic tunicates from the Alboran, but Jansa (1985) collected 13 species of larvaceans and 1 salp in the region west and south of Mallorca in the Catalan Sea. Most abundant were Oikopleura longicauda, O. dioica and Fritilaria borealis. The only salp collected was Thalia democratica. Most species known from the Mediterranean are widely distributed there (and in other oceans), and probably occur in the Alboran. Table 6 lists 54 species of Thaliaceans and Larvaceans, and 38 of these are described and ilustrated. The larvaceans are better represented in net collections because they are smaller and more numerous. Except in periodic swarms, salps are likely to be sparsely distributed. Doliolids can also form dense populations, but are more likely to be scattered in midwater. Pyrosomes are intensely luminescent, but luminescence of salps and doliolids is doubtfuL. A few species, Pyrosoma atlantica, 5a1pa fusiformis, lasis zonaria and possibly Thetys vagina are vertical migrators. Off Toulon, Franqueville (1971) found P. atlantica and S. fusiformis between 300 -900 m during the day and in the top 200 m at night. Other species reported (5. maxima, P. bicaudata, T. democratica, i. punctata) were generally at shallower depths. Maximum abundance of the salps was generally in the spring, but most pyrosomes were collected in autumn. Salps have been rather infrequently seen from submersibles in the Mediterranean. Bernard (1958) reported a few Thalia at 310m; Tregouboff (1956, 1957) saw these, as well as 5. maxima and P. bicaudata. Laval et al. (1989) found only a few S. fusiformis and 7 pyrosome colonies. On the other hand, small pyrosomes (to 10 cm), were quite common on dives made by Tregouboff (1956, 1957). Larvaceans and their houses are much more commonly reported from submersibles. Tregouboff and Bernard saw species of Fritilaria, Megalocercus and Stegosoma, some to depths of 300 m. Laval et al. (1989) observed very high densities of houses of Oikopleura albicans and other oikopleurids in the upper layers. They estimated that abundances ranged from 200 to 1 milion houses per 1000 m3, and that over 50% of them were abandoned. Similar densities were reported in surface waters by Scuba divers (Biggs et aI., 1987). Much larger houses, attributed to Megalocercus abyssorum or 5tegosoma magnum were seen from 300-450 m. These houses were up to 4 cm in diameter, and were seen in densities up to 59 per 1000 m3. Both the larvaceans themselves and the houses (in some species at least) are luminescent (Galt, 1989). 28 TABLE 6 PELAGIC TUNICATES. Species Figur e Geographic Occurrence Lu m Alb ora n Gib ral ter Cata lan Lyon Ligu 'lyrr rian heni an Adr iat ic Pyrosomas Pyrosoma atlanticum T-l a a pyrosomes X X X X X X X X X X X X Doliolids Dolioletta gegenbauri T-2 c Doliolum denticulatum T-3 b Doliolum mulleri T-4 b X X b X X Doliolum nationalis Salps Cyclosalpa affinis Cyclosalpa pinnata Cyclosalpa polae T-5 b T-6 b T-7 b X X X X Helicosalpa virgula T-8 Iasis zonaria T-9 Ihlea punctata Pegea bicaudata Pegea confoederata Pegea socia T-IO X X T-ll X X . Salpa fusi"formis Salpa maxima Thalia democratica Thalia orientalis Thetys vagina X X T-12 T-13 X X X T-14 X X X X T-15 X X X X X T-16 X X X X X T-17 X T-18 X - 29 Species Figur e Appendicularians Appendicularia sicula Appendicularia T-19 Folia gracilis T-20 Lu m A1b Gib n ter ora rai Cata ian Lyon aequatorialis Fritillaria borealis Fritillaria charybdae Fritillaria fagei Fritillaria formica Fritillaria fraudax Fritillaria gracilis Fritillaria haplostoma Fritillaria Fritillaria megachile X X Fritil1aria b Kowalevskia tenuis Kowalevskia oceanica Megalocercus iat ic X X X X X T-22 X X X X X X X X X T-23 T-24 X X X X X X X T-25 X X X X X X X X T-26 X X X X X X X T-27 X X T-28 X X X X X X X X X X X T-29 c X X X X X albicans T-30 a X X X X X cophocerca T-31 T-32 T-33 a X X X a X X X X X a X X X X X X X X X abyssorur Oikopleura Oikopleura Oikopleura Oikopleura Oikopleura X T-21 spp. Fritillaria tenella Fritillaria urticans Fritillaria- . venusta X Adr X messanensis Fritillaria pellucida Tyrr rian heni an tregouboffi Fritillaria Ligu dioica fusiforms a graciloides Oikopleura intermedia T-34 a X X X 30 Species oikopleura roediterranea Oikopleura parva Oikopleura rufescens Oikopleura sp. Figur e m Alb ora n Gib rai ter Cata ian Lyon Ligu an X T-36 a X T-37 a X T-38 a c a Tectillaria fertilis Tyrr rian heni a Pelagopleura haranti Stegosoma magnum Lu X X X X X Adr iat ic X X X X X X X X X ~i:erences General: Alboran: Catalan: Lyon: Ligurian: Adriatic: Fenaux ' 67 Rodriguez ' 83, Rodriguez et al ' 82, Harbison, pers. comm. Jansa ' 85, Trepat ' 83, Godeaux ' 85 Bernard '58, Franqueville '70,'71, Razouls & Thiriot '68, Casanova '70 Tregouboff ' 56,' 58, Laval et al ' 89, Bracconot ' 70,' 73, Fenaux ' 59 Hure ' 55, Godeaux ' 87 " ',' 31 7. Crustaceans As elsewhere, the crustaceans, particularly copepods, make up most of the numbers, biomass and diversity of the zooplankton. Although there is a much larger historical literature concerning copepods and other crustaceans in the Mediterranean, only recent studies concerned with the Alboran Sea and adjacent areas are considered here. The papers by Rodriguez (1983) and Rodriguez et al. (1982) are mainly concerned with copepods in the Alboran. Table 7 lists 90 species of hyperiid amphipods, euphausiids, mysids, copepods, ostracods and decapod shrimp; 45 of these are described and ilustrated. Cladocerans have no known luminescent genera and are not included. One hyperiid amphipod genus is reportedly luminescent, as are a few copepods. All the listed genera of euphausiids, ostracods and almost all the decapods are also bioluminescent. The most abundant copepods reported in the Alboran Sea in March, April and May from tows taken in the top 20 m were Paracalanus parvus, C/ausoca/anus spp., Centropages chierchiae, Acartia c/ausi, Temora sty/ifera, Oncaea spp. and Oithona spp. (Rodriguez, 1983). Furnestin (1968) cites P. parvus, C/ausocalanus arcuicornis and T. stylifera as the species constituting most of the copepod biomass in the Alboran in early summer. The only ostracod reported by Rodriguez (1983) was Conchoecia sp., which had maximal abundance in March and ApriL. Another genus, Cypridina, is distributed throughout the Mediterranean but in deeper water than sampled by Rodriguez. Euphausiids were collected with midwater trawls by Wiebe and D'Abramo (1972) in several parts of the Mediterranean. The dominant species occuring in the Alboran Sea were Euphausia krohni, Nematoscelis mega/ops, Sty/ocheiron abbreviatum and S. suhmii. Vertical distribution of larger crustaceans near Toulon is reported by Franqueville (1971). With the exception of Sty/ocheiron maximum, which was always found between 200 - 500 m, euphausiids collected by Franquevile were diel migrators, moving from 400 - 2400 m by day to near surface waters at night. Most abundant species were Meganyctiphanes norvegica, with maximum abundances in summer of 100 per 5000 m , Euphausia krohnii and Nematoscelis megalops. The hyperiid amphipods Phronima sedentaria and Scina crassicornis exhibited diel migration between 400 - 1400 m by day and 0 - 200 m at night. Maximum seasonal abundances of P. sedentaria and Phrosina semilunata were in spring and falL. Decapod shrimp in Franquevile's samples were dominated by Sergestes arcticus, with maximum abundance in summer, and Gennadas elegans, most common in winter and spring. Both migrate from daytime depths as great as 1400 m to the top 100 m at night. Submersible observations (Bernard, 1955, 1958; Tregouboff, 1956, 1957) have included reports of "large copepods", mainly calanoids 3 - 5 mm long and mainly below 900 m, Sapphirina sp. at 300 and 1900 m, euphausiids common between 600 and 2000 m, the peneid Gennadas elegans between 500 - 600 m, and sergestids below about 600 m. Laval et al. (1989) saw Phronima in barrels from their submersible. .. 32 TABLE 7. CRUSTACEAS. Geographic Occurrence Figur Lu Alb Gib Cat Species m ora rai aia e n ter n Lyon Ligu Tyrr rian heni an Adr iat ic Amhipods Amhithyrus bispinosus Amhithyrus simlis Brachyscelus crusculum X X CR-l X X X X X X Calamorhynchus X rigidus Euprimo macropus Eupronoe minuta c X X X X Glossocephalus milne-edwards i Hyperia schizogeneios Hyperia hydrocephala Hyperioides longipes Lycaeopsis X Paralycaea gracilis X X X X X X X themistoides X Paraphronima gracilis Phronim atlantica CR-2 X Phronima sedentaria CR-3 X Phronimella elongata CR-4 X X X Phronimopa"is X spinifera Phrosina semilunata Platyscelus ovoides CR-S X X CR-6 X X X Platyscelus serratulus Pseudolycea pachypoda X X X X X X CR-7 X X X X Rhabdosoma X brevicaudatum Scina borealis Scina crassicornis CR-8 Streetsia challengeri CR-9 a X a X X X X X X X X 33 Species Figur e Lu m Al ora n Gib ra1 ter Cat a1a n Lyon Ligu Tyrr rian heni an vibilia armta X X X Euphausiids a CR-IO CR-l1 atlantica Nematoscelis megalops Stylocheiron abbreviatum Stylocheiron longicorne Stylocheiron maximum Stylocheiron suhmii Thysanopodå aequalis CR-12 CR-14 X X a X a X a X a X a X X X a X X X X a X a X X X X Mysids Boreomysis semicaeca X Euchaetomeropsis merolepis Eucopia hanseni Lophogaster typicus X X c X X a a CR-13 X X a euphausiids norvegica Nematoscelis ic X X Vibilia jeangerardi Vibilia viatrix Meganyct iphanes iat X Tetrathyrus forcipatus Euphausia brevis Euphausia hemigibba .. Euphausia krohnii Adr X X 34 Species Figur e Lu m Al ora n Gib Cat ral ala ter n Lyon Ligu Tyrr rian heni an Adr iat ic Copepods Acartia clausi Acartia grani CR-1S X Aetidius armatus Calanus brevicornis Calanus helgolandicus Calanus minor Calocalanus sp. Centropages chierchiae Centropages kroyeri Centropages typicus Clausocalanus arcuicornis Clausocalanus sp. Coryceus sp. Ctenocalanus vanus Eucalanus elongatus Eucalanus hyalinus Eucalanus monachus Euterpina acutifrons X X X X CR-16 c X c X c X X X X X CR-17 X CR-18 X CR-19 X X CR-20 X X X X X X CR-21 b X X X CR-22 X X X X X X X X X X X X X X X Haloptilis.. acutifrons Lucicutia flavicornis CR-23 a X X CR-24 a X X oi thona sp. CR-2S b X X Oncaea sp. CR-26 a X X ;, - Onchocalanus spp. Paracalanus parvus CR-27 P leuromam borealis CR-28 a X gracilis CR-29 a X P leuromamma X P seudocalanus elongatus Rhincalanus nasutus Sapphirina sp. X X X X CR-30 X CR-31 X X " ìj, 35 Species Scolecithrix bradyi Figur e CR-32 Lu m c Alb ora n Gi. Cat ral ala ter n Lyon Ligu Tyrr rian heni an Adr iat ic X X Temora longicornis CR-33 X Temora stylifera CR-34 X X X X Ostracods Conchoecia sp. CR-35 a Cypridina castanea CR-36 a CR-37 a Decapods Acanthephyra pelagica X Fi.nchalia sp. Gennadus elegans X CR-38 b CR-39 c X sivado CR-40 c X arcticus CR-41 a X corniculum a X mollis robustus sargassi a X CR-42 a X CR-43 a X a X a X multidentata Pasiphaea Sergestes Sergestes Sergestes Sergestes Sergestes Sargestes ~spp . Sergestes vigilax References General: Alboran: Lyon: Ligurian: Adriatic: X X Lucifer typus Pasiphaea X CR-44 X X Stephensen ' 25, Rose ' 33, Crosnier & Forest ' 73, Wiebe & D' Abramo , 72 Rodriguez ' 83, Rodriguez et al ' 82, Furnestin ' 68 Bernard' 55, Franqueville ' 70,' 71, Casanova, Razouls & Thiriot ' 68 Tregouboff ' 56, , 58 Hure ' 55 36 Ilustrated systematic guide to zooplankton of the Alboran Sea and adjacent areas. The following guide is intended for use in the field, with live animals or images of them. An effort was made to keep the descriptions concise, specific and free of specialized terminology or abbreviations. Some terms specific to major groups are defined in the beginning of each taxonomic section. For some groups pods and larvaceans, specialists differentiate species on the basis of rather like cope obscure or morphometric characters. These have been avoided here wherever possible. The illustrations are compiled from a variety of sources, indicated for each section. Wherever possible, a picture of the whole animal was used, but in the case of some siphonophores, larvaceans and crustaceans, only illustrations of parts were available. Original captions (not always in English) have been left on the ilustrations in some instances. 37 Hydromedusae and Scyphomedusae What are commonly called jellyfish are medusae belonging to two Classes of the Cnidaria -- the Hydrozoa and the Scyphozoa. Hydromedusae possess a velum around the umbrella opening that scyphomedusae lack. Since the morphology and lie history is broadly similar, it is most practical to treat them as one group here. There are perhaps 1000 species of hydro- and scyphomedusae, with undoubtedly more to be discovered, especially in deep or polar waters (e.g. Larson et at. 1988; Larson and Harbison, 1990). Some meso- or bathypelagic species known from other regions may occur in the Mediterranean, but have not yet been reported, and are not included here. Many species are luminescent, some very conspicuously. Many of these medusae are part of a life history that alternates between a sessile, benthic, asexually reproducing polyp and a sexually reproducing and dispersing planktonic medusa. However, many oceanic medusae have lost the polyp stage and have evolved a variety of sexual and asexual reproductive mechanisms that do not require a benthic habitat. In many cases polyp and medusa stages were described separately, with different names, and there are stil many instances in which the two stages have not been recognized as belonging to the same species. There are two classifications for Hydromedusae, based either on the polyp (hydroid) or medusoid forms. In this description, the classification follows that of Kramp (1961) based on medusoid stages. HYDROMEDUSAE 1. Anthomedusae. This order includes relatively small forms ranging in size from less than 1 mm to several em. The umbrella is usually a tall bell shape, and gonads are almost always found on the sides of the central stomach. There are 4 radial canals connecting the stomach to a marginal ring canaL. Tentacles occur in varying numbers around the umbrella margin and sometimes around the mouth. Anthomedusae alternate with polyp forms, but some also bud medusae directly. 2. Leptomedusae. These medusae are generally flatter than a hemisphere. They usually have 4 radial canals, but sometimes 8 or more, or canals that are branched. Gonads are located on the radial canals, and there may be various sense organs on the margin. The stomach is sometimes flat, and sometimes mounted on a peduncle which can be quite long. There are tentacles around the margin but not the mouth. Leptomedusae also alternate with hydroids, but again there are instances of direct production of new medusae by budding or fission. 3. Limnomedusae. Both high and low umbrella shapes are found in this group. There are usually 4 radial canals, sometimes branched. Centripetal canals occur in some species. Gonads are either on the stomach or the radial canals. There is alternation of generations. Many limnomedusae live in brackish or even fresh water, but there are marine genera. 4. Trachymedusae. These medusae do not alternate generations, but develop young medusae directly from planula larvae. The umbrella is often high, with stiff mesoglea and well developed muscle fibers. Most have 8 unbranched radial canals and gonads located on them. Many trachymedusae live in deep water and are heavily pigmented. 38 5. Narcomedusae. Narcomedusae also have direct development of medusae from planulae, and larvae are often parasitic on other medusae. There are no radial canals, but the flat central stomach is very wide and, in some genera, extends into radial stomach pouches. The umbrella margin is divided into lobes by grooves. Tentacles are solid and stiff, and often extend aborally. Narcomedusae are common in epipelagic and mesopelagic environments; some are strong vertical migrators. SCYPHOMEDUSAE 6. Coronatae. This order of scyphomedusae includes mainly deepwater forms. The umbrella is divided into a high central part and a thinner marginal part by a coronal groove. The margin of the bell is divided into lappets; sense organs and solid tentacles arise from the cleft between lappets. The mouth has simple lips and the gastrovascular cavity is often deeply pigmented. 7. Semaeostomae. The familiar large jellyfish are mainly in this order of the Scyphozoa. The umbrella margin is divided into lappets, and bears sense organs and hollow tentacles. There is no coronal groove around the umbrella. The mouth opening is surrounded by four long oral arms, often frilled. Gonads are in folds of the subumbrella. The classification and nomenclature used here follows Kramp (1961). Descriptions, illustrations and distributions are mainly from Kramp (1959, 1961), Goy (1983), Russell (1953, 1970) and Tregouboff and Rose (1957). Terminology: abaxial - outer surface of tentacle or bulb, away from umbrella aboral - the side of the umbrella opposite the mouth bell or umbrella - the main gelatinous body of a medusa centripetal canals - radial canals that begin at the bell margin and run partway to the apex cirri - small tentacle-like structures between true tentacles on the margin cordyli - club-shaped marginal structures located between tentacles coronal groove - a groove separating the central part of the bell from the peripheral in coronate scyphomedusae exumbrellar - the upper or aboral surface of the umbrella interradial - aligned between the 4 primary radii lappets - separated sections of the umbrella margin manubrium - central part of the medusa containing stomach and mouth 39 margin - the edge of the umbrella marginal clubs - short clublike structures around the margin, between tentacles marginal vesicles - spherical sensory structures arranged around the margin nematocyst knobs '. clump of nematocysts at the end of the tentacles nematocyst rings . thickened rings of nematocysts around the shaft of the tentacles nematocyst tracks - rows of nematocysts, usually on the umbrella surface ocell - light sensitive structures around the margin or at the bases of the tentacles ,', oral arms. extended lips hanging down from the mouths of semaeostome scyphomedusae oral tentacles - tentacles arranged around the mouth in anthomedusae otoporpae - linear structures, possibly sensory, on the marginal lappets of some narcomedusae perradial - aligned with the 4 principal radial canals pyriform - pear-shaped radial canals - the gastrovascular canals in the umbrella, extending from the stomach to the margin rhopalia - complex sensory structures on the margin of scyphomedusae '" :... :¡t statocysts - gravity-sensing vesicles on the margin stomach pouches - radial extensions of the central gastrovascular cavity subumbrella - the under or oral side of the umbrella tentacle bulbs - the swellngs on the margin from which the tentacles arise tentacle rudiments - undeveloped tentacle bulbs tentaculae - small tentacles l, 40 Fig. M-1 SPECIES: Amphinema dinema FAMILY: Pandeidae ORDER: Anthomedusae SIZE: to 6 mm high, 4 mm wide DESCRIPTION: globular bell with long, conical apica projection, 2 long tentacles with long conical bulbs, flask-shaped stomach with 4 recurved lips, simple adradial gonads LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: N. Atlantic, Indian, Med. Fig. M-2 SPECIES: Amphinema rubra FAMILY: Pandeidae ORDER: Anthomedusae SIZE: 7 mm high, 4.5 mm wide DESCRIPTION: ovoid bell coming to apical point, thick jelly, 2 tentacles with large conical bulbs, 6 small tentaculae, stomach barrel shaped, dark reddish-brown. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: .Antarctic, in deep water. Fig. M-3 SPECIES: Amphinema rugosum F AMIL v: Pandeidae ORDER: Anthomedusae SIZE: 5 mm high, 3 mm wide DESCRIPTION: domed bell with conical apical projection, 2 long tentacle with long bulbs, 16-24 solid tentaculae, stomach flaskshaped, gonads with 3-4 folds. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: N. Atlantic, W. Pacific, Med. 41 Fig. M-4 SPECIES: FAMILV: ORDER: SIZE: Amphinema turrida Pandeidae Anthomedusae 4-7 mm high, slightly less wide DESCRIPTION: domed bell, conical projection, 2 long tentacles, 14 small tentaculae, pyriform stomach with large lips, gonads folded, extending along radial canals LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: tropical Atlantic, Pacific, Med. Fig. M-5 SPECIES: Bougainvilla ,amosa FAMILV: Bougainvilliidae ORDER: Anthomedusae SIZE: 2 - 3.5 mm high and wide DESCRIPTION: globular bell, thick jelly, 3-4 long tentacles from each bulb, stomach short, oral tentacles short, divided 1-2 times, gonads globular in female, elongate in male LUMINESCENCE: Herring (1987) lists Lizzia in this family as uncertain. DISTRIBUTION:-N. Atlantic, Med. Fig. M-6 SPECIES: Bythotia,a murrayi F AMIL v: Calycopsidae ORDER: Anthomedusae SIZE: to 20 mm high and wide DESCRIPTION: globular bell, thick walls, 4 bifurcate radial canals, 8 or more long tentacles with end knobs, small tentaculae, small stomach, gonads with transverse furrows. LUMINESCENCE: unknown DISTRIBUTION: E. Atlantic, Med. in deep wate r 42 Fig. M-7 SPECIES: Calycopsis simplex F AMIL v: Calycopsidae ORDER: Anthomedusae SIZE: 8 mm high and wide DESCRIPTION: globular bell, 4 centripetal canals, 8 tentacles, stomach short, gonads with few transverse folds. LUMINESCENCE: unknown DISTRIBUTION: Norway, in deep water Fig. M-8 SPECIES: Cladonema radiatum F AMIL v: Cladonematidae ORDER: Anthomedusae SIZE: 4 mm high, 3 mm wide DESCRIPTION: thin walled bell, 4-5 bifurcate or 8-10 si mple radial canals, 8-10 tentacles with 4-6 branches, nematocyst knobs, 4-5 oral tentacles, gonad with 4-5 sacs. LUMINESCENCE: unknown DISTRIBUTION:.N. Atlantic, Med., Black Sea, creeps and swi ms Fig. M-9 SPECIES: Cytaeis tetrastyla FAMIL v: Cytaeidae ORDER: Anthomedusae SIZE: 6 mm high, 5 mm wide DESCRIPTION: domed bell, 4 tentacles with large, black bulbs, to 32 oral tentacles with nematocyst knobs, large stomach with medusa buds on upper part. LUMINESCENCE: unknown DISTRIBUTION: tropical and subtropical Atlantic, Pacific, Indian, Med. 43 Fig. M-10 SPECIES: Dipurena halterata FAMILY: Corynidae ORDER: Anthomedusae SIZE: 8 mm high, 6 mm wide DESCRIPTION: bell-shaped, thick jelly, 4 tentacles with 3-6 nematocyst rings and terminal knob, stomach on very long manubrium, gonads halfway down manubrium LUMINESCENCE: unknown DISTRIBUTION: N. Atlantic, Med. Fig. M-11 SPECIES: Dipurena ophiogaster FAMILY: Corynidae ORDER: Anthomedusae SIZE: 5 mm high, slightly less wide DESCRIPTION: bell-shaped, 4 tentacles with small, irregular nematocyst clusters, stomach on long manubrium, gonad with 26 segments on manubrium. LUMINESCENCE: unknown DISTRIBUTION: N. Atlantic, Pacific, Med. \:~~ , t Fig. M-12 SPECIES: Ectopleura dumortieri FAMILY: Tubulariidae ORDER: Anthomedusae SIZE: 2-3 mm high and wide DESCRIPTION: spherical bell, thick jelly, 4 tentacles with large bulbs, nematocyst clusters along length, 8 nematocyst tracks on exumbrella, stomach short. LUMINESCENCE: Herring (1987) lists Euphysa in this family as definite. DISTRIBUTION: Atlantic, Indian, Pacific, Med. 44 Fig. M-13 Eucodonium brownei SPECIES: FAMILY: Tubulariidae ORDER: Anthomedusae SIZE: 1 mm high and wide DESCRIPTION: globular bell, thin walls, 4 thin tentacles with terminal nematocyst knobs, stomach on short peduncle, with medusa buds on upper part, simple mouth. LUMINESCENCE: Herring (1987) lists Euphysa in this family as definite. DISTRIBUTION: N. Atlantic, Med. Fig. M-14 Euphysa aurata SPECIES: FAMILY: Tubulariidae ORDER: Anthomedusae SIZE: 4 mm high, slightly less wide DESCRIPTION: tall bell, thick jelly, 1 tentacle with rings of nematocysts, stomach tubular, encircled by gonad. LUMINESCENCE: Herring (1987) lists as definite. . DISTRIBUTION: Atlantic, Pacific, Med. dJ Fig. M-15 SPECIES: Haliiara formosa FAMILY: Pandeidae ORDER: Anthomedusae SIZE: 3 mm high DESCRIPTION: pear-shaped bell, solid apical projection, 4 hollow main tentacles, 24-35 short, solid, tightly-coiled tentacles, stomach half as long as bell, mouth simple. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: tropical Atlantic, Pacific, Indian, Med. 45 Fig. M-16 SPECIES: Hybocodon prolifer FAMILY: Tubulariidae ORDER: Anthomedusae SIZE: 4 mm high, 3 mm wide DESCRIPTION: bell-shaped, margin oblique, 1 bulb with 1-3 tentacles, 5 exumbrellar nematocyst tracks, cylind.rical stomach gonad, medusa buds on tentacle bulb. and LUMINESCENCE: Herring (1987) lists Euphysa in this family as definite. DISTRIBUTION: temperate and subarctic Atlantic, Pacific Fig. M-17 SPECIES: Koellkerina fasciculata FAMILY: Bougainvillidae ORDER: Anthomedusae SIZE: 8 mm high, 9 mm wide DESCRIPTION: barrel-shaped, thick walls, 8 tentacle bulbs, each with 10-13 tentacles, stomach on short peduncle, with oral tentacles divided 7 times, 4 horseshoe shaped gonads. LUMINESCENCE: Herring (1987) lists Lizzia in this family as uncertain. DISTRIBUTloN:-Med., Atlantic, Black Sea. Fig. M-18 SPECIES: FAMILY: ORDER: SIZE: Leuckartiara nobils Pandeidae Anthomedusae to 27 mm high and 20 mm wide DESCRIPTION: bell-shaped with conical apical projection, about 40 tentacles of various sizes, dark red ocelli, large manubrium, folded lips, folded gonads cover stomach. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: N. Atlantic, Pacific, Med. 46 Fig. M-19 SPECIES: Leuckartiara octona FAMILY: Pandeidae ORDER: Anthomedusae SIZE: to 20 mm high DESCRIPTION: bell-shaped, with conical or spherical apical projection, 12-24 (usu.16) tentacles, 16+ tentacle rudiments, red ocell, furrowed gonads cover broad stomach. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. M-20 SPECIES: Lizzia blondina FAMILY: Bougainvilliidae ORDER: Anthomedusae SIZE: 1-2 mm high and wide DESCRIPTION: globular bell, thick apex, 8 tentacle bulbs, perradial with 1-3 tentacles, interradial with 1, stomach on short peduncle with' oral tentacles, medusa buds. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION:.NE Atlantic, Med. Fig. M-21 SPECIES: Lizzia fulgurans FAMILY: Bougainvilliidae ORDER: Anthomedusae SIZE: 1 mm high DESCRIPTION: soft, globular bell, 8, sometimes 16, stiff recurved tentacles, small stomach on pyramidal peduncle, 4 oral tentacles, medusa buds on stomach. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: NW Atlantic. 47 Fig. M-22 SPECIES: Merga tergestina FAMILY: Pandeidae ORDER: Anthomedusae SIZE: 7 mm high, 4 mm wide DESCRIPTION: bell with thin walls and high, pointed apical projection, 4-8 tentacles with large bulbs, also some rudimentary bulbs, stomach short, gonads smooth. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: E. tropical Atlantic, Med. !- Fig. M-23 SPECIES: Merga vio/acea FAMILY: Pandeidae ORDER: Anthomedusae SIZE: to 11 mm high, 7 mm wide DESCRIPTION: bell with domed apex, 8-12 long and 24-36 rudimentary tentacles, stomach half length of bell, cross-shaped in section, smooth adradial gonads. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: Atlantic, Pacific, Indian, Med. e. Fig. M-24 SPECIES: Neoturris pi/eata FAMILY: Pandeidae ORDER: Anthomedusae SIZE: to 40 mm high, 25 mm wide DESCRIPTION: bell with variable apical projection, 60-80 tentacles with elongated bulbs, radial canals with short branches, stomach broad, complex lips, gonads pitted. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: Atlantic, Med. 48 Fig. M-25 SPECIES: Niobia dendrotentaculata F AMIL V: Pandeidae ORDER: Anthomedusae SIZE: 4 mm wide DESCRIPTION: very flat bell, 2 of 4 radial canals bifurcate, so 6 reach margin, 12 tentacles, medusa buds develop from tentacle bulbs, stomach elongate, gonads interradiaL. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: W. Atlantic, Indian Fig. M-26 SPECIES: Oceania armata FAMILV: Clavidae ORDER: Anthomedusae SIZE: to 10 nmm high DESCRIPTION: bell with thin walls, flat top, 60-100 crowded tentacles, stomach flaskshaped, on short peduncle, lips with nematocyst knobs. LUMINESCENCE: unknown DISTRIBUTION:.Atlantic, Pacific, Med. Fig. M-27 SPECIES: Pandea conica FAMILV: Pandeidae ORDER: Anthomedusae SIZE: to 21 mm high, 10 mm wide DESCRIPTION: conical bell with apical projection, ridges on exumbrella, 16-24 tentacles with abaxial ocelli, stomach in upper bell, with folded lips, reticulate gonads around stomach. LUMINESCENCE: Herring (1987) lists 2 other pandeids as definitely luminescent DISTRIBUTION: Atlantic, Pacific, Med. 49 Fig. M-28 SPECIES: Paragotoea bathybia FAMILY: Tubulariidae ORDER: Anthomedusae SIZE: 2 mm high, 3 mm wide DESCRIPTION: bell with thin walls, nematocyst clusters on exumbrella, 1 solid tentacle with nematocyst knob, stomach short with simple mouth, gonads surround stomach. LUMINESCENCE: Herring (1987) lists Euphysa in this family as definite. DISTRIBUTION: boreal Atlantic in deep water Fig. M-29 Podocoryne carnea SPECIES: FAMILY: Hydractiniidae ORDER: Anthomedusae SIZE: 1 mm high and wide DESCRIPTION: bell with thin walls, 4-16 tentacles, stomach cylindrical with simple mouth arms, gonads interradiaL. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. .. Fig. M-30 SPECIES: Podocoryne hart/a ubi FAMILY: Hydractiniidae ORDER: Anthomedusae SIZE: 3.5 mm high and wide DESCRIPTION: domed bell, thick at top, 8 large tentacles, up to 50 smaller ones, mouth with 4 simple arms, gonads on stomach, extend partway along radial canals. LUMINESCENCE: unknown DISTRIBUTION: NE Atlantic, Med. 50 Fig. M-31 SPECIES: Podocoryne minima FAMILY: Hydractiniidae ORDER: Anthomedusae SIZE: to 1 mm high and wide DESCRIPTION: bell with slightly thicker apex, 4 tentacles, stomach on peduncle, 4 mouth arms, medusa buds on interradial sides of stomach. LUMINESCENCE: unknown DISTRIBUTION: North Sea, Med. Fig. M-32 SPECIES: Podocoryne minuta FAMILY: Hydractiniidae ORDER: Anthomedusae SIZE: 0.3 mm high DESCRIPTION: bell pear-shaped, with solid apex, 8 equal tentacles, stomach on short peduncle, mouth with 4 arms, medusa buds on sides of stomach. LUMINESCENCE: unknown DISTRIBUTION:.Atlantic, Med. Fig. M-33 SPECIES: Rathkea octopunctata FAMILY: Rathkeidae ORDER: Anthomedusae SIZE: 3-4 mm high DESCRIPTION: bell pear-shaped with solid apex, 8 groups of dark pigmented tentacles, with 3 in interradial and 3-5 in perradial groups, mouth with 4 lips, medusa buds on stomach. LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION: Atlantic, Pacific, Black Sea, Med. 51 Fig. M-34 SPECIES: Sarsia eximia FAMILY: Corynidae ORDER: Anthomedusae SIZE: 3-4 mm high DESCRIPTION: bell-shaped, 4 tentacles with large oval bulbs, ocell, nematocyst warts and terminal knob, stomach cylindrical, surrounded by gonad. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. Fig. M-35 SPECIES: Sarsia gemmifera FAMILY: Corynidae ORDER: Anthomedusae SIZE: to 5 mm high DESCRIPTION: thick walls, 4 tentacles with nematocyst warts and terminal knob, very long manubrium with medusa buds along it, gonads around manubrium above stomach. LUMINESCENCE: unknown DISTRIBUTION:""N. Atlantic, Med. Fig. M-36 SPECIES: Sarsia prolifera FAMILY: Corynidae ORDER: Anthomedusae SIZE: to 4 mm high and wide DESCRIPTION: bell-shaped, thin walls, 4 tentacles with nematocyst warts, medusa buds from tentacle bulbs, manubrium short, gonads surround it. LUMINESCENCE: unknown DISTRIBUTION: N. Atlantic, Black Sea 52 Fig. M-37 SPECIES: Sarsia tubulosa FAMILY: Corynidae ORDER: Anthomedusae SIZE: to 18 mm high DESCRIPTION: bell-shaped, fairly thick walls, 4 long tentacles with nematocyst warts, no terminal knob, manubrium very long, gonads surround it, no medusa buds. LUMINESCENCE: unknown DISTRIBUTION: N. Atlantic, Pacific Fig. M-38 SPECIES: Steenstrupia nutans FAMILY: Tubulariidae ORDER: Anthomedusae SIZE: 5-6 mm high, 3-4 mm wide DESCRIPTION: bell with conical apical . projection, 1 long tentacle with nematocyst rings, 3 undeveloped bulbs, stomach on short peduncle, surrounded by gonad. LUMINESCENCE: Herring (1987) lists Euphysa in this family as definite. DISTRIBUTION: .N. Atlantic, Black Sea, Med. ,~, . Fig. M-39 SPECIES: Tiaranna rotunda FAMILY: Tiarannidae ORDER: Anthomedusae SIZE: to 20 mm wide DESCRIPTION: hemispherical bell, thick jelly, 16-28 tentacles, 2-3 cordyli between each, broad cruciform stomach with large lips, gonads in folds, extend under belL. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Antarctic, Med. in deep water t 53 Fig. M-40 SPECIES: Turritopsis nutricula FAMILY: Clavidae ORDER: Anthomedusae SIZE: 4-5 mm high and wide DESCRIPTION: bell-shaped, thin walls, 80-90 tentacles, large cruciform stomach, 4 lips with nematocyst knobs. LUMINESCENCE: unknown DISTRIBUTION: N. Atlantic, Pacific, Indian, Med. Fig. M-41 SPECIES: Zanclea costata FAMILY: Zancleidae ORDER: Anthomedusae SIZE: to 3 mm high and wide DESCRIPTION: bell-shaped, thick jelly, 2 or 4 tentacles with stalked nematocyst capsules along length, patches or tracks of nematocysts on exumbrella, stomach cylindricaL. LUMINESCENCE: unknown DISTRIBUTION:. Àtlantic, Pacific, Indian, Med. Fig. M-42 SPECIES: Aequorea aequorea F AMIL v: Aequoreidae ORDER: Leptomedusae SIZE: up to 175 mm wide DESCRIPTION: disk shape, thicker in center, usually 60-80 radial canals, tentacles usually fewer than canals, with elongated ' bulbs, stomach half width of umbrella. LUMINESCENCE: Herring (1987) lists this genus as definite. Source of aequorin. DISTRIBUTION: Atlantic, Med. 54 Fig. M-43 SPECIES: Eirene viridula At FAMILY: Eirenidae ORDER: Leptomedusae SIZE: 20-30 mm wide DESCRIPTION: bell hemisphencal, thick at center, 60+ tentacles of various sizes, 40+ marginal vesicles, stomach on gelatinous peduncle, gonads along radial canals. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. p r/' Eirene Diridula, with stomach amI bell margin (after KÜNNE). Fig. M-44 SPECIES: Eucheilota paradoxica FAMILY: Lovenellidae ORDER: Leptomedusae SIZE: 4 mm wide DESCRIPTION: globular bell, 4 tentacles with lateral cirri, 4 rudimentary bulbs with cirri, stomach small, gonads in middle of radial canals, medusa buds from gonads. LUMINESCENCE: Herring (1987) lists Lovenella in this family as definite. DISTRIBUTION:~Atlantic, Pacific Fig. M-45 SPECIES: Eutima gegenbauri FAMILY: Eutimidae ORDER: Leptomedusae SIZE: 20 mm wide DESCRIPTION: bell hemispherical, thick jelly at apex, 8-16 tentacles and 60-80 marginal warts, both with 1-2 cirn, stomach on long gelatinous peduncle, gonads on radial canals. LUMINESCENCE: Herring (1987) lists Tima in this family as definite. DISTRIBUTION: N. Atlantic, Med. 55 Fig. M-46 SPECIES: Eutima gracils FAMILY: Eutimidae ORDER: Leptomedusae SIZE: to 13 mm wide DESCRIPTION: bell flatter than hemisphere, jelly thick, 2-4 long tentacles, 40-80 marginal warts, both with cirri, stomach on long narrow peduncle, gonads along peduncle. LUMINESCENCE: Herring (1987) lists Tima in this family as definite. DISTRIBUTION: N. Atlantic, Med. Fig. M-47 SPECIES: o ~, Helgicirrha schulzei FAMILY: Eirenidae ORDER: Leptomedusae SIZE: 30-40 mm wide DESCRIPTION: bell flatter than hemisphere, jelly thin, 30-40 large tentacles, 100+ small tentacles or bulbs with lateral cirri, stomach small, gonads linear along radial canals. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. - .'.' -- ~.... . .~. ~"."....,_. Helgicirrha schului, with stomach and bell margin (after KÜNNE). Fig. M-48 SPECIES: Krampella dubia FAMILY: Laodiceidae (?) ORDER: Leptomedusae SIZE: 3 mm wide DESCRIPTION: bell hemispherical, 8 tentacles with swollen bases, 3-4 cirri between tentacles, gonads along length of broad radial canals, systematic position uncertain. LUMINESCENCE: Herring (1987) lists two genera in this family as uncertain. DISTRIBUTION: Atlantic 56 Fig. M-49 SPECIES: Laodicea neptuna FAMILY: Laodiceidae ORDER: Leptomedusae SIZE: 2.5 mm wide DESCRIPTION: bell nearly hemispherical, 8 short tentacles, 8 rudimentary bulbs, numerous cirri, stomach large, lips with 4 nematocyst clusters, gonads on upper parts of radial canals. ' LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION: Atlantic Fig. M-50 SPECIES: Laodicea ocellata FAMILY: Laodiceidae ORDER: Leptomedusae SIZE: 3.5 mm wide DESCRIPTION: bell globular, thin jelly, 7-14 tentacles, 10-18 rudimentary bulbs, large black ocell on bulbs, lips short, thick clubshaped gonads along radial canals. LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION:. Med. Fig. M-51 SPECIES: Laodicea undulata FAMILY: Laodiceidae ORDER: Leptomedusae SIZE: to 37 mm wide DESCRIPTION: bell flatter than hemisphere, 400-600 tentacles, spiral cirri and cordyli between tentacles, stomach short, long sinuous gonads along radial canals, ' reaching stomach. LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION: Atlantic, Med. 57 Fig. M-52 SPECIES: Lovenella cirrata FAMIL V: Lovenelidae ORDER: Leptomedusae SIZE: to 16 mm wide DESCRIPTION: bell hemispherical, 8-16 tentacles with 3-4 pairs spiral cirri and 3 rudimentary bulbs, stomach urn-shaped, gonads spindle-shaped, on distal radial canals. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. Fig. M-53 SPECIES: Mitrocoma annae FAMILV: Mitrocomidae ORDER: Leptomedusae SIZE: 30-40 mm wide DESCRIPTION: bell flatter than hemisphere, 60-100 tentacles with 3-8 cirri between them, 60-100 marginal vesicles, stomach small, gonads sinuous along distal 'radial canals. LUMINESCENCE: Herring (1987) lists Halistaura in, this family as definite. DISTRIBUTION:' Med. Fig. M-54 SPECIES: Mitrocomella brownei fAMILV: Mitrocomidae ORDER: Leptomedusae SIZE: 4-7 mm wide DESCRIPTION: bell flatter than hemisphere, 16-24 tentacles with 6-8 cirri between them, 8 marginal vesicles, stomach small, gonads oval, near distal ends of radial canals. LUMINESCENCE: Herring (1987) lists Halistaura in this family as definite. DISTRIBUTION: Atlantic, Mad. 58 Fig. M-55 SPECIES: Obelia spp. FAMILY: Campanulariidae ORDER: Leptomedusae SIZE: to 6 mm wide . DESCRIPTION: bell flat, jelly thin, numerous stiff, solid tentacles, 8 marginal vesicles, stomach short with square base, gonads spherical, on middles of radial canals. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide except polar Fig. M-56 SPECIES: Octophialucium funerarium FAMILY: Phialuciidae ORDER: Leptomedusae SIZE: 30-40 mm wide DESCRIPTION: bell lens-shaped, jelly thick, 8 radial canals, 64-128 tentacles, 2 marginal vesicles between tentacles, stomach small, gonads on distal part of radial canals. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: .N. Atlantic, Med. in deep water Fig. M-57 SPECIES: Phialidium hemisphaericum FAMILY: Campanulariidae ORDER: Leptomedusae SIZE: to 20 mm wide DESCRIPTION: bell hemispherical, jelly thin, 16-58 tentacles with 2 marginal vesicles between them, stomach small with simple lips, gonads oval or linear, along distal radial canals. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Indian, Med. ,~ '~~~,i,~,~Ä , ø¡ '--, l 1 -'l~~~ri . 59 Fig, M-58 SPECIES: Phialidium mccradyi FAMILY: Campanulariidae ORDER: Leptomedusae SIZE: 15 mm wide DESCRIPTION: bell lens-shaped. 16-24 tentacles. 1-2 marginal vesicles between them. stomach short with 4 lips. small gonads on radial canals., with hydroid buds. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: N. Atlantic Fig. M-59 SPECIES: rima lucullana FAMILY: Eutimidae ORDER: Leptomedusae SIZE: to 74 mm wide DESCRIPTION: bell flatter than hemisphere, jelly thin. radial canals extend onto peduncle. 60-70 short tentacles with 7 marginal warts between them. gonads along radial canals. LUMINESCENCE: Herring (1987) lists this genus as definite. \\~) ~\~) l';'..J DISTRIBUTION:' Med. Fig. M-60 SPECIES: Gonionemus vertens FAMILY: Olindiadidae ORDER: Limnomedusae SIZE: 15-20 mm wide DESCRIPTION: bell flatter than hemisphere. 60-80 long, stiff tentacles with adhesive pads on bent ends. stomach with 4 ruffled lips. folded gonads along most of radial canals. LUMINESCENCE: unknown DISTRIBUTION: world-wide temperate 60 Fig. M-61 SPECIES: Odessia maeotica FAMILV: Moerisiidae ORDER: Limnomedusae SIZE: to 18 mm wide DESCRIPTION: bell almost hemispherica, jelly thick, 16-32 tentacles, lobes of stomach extend along radial canals, gonads on radial canals and stomach walls. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Black Sea, Med. in ' brackish water Fig. M-62 SPECIES: Ollndms phosphonca FAMILV: Olindiadidae ORDER: Limnomedusae SIZE: 40-60 mm wide DESCRIPTION: bell hemispherical, 40-80 centripetal canals, 50-60 primary tentacles project aborally, 100-120 secondary tentacles, 100-170 marginal clubs. LUMINESCENCE: unknown DISTRIBUTION:-Atlantic and Med. Fig. M-63 SPECIES: Proboscidactyla ornata F AMIL v: Proboscidactylidae ORDER: Limnomedusae SIZE: 5 mm wide DESCRIPTION: jelly thick, 4 radial canals branch to 16-20, 16-20 tentacles. nematocyst tracks on umbrella, stomach with 4 radial lobes, medusa buds on stomach or canals. LUMINESCENCE: unknown DISTRIBUTION: world-wide in coastal waters 61 Fig. M-64 SPECIES: Scolionema suvaensis FAMILY: Olindiidae ORDER: Limnomedusae SIZE: 6 mm high, 9 mm wide DESCRIPTION: jelly thick, 40-70 tentacles of various lengths, with nematocyst rings and bent tips, cruciform stomach with small lips, gonads along distal radial canals. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. M-65 SPECIES: Aglantha digitale FAMILY: Rhopalonematidae Trachymedusae ORDER: SIZE: 10-40 mm high, 5-20 mm wide DESCRIPTION: thimble-shaped bell, clear, pink or red, 8 radial canals, 80+ tentacles, stomach on long peduncle, sausage-like gonads hang inside belL. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION:"1\tlantic, Pacific, surface to deep water Fig. M-66 SPECIES: Aglaura hemistoma FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: 4-6 mm high, 3-4 mm wide DESCRIPTION: bell with flat top, jelly thin, 8 radial canals, 48-85 tentacles, peduncle shorter than bell, stomach with 4 simple lips, sausage-like gonads attached above stomach. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: world-wide in surface layers 62 Fig. M-67 SPECIES: Arctapodema ampla FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: to 15 mm wide DESCRIPTION: bell flatter than hemisphere, thin walls, 8 radial canals, 100 tentacles, stomach with radial lobes, 8 gonads adjacent to stomach. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: Atlantic, Antarctic, Med. in deep water Fig. M-68 SPECIES: Geryonia proboscidalis FAMILY: Geryonidae ORDER: Trachymedusae SIZE: 35-80 mm wide DESCRIPTION: bell hemispherical, jelly thick, 6 radial canals with up to 7 centripetal between, 6 long and 6 small tentacles, stomach on long peduncle, gonads heartshaped on canals. LUMINESCENCE: Herring (1987) lists this genus as uncertain. -:~-,-f\.n '- DISTRIBUTION:" world-wide, tropical and subtropical Fig. M-69 SPECIES: Haliscera bigelowi FAMILY: Halicreatidae ORDER: Trachymedusae SIZE: 10 mm high, 17 mm wide , DESCRIPTION: high bell with thick apex, 8 broad radial canals, 96 tentacles, 24 statocysts, broad circular stomach, long oval gonads on canals. LUMINESCENCE: unknown DISTRIBUTION: N. Atlantic, Pacific in deep water 63 Fig. M-70 SPECIES: Ha/iscera conics FAMILY: Halicreatidae ORDER: Trachymedusae SIZE: to 18 mm wide DESCRIPTION: low bell with blunt conical apex, stiff jelly, 8 broad radial canals, 6472 tentacles, 16 statocysts, broad circular stomach, oval gonads in middle of canals. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Pacific, Med. in deep water Fig. M-71 SPECIES: Liriope tetraphylla FAMILY: Geryonidae ORDER: Trachymedusae SIZE: 10-30 mm wide DESCRIPTION: hemispherical bell, thick jelly, 4 radial canals, 4 long and 4 short tentacles, small stomach on long peduncle, gonads of variable leaf-like shape, on radial canals. LUMINESCENCE: Herring (1987) lists this genus as unpertain DISTRIBUTION:~ world-wide in warm water Fig. M-72 SPECIES: Persa inc%rata FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: 3 mm high, 2 mm wide DESCRIPTION: high bell with thin walls, 8 radial canals, to 48 long tentacles with nematocyst knobs, tubular stomach on short peduncle, 2 oval pendent gonads on radial canals. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: Atlantic, Indian, Med. 64 Fig. M-73 SPECIES: Ransonia krampi FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: 15 mm high, 8 mm wide DESCRIPTION: high conical bell, thin walls, solid apical projection, 8 radial canals, 88 tentacles, small stomach on long peduncle, gonads along radial canals on peduncle LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: Atlantic, Med. in deep water Fig, 284, Ransania krampi, medusa and peduncle with gonads (alter RA"SO"i. Fig. M-74 SPECIES: Rhopalonema funerarium FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: to 17 mm wide, 14 mm high DESCRIPTION: bell domed, 8 radial canals, 8 main tentacles, 24 smaller cirri with terminal knobs, stomach narrow, linear gonads along distal radial canals. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION:.Atlantic, Indian, Pacific, Med.? in deep watèr Fig. M-75 SPECIES: Rhopalonema velatum FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: 8-10 mm wide DESCRIPTION: bell flatter than hemisphere, with apical knob, 8 radial canals, 8 clubshaped tentacles and 8-16 cirri, stomach long and narrow, gonads on radial canals. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: N. Atlantic, Med., surface to deep 65 Fig. M-76 Sminthea eurygaster SPECIES: FAMILY: Rhopalonematidae ORDER: Trachymedusae SIZE: to 6 mm wide, 3 mm high DESCRIPTION: bell with small apical knob, 8 radial canals, 8 tentacles and statocysts, short stomach with 4 short lips, globular gonads on distal radial canals. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: Atlantic, Indian, Med. in deep water Fig. M-77 Cunina globosa SPECIES: FAMILY: Cuninidae ORDER: Narcomedusae SIZE: to 18 mm wide DESCRIPTION: globular bell, thick jelly, no radial canals, 16 tentacles, stomach on broad gelatinous peduncle, 10-14 stomach pouches with square outline. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: tropical Atlantic, Pacific, Med. .~ Fig. M-78 SPECIES: Pegantha rubiginosa FAMILY: Solmarisidae ORDER: Narcomedusae SIZE: to 16 mm wide DESCRIPTION: domed bell, jelly thick, no radial canals, 12-16 rectangular margi nal lappets and tentacles, 2 long & 2 short otoporpae on each lappet, stomach without pouches. LUMINESCENCE: unknown DISTRIBUTION: tropical Atlantic, Med. 66 Fig. M-79 SPECIES: Solmaris f1avescens FAMILV: Solmarisidae ORDER: Narcomedusae SIZE: 15-23 mm wide DESCRIPTION: flat, lens-shaped bell, thick jelly, 12-17 tentacles, no radial canals, marginal lappets thin, with 2 statocysts, stomach without pouches. LUMINESCENCE: unknown DISTRIBUTION: Med. and adjacent Atlantic Fig. M-80 SPECIES: Solmaris leucostyla FAMILV: Solmarisidae ORDER: Narcomedusae SIZE: 3 mm wide DESCRIPTION: flat to hemispherical bell, no radial canals, 12-26 tentacles, 12-26 marginal lappets with 1 statocyst, stomach without pouches, annular gonad. LUMINESCENCE: unknown DISTRIBUTION: .Med. Fig. M-81 SPECIES: Solmaris solmaris F AMIL v: Solmarisidae ORDER: Narcomedusae SIZE: to 35 mm wide DESCRIPTION: flat, lens-shaped bell, no radial canals, 18-20 tentacles, marginal lappets with 6-8 statocysts, stomach without pouches, annular gonad. LUMINESCENCE: unknown DISTRIBUTION: Mad. 67 Fig. M-82 SPECIES: Solmissus albescens FAMILY: Cuninidae ORDER: Narcomedusae SIZE: 25-30 mm wide DESCRIPTION: lens-shaped bell, with warts on exumbrella, no radial canals, 14-16 stomach pouches and tentacles, marginal lappets rectangular with 5-8 statocysts. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Med. common in deep water Fig. M-83 SPECIES: Solmundella bitentaculata FAMILY: Aeginidae ORDER: Narcomedusae SIZE: to 12 mm wide DESCRIPTION: high bell with thick apex, no radial canals, 2 tentacles attached near apex and held aborally, 8-16 statocysts, 8 stomach pouches. LUMINESCENCE: Herring (1987) lists two genera in this family as definite. DISTRIBUTION: world-wide tropical-temperate, surface to dè"ep water Fig. M-84 SPECIES: Atalla wyilei FAMILY: Atolldae ORDER: Coronatae SIZE: to 150 mm wide DESCRIPTION: disc-shaped bell, deep coronal groove between center and margin, 22 tentacles and sense organs, stomach pigmented deep red, remainder brownish red. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide in deep water 68 Fig. M-85 SPECIES: Nausithoe punctata FAMILY: Nausithoidae ORDER: Coronatae SIZE: 9-15 mm wide DESCRIPTION: disk-shaped with thick center, 8 tentacles and 16 marginal lappets, 16 stomach pouches, large round yellow gonads. 'rt t~~~") (\ ';''!~ ,~~\ ~' lc: ~TF;:', ~ ~ .' ~'C (E~~'~','G '~~;¡ , ,:"'&)i". ..~.. -_/ LUMINESCENCE: unknown C" i~ DISTRIBUTION: world-wide Fig. M-86 SPECIES: Paraphyllna intermedia FAMILY: Paraphyllnidae ORDER: Coronatae SIZE: 15 mm wide, 8 mm high DESCRIPTION: domed bell, deep coronal groove, 12 tentacles and 16 marginal lappets, stomach reddish brown, 4 pairs of ovoid gonads. LUMINESCENCE: unknown DISTRIBUTION: Pacific, Indian, Med. in deep water SPECIES: Periphylla periphylla FAMILY: Periphylldae ORDER: Coronatae SIZE: to 200 mm high DESCRIPTION: high domed or conical bell, 12 stiff tentacles often held aborally, 16 marginal lappets, stomach and subumbrella dark red or purple, 8 U-shaped gonads. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide in deep water 69 SPECIES: Chrysaora hysoscella FAMILY: Pelagiidae ORDER: Semaeostomeae SIZE: to 200 mm wide DESCRIPTION: saucer-shaped bell, smooth surface, 16 broad radial brown bands on exumbrella, 24 tentacles in 8 groups of 3, 32 marginal lappets, long frilled oral arms. LUMINESCENCE: Herring (1987) lists Pelagia in this family as definite. DISTRIBUTION: Atlantic, Med. Fig. M-89 SPECIES: Discomedusa lobata FAMILY: Ulmaridae ORDER: Semaeostomeae SIZE: 150 mm wide DESCRIPTION: disk-shaped bell, 24 tentacles, lappets, 8 rhopalia 32 marginal LUMINESCENCE: Herring (1987) lists Poralia in this family as definite. DISTRIBUTION: Atlantic, Med. Fig. M-90 SPECIES: Pelagia noctiluca FAMILY: Pelagiidae ORDER: Semaeostomeae SIZE: to 65 mm wide DESCRIPTION: bell flatter than hemisphere, yellow, brown or pink, nematocyst warts on outer surface, 8 tentacles and sense organs, 16 marginal lappets, 4 long oral arms. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide 70 Fig. M-91 SPECIES: Rhizostoma pulmo F AMIL v: Rhizostomatidae ORDER: Rhizostomeae SIZE: to 600 mm wide DESCRIPTION: domed bell, very thick jelly, nematocyst warts on sunace, no tentacles, 64-72 marginal lappets, oral arms divided into multiple mouths. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. 71 Siphonophores The siphonophores compose an order of the class Hydrozoa, and are thus most closely related to hydromedusae. Their complex life cycle and colonial morphology are very different from the relatively simple hydromedusae and for most practical purposes it is easier to consider the siphonophores as a separate group. The colonial, or polygastnc, phase of the life cycle is the largest and most familar, and the part which is descnbed here. Siphonophores consist of a collection of medusoid and polypoid zooids (see terminology) which are budded asexually from a founding larval polyp. The colony may include a gas float, nectophores or swimming bells, and a series of stem groups that include the feeding polyps and tentacles. In some siphonophores the stem groups break off as secondary dispersal and sexually reproductive stages called eudoxids. The colony can be thought of as an overgrown, polymorphic juvenile stage which eventually bears the sexually ids called gonophores which produce reproductive adults. These are medusoid zoo gametes. In different groups, the gonophores may remain attached to the colony, detach as part of a eudoxid or detach as individual medusae. Siphonophores range in size from a few mm to over 30 m in length, and occur throughout the water column. All are predators on other small zooplankton, and many genera are known to be luminescent. The colonies are fragile, and usually break up into their various units when collected in plankton nets. For this reason, much of the taxonomy is based on the morphology of the pieces, pnncipally nectophores, and some species are known only from a few such pieces. As a result, the appearance of the intact colonial stage is not always known. Where possible, ilustrations of intact siphonophres are provided here, but in some cases only pictures of pieces are available. In recent years many new deepwater species collected with submersibles have been descnbed (Pugh and Harbison, 1987; Pugh and Youngbluth, 1988). Although not yet reported from the Mediterranean (or included here), these, or other new species, may well be encountered at depth. The Order Siphonophora is divided into 3 suborders and 15 families. 1. Cystonectae. This suborder includes siphonophores which possess a float but no swimming bells. The Portuguese man-o-war is the most familar example. The float is so large that the animal floats on the surface. It is not generally taken in plankton collections and is not included here. 2. Physonectae. These siphonophores have more complex colonies, comprising a small apical float, numerous swimming bells that form a nectosome, and a stem containing several groups of gastrozooids, tentacles, bracts etc. The stem typically contracts when the animal is swimming, and then relaxes so that the stem and tentacles extend to maximum length for fishing. Many physonects are strong swimmers and vertical migrators. 3. Calycophorae. In this group, the float is absent, and the nectophores are reduced to a small number, most frequently two. The stem can be retracted completely into a cavity in the nectophores. A sequence of stem groups are budded, and break free as eudoxids. Calycophorans are the most diverse, widely distributed and abundant siphonophores. 72 The classification used here is based on Totton (1965). Descriptions and ilustrations are compiled from Bigelow and Sears (1937), Biggs (1977), Carré (1979), Pugh and Harbison (1986), Totton (1965) and Tregouboff and Rose (1957). Distributional data came mainly from Alvariño (1971), Bigelow and Sears (1937, Pugh (1974) and Totton (1965). Terminology: basal tooth - a tooth or projection from the ostial surface of a nectophore bract - a flattened, leaf-like zooid with little internal structure, for protection of stem groups and buoyancy cnidoband - folded or coiled band of nematocysts that is part of a tentilum cormidia - stem groups on the siphosome, usually consisting of gastrozooids, palpons, bracts and gonophores eudoxid - a stem group released from calycophorans as a free-swimming dispersal stage gastrozooid - polypoid feeding zooid with a single tentacle that catches and ingests prey hydroecium - cavity in the nectophore of calycophorans that houses the retracted stem nectophore - an asexual medusoid zooid that provides locomotion by jet propulsion nectosac - th.e cavity in the nectophore from which water is expelled for propulsion ostium - the opening of the nectophore palpon - a reduced gastrozooid with a simple tentacle and no ingestive capabilty pneumatophore - the gas float of a cystonect or physonect siphosome - the part of the stem with the gastrozooids, tentacles, bracts etc (cormidia) somatocyst - a part of the gastric cavity which occurs in the nectophores of calycophorans tentila - a side branch of the tentacle which may be simple or consist of a cnidoband and other terminal appendages terminal filaments - filaments attached to the sac containing the cnidoband on a tentillum 73 tricornuate - tentilum having three appendages off the cnidoband unicornuate - tentillum having one appendage off the cnidoband '. f 74 Fig. 5-1 SPECIES: Rhlzophysa fiIformis FAMILY: Rhizophysidae SUBORDER: Cystonectae SIZE: 2-50 cm long DESCRIPTION: apica pneumatophore 12 mm high, no nectophores, gastrozooids 25 mm apart on highly contractile stem, 1 tentacle per gastrozooid, with 3 types of tentila. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. Fig. 5-2 SPECIES: Agalma elegans FAMILY: Agalmidae ¿-.~v ,?' ../-' "...¿~JI, .c SUBORDER: Physonectae SIZE: to 1 m long ,,( ..' y /,"?J DESCRIPTION: nectophores arranged in 2 /~~ /(~ ~ 't.7 .(', " ') rows, slightly rounded with triangular nectosac, 2 rows of triangular bracts with 3 f/"Lr ridges, brick-red tricornuate tentila. LUMINESCENCE: Herring (1987) lists this genus as definite, DISTRIBUTION: ~Atlantic, Med. ilr'l i¡l ' 1J Fig. 5-3 SPECIES: Agalma okeni FAMILY: Agalmidae SUBORDER: Physonectae SIZE: to 30 cm long DESCRIPTION: prismatic nectophores form dodecagonal nectosome, Y -shaped nectosac, thick, faceted bracts, brick-red bicornuate or tricornuate tentila. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide in warm regions 75 Fig. 5-4 SPECIES: Apolemia uvaria FAMILY: Apolemiidae SUBORDER: Physonectae SIZE: to 30 m long DESCRIPTION: about 12 nectophores with tentacles. white cormidia including 3-4 gastrozooids. 2-40 bracts and 20-40 red pal pons are widely spaced on siphosome. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic. Pacific. Indian. Med. Fig. 5-5 SPECIES: Athorybia rosacea FAMILY: Athorybiidae SUBORDER: Physonectae SIZE: to 3 em wide DESCRIPTION: large central pink-red pneumatophore. no nectophores, no siphosome,. elongate bracts like overlapping petals, bi- or tncornuate tentHla. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. Fig. 5-6 SPECIES: Cordagalma cordiformis FAMILY: Agalmidae SUBORDER: Physonectae SIZE: DESCRIPTION: very small, heart-shaped nectophores "i LUMINESCENCE: Herring (1987) lists three other genera in this family as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. Fic, 25. (;d4"' cotl!.. Tonon A,..in aD B, -w YÎ of. i- fro th Gret Ba Re, x 47 76 SPECIES: Forskalia edwardsl FAMILY: Forskalidae SUBORDER: Physonectae SIZE: to 3 m long when extended DESCRIPTION: cylindrical or conica nectosome of numerous small nectophores with yellow spots on the orifices, long gastrozooids, palpons release red liquid. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, common in Med. ~" ,,"-' ~', ' ~.. ,,~. ~'" ,. . ,- : ~~,(""" '~~~~ Fig. 8-7 ~f~~-. "''"'Ø.. \~~'~~1*fl'': ,.. ,,"' Fl!. :"~ , "" '\ -"' "' .-:: ~"\, "\ 'f'#'V If '"",, '~, ."' "\ ..~ , \,~ A.\;~ :,,: ,. , ,:\~.)~-i~/' , '. , ~\\.~~~-~. \'\ '.'ì t--." '" . ,\~~,) . ..' . . Fig. 8-8 SPECIES: Halistemma rubrum FAMILY: Agalmidae SUBORDER: Physonectae SIZE: to 2 m long DESCRIPTION: nectosome of up to 60 nectophores, colony often rose colored, tentacles with vermilion, unicornuate tentila, palpons long and extensile. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION:~ Atlantic, Med. i .l Fig. 5-9 SPECIES: Lychnagalma utrlcularla FAMILY: Agalmidae SUBORDER: Physonectae SIZE: to 20 cm long DESCRIPTION: 11-25 nectophores in 2 rows, bracts flimsy, with 2 distal points, tentila unique, with red cnidoband and 8 terminal filaments, gastrozooids on stalks. LUMINESCENCE: Not, but three other genera in this family are definite (Herring, 1987). DISTRIBUTION: Atlantic, Indian, Med. in deep water r 77 Fig. S-10 SPECIES: Marrus orthocanna FAMILY: Agalmidae SUBORDER: Physonectae SIZE: large DESCRIPTION: large nectophores, long spindle or club-shaped gastrozooids, scarlet colored stem. LUMINESCENCE: Herring (1987) lists three other genera in this family as definite. DISTRIBUTION: arctic Atlantic in deep water '" 1 \ r I. \ FICJO JI_-lp:-i).~ ,¿ _ ... d _ .. .. t.. _ A.....;B.-... ft r- Qi; t.,... pt ~ Fig. S-11 SPECIES: Nanomia bijuga FAMILY: Agalmidae SUBORDER: Physonectae SIZE: 10-45 cm long DESCRIPTION: nectosome 1/5 of total length, square nectophores in 2 rows, unicornuate tentilla, dark red splotches on stem. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. Fig. S-12 SPECIES: Nanomia cara FAMILY: Agalmidae SUBORDER: Physonectae SIZE: to 50 cm long DESCRIPTION: to 30 nectophores in 2 rows, horizontally flattened, nectosome about 1/5 total length, unicornuate tentila with pigmented cnidoband. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. 78 Fig. 5-13 SPECIES: Physophora hydrostatica FAMILY: Physophoridae SUBORDER: Physonectae SIZE: to 12 cm high DESCRIPTION: conspicuous plum-color apical pneumatophore, 8-12 nectophores in 2 rows, large green-pink palpons around base of nectosome, tentacles below it. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. 5-14 SPECIES: Abyla haeckeli FAMILY: Abylidae SUBORDER: Calycophorae SIZE: ant. nectophore 5 mm high DESCRIPTION: 11-faceted anterior nectophore, tubular nectosacs, stem withdraws into hydroecium. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian. Med. Fig. 5-15 SPECIES: Abylopsis eschscholtzi FAMILY: Abylidae SUBORDER: Calycophorae SIZE: 6 mm high DESCRIPTION: cubic anterior nectophore, nectosac directed laterally, larger faceted posterior nectophore with finely toothed edges. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Red Sea, Med. in shallow water 79 Fig. S-16 SPECIES: Abylopsis tetragona FAMILY: Abylidae SUBORDER: Calycophorae SIZE: to 35 mm high DESCRIPTION: small cuboidal anterior nectophore, larger posterior nectophore 3x long as broad, with 5 terminal teeth of various lengths. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: common in Atlantic, Med. Fig. S-17 SPECIES: Amphiearyon aeaule FAMILY: Prayidae SUBORDER: Calycophorae SIZE: about 5 mm diameter DESCRIPTION: 1 large rounded nectophore and 2 small, flattened vestigial nectophore with nectosac not open to exterior. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med.' Fig. S-18 SPECIES: Bassia bassensis FAMILY: Abylidae SUBORDER: Calycophorae SIZE: to 15 mm high DESCRIPTION: small, cuboidal anterior nectophore, faceted posterior nectophore, teeth. " 2x long as broad, fairly short terminal LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide, usually near su rface 80 Fig. S-19 SPECIES: Ceratocymba sagittata F AMIL v: Abylidae SUBORDER: Calycophorae SIZE: to 60 mm high DESCRIPTION: anterior nectophore with long, pyramidal apical projection, long tubular nectosac, posterior nectophore with large ventral terminal tooth and toothed margins. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. Fig. S-20 SPECIES: CheJophyes appendicuJata FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: 30 mm high DESCRIPTION: large anterior nectophore with 3 ridges and large nectosac, smaller posterior nectophore has ventral ridges that end in terminal teeth. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide, "the commonest of all siphonophores" SPECIES: CheJophyes contorta FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: 10 mm high DESCRIPTION: large anterior nectophore with 3 ridges, ventral facet slightly twisted, smaller posterior nectophore with 2 terminal teeth. zmm LUMINESCENCE: Herring (1987) lists this genus as definite. DISTIBUTION: Atlantic, Pacific, Indian, Med. j . 81 Fig. 8-22 SPECIES: Chuniphyes multidentata FAMILV: Clausophyidae SUBORDER: Calycophorae SIZE: to 60 mm high DESCRIPTION: anterior nectophore with pointed apex, 4 ridges branching to 6, posterior nectophore with 3 ridges branching to 6, ending in 6 terminal teeth. LUMINESCENCE: unknown DISTRIBUTION: world-wide in deep water Fig. 8-23 SPECIES: Clausophyes ovata FAMILV: Clausophyidae SUBORDER: Calycophorae SIZE: to 40 mm high DESCRIPTION: soft, pear-shaped anterior nectophore, larger posterior nectophore with tapered apex. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. in deep water Fig. 5-24 SPECIES: Diphyes dispar FAMIL v: Diphyidae SUBORDER: Calycophorae SIZE: to 50 mm high DESCRIPTION: anterior nectophore laterally compressed with 5 ridges, dorsal ridge serrated with terminal tooth, posterior nectophore with smooth edges and teeth. LUMINESCENCE: Herring (1987) lists this' genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. 82 Fig. S-25 SPECIES: Enneagonum hyalinum F AMIL v: Abylidae SUBORDER: Calycophorae SIZE: 15 mm high DESCRIPTION: consists of cuboidal anterior nectophore only, with strong dorsal ridge and serrated basal edges and teeth. LUMINESCENCE: Herring (1987) lists 4 genera in this family as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. Fig. 5-26 SPECIES: Eudoxoides spiralis FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: to 11 mm high DESCRIPTION: anterior nectophore with 5 twisted longitudinal ridges, 4 of which reach apex, no posterior nectophore. LUMINESCENCE: Herring (1987) lists 3 genera in this family as definite. DISTRIBUTION: world-wide and common SPECIES: FIG. 128. Eudoxoies spira/is (Bigelow) Polygastric phase, X 10 (from Totton & Fraser, 1955). Hippopodius hippopus FAMILV: Hippopodiidae SUBORDER: Calycophorae SIZE: to 30 mm high DESCRIPTION: up to 12 horseshoe-shaped nectophores stacked above each other, without teeth or serration, no bracts, mesoglea turns opaque white on contact. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide Fig, '18. Hippopodius hippopus; dorsal view or iieclophore 83 Fig. 5-28 SPECIES: Lensia campanella FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: to 6 mm high r DESCRIPTION: anterior nectophore twisted at apex (or possibly not in live specimens), ridges indistinct, orange-red spots on nectophores. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fic. '00, i- -- (M_~ Poyp pb GL il Ro A. Id la 'I of th ante neopho. x )0; B, ri¡hi 1a 'fnr of th por DChore x ..6 (fro Toa 1932. fi. 35) Fig. 5-29 SPECIES: Lensia conoidea i i .":" ~,I:1 FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: to 45 mm high DESCRIPTION: pyramidal anterior nectophore with 5 ndges and smooth facets, large nectosac, postenor nectophore with 5 ridges and facets. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. FiB_ 2;. ulUio coniidm; latt'ral vi_ of auptor ncdpbor. i4.~ mm. )onii. frem -Thor" St. 36. F1l. 28. ulUia t'idfØ; nntnl'"kw Fig. 5-30 SPECIES: Lensia fowleri FAMIL v: Diphyidae SUBORDER: Calycophorae SIZE: to 12 mm high DESCRIPTION: anterior nectophore with 5 ridges, smooth facets, no basal teeth, posterior nectophore 3/4 length of antenor. LUMINESCENCE: unknown Fig. 36. umia IDllui; "ten) vi"" or superior Deopbo~ about .. mm. blgh. colleced by H. Ji. S. "'Res" Ja the Bay 01 Bisy (Mas, Comp, Zool, Cat, No, 775). Fig 37. ùlUia fowlu;; lat.er view ot bas portion or iu~rior neclo. pbOl about 7.6mm. blgb, collected DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Mad. by H. M. S. "Re." in Uie Bs:r of Bisy (M~. Comp. Zo!. Cat. No, 775). 84 Fig. 5-31 SPECIES: Lensia meteori FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: ' to 5 mm high DESCRIPTION: anterior nectophore with indistinct ridges and smooth conical surface, posterior nectophore non-existent or unknown. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. Fig. 5-32 SPECIES: Lensia multicristata FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: to 20 mm high DESCRIPTION: anterior nectophore with 7 ridges, 5 reaching the apex and basal margin, posterior nectophore with 5 ridges. LUMINESCENCE: unknown Fig. 40. un~ia mulUcrislala: su- DISTRIBUTION: Atlantic, Pacific, Indian, Med. ptrlor ntdophort. i 1 mm. high. uTbor" St. 217. Fig. 5-33 SPECIES: Lensia subtils FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: to 20 mm high DESCRlpT:0N: anterior nectophore with 5 indistinci ridges, smooth surface and rounded apex, posterÏr1r nectophore with 5 ridges, yellow pigmelL" .,In. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. near surface 85 Fig. 5-34 SPECIES: Lensia subtioides FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: to 7 mm high DESCRIPTION: antenor nectophore with 5 ndges, less distinct at apex, no basal tooth on dorsal ndge, postenor nectophore with Fig. 15, un.ia sublioid..; latera view of superior nec- tophore. 6 mm. long, from "Thor" St, 80. 5 ridges. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. near surface Fig. 5-35 SPECIES: Li/yopsis rosea F AMIL v: Prayidae SUBORDER: Calycophorae SIZE: to 20 cm (?) DESCRIPTION: 2 large, equal, opposed nectophores of roughly conical shape, with large nectosacs, stem with large bracts, red pigment spots on stem eudoxids. LUMINESCENCE: Herring (1987) lists 5 genera in this family as definite. DISTRIBUTION: Med., rare " ~. Fig. 8-36 SPECIES: Muggiaea at/antica FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: to 7 mm high DESCRIPTION: antenor nectophore with 5 serrate ndges, somatocyst reaches top of nectosac, no posterior nectophore. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. common ,¡, r 86 Fig. 8-37 SPECIES: Muggiaea kochi FAMILV: Diphyidae SUBORDER: Calycophorae SIZE: to 5 mm high DESCRIPTION: anterior nectophore with 5 ridges, somatocyst reaches halfway up nectosac, no posterior nectophore. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. common Fig. 8-38 SPECIES: Rosacea cymbiformis F AMIL v: Prayidae SUBORDER: Calycophorae SIZE: to 2 m long extended DESCRIPTION: 2 large, unequal, oblong nectophores with small nectosacs, soft jelly, somatocyst extends below nectosac, numerous stem groups with large bracts. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. Fig. 8-39 SPECIES: Rosacea plicata FAMILV: Prayidae SUBORDER: Calycophorae SIZE: 1-2 m extended (?) DESCRIPTION: 2 large, unequal oblong nectophores, small nectosacs, somatocyst stops above nectosac, numerous stem groups with large bracts. LUMINESCENCE: Herring (1987) lists this genus as definite. Fig. 9. RNa~a plicaa; pair ot nectopbore. 16 mm. DISTRIBUTION: Atlantic, Pacific, Indian, and 13 mm., sWI connected, trom Berng Sea. uAJba. tross" St. 47671. Antarctic, Med. 87 Fig. 5-40 SPECIES: Sphaeronectes gracils FAMILY: Sphaeronectidae SUBORDER: Calycophorae SIZE: 8 mm diameter DESCRIPTION: single, spherical nectophore with hemispherical nectosac and curved, elongate somatocyst. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Med. SPECIES: Sphaeronectes irregularis FAMILY: Sphaeronectidae SUBORDER: Calycophorae SIZE: 7 mm diameter DESCRIPTION: single, pear-shaped nectophore and nectosac, short, straight somatocyst. LUMINESCENCE: unknown ~ Fig. 5-41 _./;f i ¡¡-//;~""\ " / ,j,";¡'" \ - _/:~ i '. fl----~ ' \... ";.-=- . ~..' DISTRIBUTION: Atlantic, Pacific, Med. Fig. 5-42 SPECIES: Sulculeolaria biloba FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: to 1 + m long, extended DESCRIPTION: conical anterior nectophore, nectosac opens obliquely, short somatocyst, 2 large basal lobes, posterior nectophore without basal lobes or teeth. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. 88 Fig. 5-43 SPECIES: Sulculeolaria chuni // J% / . i." :: FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: 10+ cm long DESCRIPTION: conical anterior nectophore, long, thin somatocyst, 2 short basal lobes, posterior nectophore without basal lobes or teeth. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Red ~,._;:" h'Iii/1~/? / ///f i,' ..,,' \ .' ,". // i:g' //./ l' ~ ~4..:. !_;.~/ ..~~ .ILf -L'1-., ',~ _'---://~, ' ~i' t. "", " // \..~ ~!,' """', \_._',. /¡,' t~~;;,:;/ P' ,', .l,~"A'" \ '\~. t"...-. ':~ . l(/~~-:.i-"~ r'7'~:\\~ -~-"" \ Sea, Med. Fig. 5-44 SPECIES: Sulculeolaria quadrivalvis FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: 1 + m long, extended DESCRIPTION: conical anterior nectophore, medium length somatocyst, 2 large basal lobes, posterior nectophore with 2 lateral and 2 dorsal teeth at opening. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. Fig. 5-45 SPECIES: Sulculeolaria turgida FAMILY: Diphyidae SUBORDER: Calycophorae SIZE: 20 cm long DESCRIPTION: conical anterior nectophore, small somat!)cyst, 2 basal lobes, no teeth, posterior nectophore with single large basal lobe. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. 89 Fig. 8-46 SPECIES: Vogtia glabra FAMILY: Hippopodiidae SUBORDER: Calycophorae SIZE: 10 cm long (?) DESCRIPTION: up to 12 similar nectophores, partly overlapping in 2 rows, with smooth, rounded exterior, 3 lateral ridges and 2 dorsal humps. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. 20. Voglia glabra; dorsal view of neetophore, 11 mm. in length from "Thor" St. 91. Fig. 5-47 SPECIES: Vogtia pentacantha FAMILY: Hippopodiidae SUBORDER: Calycophorae SIZE: 10 cm long (?) DESCRIPTION: up to 12 similar nectophores, partly overlapping in 2 rows, pentagonal in section, with teeth on edges but not surfaces of facets. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. in deep water Fig. 5-48 SPECIES: Vogtia spinosa FAMILY: Hippopodiidae SUBORDER: Calycophorae SIZE: 10 cm long (?) DESCRIPTION: up to 12 similar nectophores, partly overlapping in 2 rows, pentagonal in section, with teeth on edges and surfaces of facets. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. in deep water 90 i t 91 Ctenophores The phylum Ctenophora comprises perhaps a hundred or more gelatinous animals, almost all of which are planktonic. With few exceptions, ctenophores are strongly bioluminescent, emitting light from the gastrovascular canals, and sometimes from luminous secretions. They occur from shallow to deep water in all oceans, and are predators on other zooplankton. Ctenophores are fragile and difficult to collect or preserve. Many new species have only recently been descnbed using in-situ methods, and undoubtedly many more species remain to be discovered, especially in deep water (Harbison and Botkin, in prep; Madin and Harbison, 1978a,b ). The classification of the phylum remains somewhat unsettled due to the recent influx of new species and higher taxa. At least five orders are represented in the Mediterranean plankton and are included here. 1. Cydippida. The cydippids generally have oval or cylindncal bodies with a mouth at one end and a statocyst at the other. They range in size from a few mm to nearly 30 cm. Cydippids have two long tentacles which are extended outside the body for fishing, but can be withdrawn into it. Division into familes is based on the structure of the tentacles , their position (emerging near the oral or the aboral end of the body), body shape, and connections of the internal gastrovascular canals. 2. Lobata. In these ctenophores the oral end of the body is enlarged into two oral lobes, which are spread out as food-catching surfaces. The external tentacles are reduced to a veil of fine side-branches or tentila which cover the surfaces of the lobes and parts of the body. Lobates have elongate, flattened bodies, and range from about 10 mm to a meter or more across. Familes are distinguished on the basis of body shape, arrangement of canals or the presence of particular structures. 3. Thalassocalycida. This order contains a single genus which occurs mainly in midwater. It is most similar to the lobates, but the oral lobes are connected to form a continuous, medusa-like belL. 4. Cestida. The two genera in this order have similar morphologies, but differ in size. The body is extremely flattened and elongate, looking like a transparent belt. The tentacles are within grooves on one edge of the body, and tentila cover the flat surfaces of the body. 5. Beroida. These ctenophores lack tentacles altogether. The body is quite flattened, and oval or conical in outline; size ranges from a few mm to 20 cm or more. Beroids have a large, expansive mouth and stomodeum with which they engulf other ctenophores as prey. The classification used here is based on Harbison and Madin (1982), Harbison (1985) and Mils (1987). Descriptions, ilustrations and distributional data are compiled from Carré and Carré (1989), Chun (1878, 1880, 1898), Fedele (1940), Harbison (pers. comm.), Komai (1918), Madin (unpubl. data), Madin and Harbison (1978a,b), Mayer (1912), Mills (1987), Moser (1910), Tregouboff and Rose (1957). 92 Terminology: auncles - 4 flattened or elongate structures on lobate ctenophores that attach near the base of the lobes colloblasts - glue-cells on tentacles and tentila which stick to prey comb rows- 8 mendional rows of ctenes which provide propulsion ctenes - plates of fused cilia that beat like paddles, arranged in comb rows diverticula - side branches off gastrovascular canals that sometimes anastomose meridional canals - 8 main gastrovascular canals running longitudinally through body or into lobes paragastnc canals - canals running along each side of the stomodeum stomodeal canals - 4 meridional canals in the stomodeal plane of the body stomodeal plane - the plane of symmetry in which the flattened stomodeum lies stomodeum - the large first part of the gut into which prey is taken tentacle sheaths - cavities in the body of cydippids into which the tentacles can be withdrawn tentacular plane - the plane of symmetry, orthogonal to the stomodeal, in which the tentacle bulbs and sheaths lie tentilla - side branches off the tentacles, may be simple, complex or coiled 93 Fig. C-1 SPECIES: Callanira bia/ata FAMILV: Mertensiidae ORDER: Cydippida SIZE: to 30 mm high DESCRIPTION: body flattened in stomodeal plane, with 2 long aboral projections, tentacles emerge near aboral end, and have many fine tentila. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. in deep water Fig. C-2 SPECIES: Eup/okamis stationis FAMILV: Euplokamidae ORDER: Cydippida SIZE: to 25 mm high DESCRIPTION: cylindncal or ovoid body, comb rows extend 2/3 body height, tentacles emerge near aboral end, with fine, helically coiled tentila. LUMINESCENCE: probable but not published DISTRIBUTION: Med. ~ ~ ¿ß_,~ Fig. C-3 SPECIES: Haecke/ia bimacu/ata F AMIL v: Haeckelidae ORDER: Cydippida SIZE: 3 mm DESCRIPTION: ellpsoidal body, tentacles lack tentila, emerge near mouth, large orange spots on stomodeum, small red spots along comb rows, no green pigmentation. LUMINESCENCE: probable, but not published DISTRIBUTION: Med. 94 Fig. C-4 SPECIES: Haeckelia rubra F AMIL V: Haeckeliidae ORDER: Cydippida SIZE: to 10 mm high DESCRIPTION: body short and squareish, large mouth, orange tentacle sh'eaths, tentacles emerge near mouth, lack tentilla and colloblasts, but have nematocysts. LUMINESCENCE: probable, but not published DISTRIBUTION: Atlantic, Pacific, Med. in shallow water Fig. C-5 SPECIES: Hormiphora plumosa FAMILV: Pleurobrachiidae ORDER: Cydippida SIZE: to 20 mm DESCRIPTION: ovoid body with elongate oral end, comb rows about 1/2 body height, tentacles emerge aborally, with simple and hand-shaped tentilla. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. in shallow water Fig. C-6 SPECIES: Hormiphora spatulata FAMILV: Pleurobrachiidae ORDER: Cydippida SIZE: to 21 mm high DESCRIPTION: ovoid body, not compressed, tentacles sheaths diverge orally from stomodeum, comb rows almost as long as body, tentacles with 2 sizes of tentila. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. ,=, 95 Fig. C-7 SPECIES: Lampea pancerina FAMILY: Lampeidae ORDER: Cydippida SIZE: to 75 mm high DESCRIPTION: cylindrical body with large, extensile mouth, comb rows 2/3 body height, tentacles emerge orally, with simple tentila that coil up. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. in shallow water Fig. C-8 SPECIES: Pleurobrachia pileus FAMILY: Pleurobrachiidae ORDER: Cydippida SIZE: to 20 mm high DESCRIPTION: ovoid body, not compressed, comb rows 3/4 body height, tentacle sheaths distant from stomodeum, tentacles emerge aborally, with fine tentila. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. in shallow water " Fig. C-9 t f SPECIES: Bathocyroe fosteri FAMILY: Bathocyroidae ORDER: Lobata SIZE: to 40 mm high I: DESCRIPTION: short body with broad oral lobes that flap to swim, auricles flat and broad, stomach red, paragastric canals extend onto inner lobe surfaces. LUMINESCENCE: yes, along comb rows DISTRIBUTION: Atlantic, Pacific, Med. in deep wate r , ' eli &l..¡..ur: do gut; oqttii,pi.inAbbrti... odea¡ i=;pp, au aurj cob nJ¡siIe.vifiofc:;I'. ca; W. pat: pole pbic; so. lt¡ Ø.lUbstt ca; n, iubtCDEa cm; rh. tmtac bulb. 96 Fig. C-10 SPECIES: Bolinopsis vitrea FAMILY: Bolinopsidae ORDER: Lobata SIZE: to 80 mm high DESCRIPTION: oval body, compressed in stomodeal plane, oral lobes 1/2 body height, auricles slender, stomodeal canals make simple loops in oral lobes. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. Fig. C-11 SPECIES: Deiopea kaloktenota FAMILY: Bolinopsidae ORDER: Lobata SIZE: to 50 mm high DESCRIPTION: wide body, strongly compressed in stomodeal plane, short comb rows with few large, widely spaced ctenes, lobes 1/2 body height. LUMINESCENCE: Herring (1987) lists this Sl:iIOC\"st ~~.. Infundibulum .. -~' ~ .; ..- I. ~ ~ Auricle ß,: :~:. ~ canai~~ ! ~. canal ~ co- _i ",.. yCombrow Meridional ¡; : \ -t. .~. Meridionoil ~.,.f '.,.., ,"', ,", : ,f~! .111'\" " - -. ,-" ", " 7\" ~ 0"1100. ¿. ~ i ~ ' Ti:ntac\e Mouth ~ \\') ; . -ø bulb Tent:icle ~ side branches\.~/)~ (i~' " ~L""","',?I~~\ genus as definite. DISTRIBUTION: Atlantic, Med. in deep water Fig. C-12 SPECIES: Eurhamphaea vexillgera FAMILY: Eurhamphaeidae ORDER: Lobata SIZE: to 150 mm high DESCRIPTION: long, narrow body with 2 aboral processes, compressed in stomodeal plane, rows of conspicuous red vesicles under comb rows that release ink. LUMINESCENCE: Herring (1987) lists this genus as definite. Ink is luminescent. DISTRIBUTION: Atlantic, Pacific, Med. 97 Fig. C-13 SPECIES: Leucothea multicornis FAMILY: Leucotheidae ORDER: Lobata SIZE: to 250 mm high DESCRIPTION: long body, flattened in stomodeal plane, voluminous oral lobes, extensile papillae on body and lobes, long, sinuous auricles, 2 aboral trailng tentacles. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. SPECIES: Thalassocalyce inconstans FAMILY: Thalassocalycidae ORDER: Thalassocalycida SIZE: to 150 mm diameter DESCRIPTION: body umbrella-shaped when expanded, contracts to spherical or bilobed form, stomodeum on peduncle, short comb rows, delicate tentacles with tentilla. LUMINESCENCE: unknown ..,...~ Fig. C-14 .~.. ,-n '" ....'''~ /~ .. ~ ~ -- \. . P i.ll~. ~';l \ .~¡ ~j;;-. ';i i'-v\f-\.' - ; j;l ¡if'f"'hi iI ( 1 l -, " ,r" L ~" J' ... \, , \ " , ;, 1- ( ¡ i'" l /I " "I \0 , 'I ;' 1 ~,: ,:,;,....,..\,.."~"'., . ,".. \", .."' ~.. ,. '." '. .... ,. c" ¡ ~". .:. .. ...... '.: .' ~L~~ "..'''¿~J'''' " " ......' ..,.. I . . DISTRIBUTION: Atlantic, Pacific, Med. Fig. C-15 SPECIES: Cestum veneris FAMILY: Cestidae ORDER: Cestida SIZE: to 1 m long (wide) DESCRIPTION: flat, belt-shaped body with central stomodeum, comb rows extend along entire aboral edge, tentacles along oral edge with tentila covering body sides. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide 98 Fig. C-16 SPECIES: Velamen parallelum FAMILV: Cestidae ORDER: Cestida SIZE: to 150 mm long (wide) DESCRIPTION: body shape like Cestum but smaller, gonads form dark dashes along aboral edge, stomodeum short, mendional canals converge in center of body. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. Fig. C-17 SPECIES: Berae farskali FAMIL v: Beroidae ORDER: Beroida SIZE: to 20 cm high DESCRIPTION: conical body with wide, flaring mouth, anastomosing diverticula from meridional and paragastric canals, dark pink color overalL. LUMINESCENCE: Herring (1987) lists this genus as definite. Details in Panceri (1872). DISTRIBUTION: Atlantic, Pacific, Med. Fig. C-18 SPECIES: Berae mitrata FAMILV: Beroidae ORDER: Beroida SIZE: to 30 mm high DESCRIPTION: compressed, mitre-shaped body, large mouth, the few mendional diverticula don't anastomose, but some join paragastncs, orange spot in mid body. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. 99 Fig. C-19 SPECIES: SeTae ovata FAMILY: Beroidae ORDER: Beroida SIZE: to 115 mm high DESCRIPTION: body mitre-shaped, moderately compressed, milky to pink, meridional diverticulae anastomose with paragastric branches, not each other. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. 100 101 Polychaetes and Nudibranchs Planktonic polychaetes include both adult forms and numerous larval stages of benthic species. The holoplanktonic species typically have large paddle-like parapodia, swim in an undulating fashion and are predators on other zooplankton. At least one widespread genus, Tomopteris is reported to be luminescent. Tomopterids in deep water attain lengths of up to 20 cm. There are six familes of polychaetes with pelagic genera that are reported to occur in the Mediterranean by Tregouboff and Rose (1957). However only 2 genera have been reported in more recent studies of the western Mediterranean plankton, and those are described here. There are two genera of holoplanktonic nudibranchs, Phyllrhoe and Cephalopyge, of which the first is luminescent. Description of Phyllrhoe is taken from Lall and Gilmer (1989) and Tregouboff and Rose (1957). 102 Fig. P-1 SPECIES: Calizonella lepidota FAMILV: Alciopidae SUBCLASS: Errantia , SIZE: DESCRIPTION: elongate body, large round red eyes, parapodia with 1 cirriform appendage. LUMINESCENCE: Herring (1987) lists 3 genera in this family as uncertain. DISTRIBUTION: Fig. P-2 SPECIES: Lopadorhynchus uncinatus F AMIL v: Phyllodocidae SUBCLASS: Errantia SIZE: to 20 mm long DESCRIPTION: broad, tapered body, 4 antennae, no palps, may be dark colored. LUMINESCENCE: unknown DISTRIBUTION: Med., Atlantic. Fig. P-3 SPECIES: Tomopteris helgolandica FAMILV: Tomopteridae SUBCLASS: Errantia SIZE: to 200 mm DESCRIPTION: Body usually transparent, with long trailing antennae, paired paddle-like parapodia with conical lobes. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide 103 Fig. P-4 SPECIES: Vanadis crystallna FAMILY: Alciopidae SUBCLASS: Errantia SIZE: DESCRIPTION: very elongate body, head with conspicuous round red eyes, parapodia with single cirriform appendages. LUMINESCENCE: Herring (1987) lists 3 genera in this family as uncertain. DISTRIBUTION: Atlantic, Med. Va... i. .. .~. -., ," -,-- .' -,.1::,-,1'"'1"";:,,1 Fig. P-5 SPECIES: Vanadis formosa FAMILY: Alciopidae .-..... SIZE: .'~ f SUBCLASS: Errantia .: "." DESCRIPTION: very elongate body, head with conspicuous round red eyes, parapodia with single cirriform appendages. .~' .' (" :~~ .; " va.r. ,: .; LUMINESCENCE: Herring (1987) lists 3 genera , ~E:d:~J in this family as uncertain. DISTRIBUTION: Atlantic, Med. Fig. P-6 SPECIES: Phyllirhoe bucephala FAMILY: Phyllroidae ORDER: Nudibranchia SIZE: to 40 mm DESCRIPTION: flattened, transparent,leaf-like body with expanded tail, conspicuous internal organs, 2 long anterior tentacles, gils absent and foot reduced. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. ~~ . ..::r; .~.,~ - .' ::....................~ '..",.,~ ..-- ',., .:~:-..-., . ... '. i.. -i .....,...... . ~~. P6.bu. 104 105 Pelagic Tunicates: Thaliacea and Larvacea The class Thaliacea includes three orders -- the colonial Pyrosomes, which may range in length up to a meter or more and are strongly bioluminescent, the doliolids, and the salps, which are filter-feeders with tubular bodies and alternating generations. The class Larvacea comprises a single order of small, tadpole-like organisms that produce an external mucous filtering structure called a house. In some genera both the animal and the house are luminescent. They are widely distributed and often abundant. 1. Pyrosomida. The colonies are made up of numerous small ascidian-Iike zooids embedded in a stiff matrix or tunic. The colony is tubular, with a single terminal opening. Water pumped through each zooid for filter-feeding passes into the lumen of the colony and out the opening for jet propulsion. External morphology of the colony is variable, and although pyrosomas are unmistakable, specific identification is difficult, and there are many uncertain species and synonyms. 2. Doliolida. This order of the Thaliacea comprises small, barrel shaped animals with circumferential muscle bands used to create jet propulsion. The life cycle involves 6 different stages, and at one point includes a large polymorphic colony of thousands of zooids, which may attain lengths over 1 m. These colonies are fragile and rarely collected intact. The taxonomy is usually based on the gonozooid (sexually reproducing) stage, which is single and free-swimming. The oozooid (asexually budding) stage develops into the "nurse" which pulls the polymorphic colony; since this form is fairly sturdy it is often collected intact. Included here are descriptions of the gonozooid (gz) and nurse stages. Doliolids are easily recognized, but not easily identified to species. 3. Salpida. This order is of larger filter feeding animals, also with circumferential muscle bands. The salps alternate between two forms, an asexually budding solitary stage and a sexually reproducing aggregate stage. The aggregate salps usually remain connected together in chains or whorls of various types. The individual animals range in size from 5 to over 100 mm, and chains can be several m long. Descriptions and illustrations of both solitary (s) and aggregate (a) forms are included here. 4. Larvacea. This class is divided into 3 familes of small (1-10 mm) animals consisting of a trunk and long, flat taiL. Much of the taxonomy is based on arrangement of internal organs, which are difficult to see without using microscopy on fixed specimens. Descriptions are included here only for the more common Mediterranean species, and those characteristics likely to be most apparent in living, whole animals are emphasized. Descriptions, ilustrations and distributional data for Thaliaceans are compiled from Bracconot (1970, 1971), Madin (1974), Madin and Harbison (1978), Madin et al. (1981), Sewell (1953), van Soest (1973, 1974a,b, 1975), Thompson (1948) and Tregouboff and Rose (1957). Information for Larvaceans is mainly from Fenaux (1967), with other material from Galt (1989), Thompson (1948) and Tregouboff and Rose (1957). 106 Terminology: body muscles - circumferential muscle bands around the tubular body of salps and doliolids, continuous in the former and interrupted ventrally in the latter caeca - blind extensions of the gut cluster - loose radial group of many aggregate Cyclosalpa polae , endostyle - ventral organ in thaliaceans and larvaceans that secretes mucus gonozooid - free-swimming sexually reproductive stage of doliolid helical chain - chain of aggregate salps arranged in double helix "light organs" - stripes of opaque tissue along sides of Cyclosalps, sometimes thought to be luminescent linear chain - chain of aggregates all aligned with zooid axes nearly parallel to chain axis longitudinal muscle - body muscle of salps that runs longitudinally on the dorsal su rface nurse - later growth stage of doliolid oozooid that loses digestive organs and serves only for locomotion of colony oblique chain - chain of aggregate salps aligned with zooid axes at oblique angle to chain axis peduncle - mid-ventral projection on aggregate cyclosalps that attaches them into whorl or cluster radial whorl - chain of 10-15 cyclosalps arranged like segments of an orange spiracles - cilated openings into the pharynx of larvaceans that pump water through the pharyngeal filter net ' stolon - strand of tissue that buds asexually produced aggregate salps, may remain attached to parent solitary salp while developing subchordal cells - large cells present in the tails of larvaceans, often in speciesspecfic numbers test or tunic - the stiff gelatinous part of the body of a salp or pyrosome transverse chain - chain of aggregate salps aligned with zooid axes perpendicular to the chain axis 107 Fig. T-1 SPECIES: Pyrosoma at/anticum ORDER: Pyrosomida CLASS: Thaliacea SIZE: colony to 60 em DESCRIPTION: cylindrical colony, colorless to pin'k or brownish, test fairly rigid with dentate processes of varying length, zooids irregularly arranged in larger colonies. LUMINESCENCE: Herring (1987) lists this genus as definite. One of the most brightly luminous organisms. DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-2 SPECIES: Dolio/etta gegenbauri ORDER: Doliolida CLAss: Thaliacea SIZE: gz 10 mm DESCRIPTION: gz: barrel shaped, 8 circular body muscles, gut mid-ventral, in tight dextral coil. nurse: with muscles 3,4 wider than the others. LUMINESCENCE: Herring (1987) lists the genus Doliolurn in this order as uncertain. I'ïf:. i. i,.t:~:~'::.i'. ::;~:i:~:'."';i:i 1.11;..:~""''''''. 1......./0...101 ....11' :".L DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-3 SPECIES: Do/io/um denticu/atum ORDER: Doliolida CLAss: Thaliacea SIZE: gz 10 mm, nurse 15 mm DESCRIPTION: gz: barrel-shaped, with 8 body muscles, scalloped oral valve, gut in a broad curve on ventral floor. nurse: body muscles fused into continuous sheet. LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION: Atlantic, Pacific, Indian, Med. \ ....Ii ~11i" " \\ ". .-, ". 108 Fig. T-4 SPECIES: Dolio/um mulleri ORDER: Doliolida CLASS: Thaliacea SIZE: gz 4 mm, nurse 8 mm DESCRIPTION: gz: barrel-shaped body, 8 muscles, gut forms upright U or S-shaped loop. nurse: body muscles fused into I. , continuous sheet. LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION: Atlantic, Med. Fig. T-5 SPECIES: Cyc/osa/pa affinis ORDER: Salpida CLASs: Thaliacea SIZE: s to 80 mm, a to 60 mm DESCRIPTION: s: cylindrical body, 7 body muscles, 1 st 2 interrupted dorsally, no "light organs". a: 4 body muscles, short ventral peduncle, gut in open loop, radial whorls, connected in chains. LUMINESCENCE: Herring (1987) lists this genus as uncertain DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-6 SPECIES: Cyc/osa/pa pinnata ORDER: Salpida CLASS: Thaliacea SIZE: s to 75 mm, a to 65 mm Q,O DESCRIPTION: s: 7 body muscles, interrupted dorsally, linear gut with 2 caeca, 5 purple "light organs" on each side. a: 4 body muscles, short peduncle, 1 light organ on each side, in radial whorls of 10-15 salps. LUMINESCENCE: Herring (1987) lists this genus as uncertain DISTRIBUTION: Atlantic, Med. 109 Fig. T-7 SPECIES: Cyclosalpa polae ORDER: Salpida CLASS: Thaliacea SIZE: s to 80 mm, a to 40 mm DESCRIPTION: s: 7 body muscles, interrupted dorsally, 6th forms longitudinal muscle, 5 white "light organs" on each side. a: 4 body muscles, long peduncle, 1 light organ each side, in clusters of up to 200 salps. LUMINESCENCE: Herring (1987) lists this genus as uncertain DISTRIBUTION: Atlantic, Med. SPECIES: Helicosalpa virgula ORDER: Salpida CLASS: Thaliacea SIZE: s to 180 mm, a to 35 mm DESCRIPTION: s: 7 body muscles interrupted by paired longitudinal muscles, 1 "light organ" on each side, linear gut with 2 caeca. a: asymmetric, 4 body muscles, testis in posterior projection, helical chain. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-9 SPECIES: lasis zonaria ORDER: Salpida CLASS: Thaliacea SIZE: s to 65 mm, a to 50 mm DESCRIPTION: s: elongate, prismatic with stiff test, 5 broad body muscles, stolon coils around compact gut. a: stiff test, asymmetrical, 5 broad muscles, in tight linear chain. LUMINESCENCE: unknown ttfEC~ ~'of. 1._...¡;I&It ("nu. .1... ..¡,... I...,ii\h.:: 11. ~'('. :?-.\.:al" (Mm. .J....I ,"",.., I,,'l'~~h.:! ,"D. t"i" .'._.\':."'01.1,, ¡..no, .1..,.1 ,....'. \l-."i 1".- !'II".. ,....".1 .ii", ~1~lr..ir !~'l~., DISTRIBUTION: Atlantic, Pacific, Indian, Med. F.i¡ i. :;_f. \'_ .\1...ri~ .i..._ 110 Fig. T-10 SPECIES: Ihlea punctata ORDER: Salpida CLASS: Thaliacea SIZE: s to 70 mm, a to 23 mm DESCRIPTION: S: 9 wide body muscles, some fused dorsally, yellow pigment band around body, round gut. a: 6 asymmetric body , muscles, orange-red spots on ventral side, linear chain. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. ~~, Fig. T-11 SPECIES: Pegea bicaudata ORDER: Salpida CLASS: Thaliacea SIZE: s to 72 mm, a to to 80 mm DESCRIPTION: s: globular test with diffuse yellow or red pigment, 4 body muscles, stolon coils around gut. a: cylindrical test with yellow pigmentation posteriorly, 2 "tails", 4 body muscles, transverse chain. LUMINESCENCE: unknown ---=~ :; - :: '/:r~,~~, ' / ¿t~ ".-~ ", ( ¿\v pJ!\ i i "".,,.j~_,7 ; \\ .'~~'i--o=", ./ ~~~'?/ /' -'~I ~/1j, ,~.. ., " , ;;," ?\~ \ i0: .~~\ i!~~. ~~\ ,~ -=1 ,,~~ J \=~'\\,! V~'\il ',. -" ~ ".,,~ ( 4~~~j) DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-12 SPECIES: Pegea confoederata ORDER: Salpida CLASS: Thaliacea SIZE: s to 90 mm, a to 110 mm DESCRIPTION: s: test more cylindrical, with reticulate brown pigmentation, reddish- brown spherical gut. a: short, plump body with thick test around gut, no processes, transverse chain. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. 111 Fig. T-13 SPECIES: Pegea socia ORDER: Salpida CLASS: Thaliacea SIZE: s to 140 mm, a to 120 mm DESCRIPTION: s: plump body with yellow band of pigment along each side, stolon coils twice around gut. a: body cylindrical, uniform gold pigmentation, no processes, transverse chain. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-14 SPECIES: Salpa fusiformis ORDER: Salpida CLASS: Thaliacea SIZE: s to 55 mm, a to 52 mm DESCRIPTION: s: smooth symmetric body, 9 body muscles, small, round, reddish gut. a: fusiform body with long anterior, posterior projections, 6 body muscles, linear chain. LUMINESCENCE: unknown DISTRIBUTION: world-wide and common Fig. T-15 SPECIES: Salpa maxima ORDER: Salpida CLASS: Thaliacea SIZE: s to 180 mm, a to 100 mm DESCRIPTION: s: smooth body, thick test, 9 body muscles parallel on dorsal side, large round, red gut. a: cylindrica with short anterior, posterior projections, 6 body muscles, round gut, linear chain LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. 112 Fig. T-16 SPECIES: Thalia democratica ORDER: Salpida CLASS: Thaliacea SIZE: s to 15 mm, a to 18 mm DESCRIPTION: s: 6 body muscles, 2 long posterior projections, shorter projections , around gut, round, blue or brown gut. a: ovoid body, 5 body muscles, posterior projection of gut, oblique chain. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-17 SPECIES: Thalia orientalis ORDER: Salpida CLASS: Thaliacea SIZE: s to 7 mm, a to 5 mm ~ DESCRIPTION: 6 body muscles, 2 very long posterior projections, 8 toothed ridges along test, no lateral projections. a: ovoid body, thick test, 5 body muscles, no gut projection, oblique chain. LUMINESCENCE: unknown ..".: -'. mk-.,., ""~"= :- - ': :.. ' , DISTRIBUTION:, Atlantic, Pacific, Indian, Med. Fig. T-18 SPECIES: Thetys vagina ORDER: Salpida CLASS: Thaliacea SIZE: s to 300 mm, a to 120 mm DESCRIPTION: s: 16-22 body muscles, body broad at mouth, tapered at posterior, with 2 lateral appendages. Test thick, greenish. a: cylindrical body, thick test of greenish hue. 5 body muscles, interrupted dorsally. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. 113 Fig. T-19 SPECIES: Appendicu/aria sicu/a F AMIL V: Fritilaridae CLASS: Larvacea SIZE: trunk 0.5 mm, entire 1.5 mm o DESCRIPTION: short, pear-shaped trunk, round mouth without lips, tail is broad, narrows near attachment to trunk. LUMINESCENCE: unknown fj i \: DISTRIBUTION: world-wide in warm or temperate water Fig. T-20 SPECIES: Fo/ia graci/is FAMILV: Oikopleuridae CLAss: Larvacea o E. SIZE: trunk 0.6 mm DESCRIPTION: ovoid trunk, flattened dorso- ventrally, narrow mouth with small ventral lip, tail pointed distally, lacks subchordal cells. LUMINESCENCE: Herring (1987) lists Oikopleura in this family as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-21 SPECIES: Fritilaria aequatoria/is FAMILV: Fritilaridae CLASS: Larvacea SIZE: trunk 0.7 mm, entire 1.0 mm DESCRIPTION: trunk long and narrow with enlarged pharynx, leaf-shaped tail with pointed end attaches at middle of trunk. \ LUMINESCENCE: unknown 0\1 DISTRIBUTION: Atlantic, Med. 114 Fig. T-22 SPECIES: Fritilaria borealis FAMILY: Fritillaridae Larvacea CLASS: SIZE: DESCRIPTION: pear-shaped trunk, mouth with rounded lip, tail rectangular, with central musculature and incised end. (I' .i. \' ! , ;1 " ì' LUMINESCENCE: unknown DISTRIBUTION: world-wide Fig. T-23 SPECIES: Fritilaria gracils FAMILY: Fritillaridae CLASS: Larvacea SIZE: trunk 0.7 mm, entire 2.7 mm o DESCRIPTION: trunk oval, broader at anterior .. end, mouth without lips, tail sharply narrowed at distal end. l- LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. T-24 SPECIES: Fritilaria haplostoma FAMILY: Fritillaridae CLASS: Larvacea SIZE: trunk 1.0 mm, entire 2.3 mm DESCRIPTION: long, narrow trunk, mouth with 1 large upper lip and 2 small lower lips, tail lanceolate, with scattered gland cells. LUMINESCENCE: unknown DISTRIBUTION: world-wide in warm water " 115 Fig. 1-25 Fritilaria megachile SPECIES: FAMILY: Fntilaridae CLAss: Larvacea SIZE: trunk 2.0 mm, entire 4.0 mm DESCRIPTION: trunk slim and elongate, not curved, mouth with large upper lip and 2 small lower lips, tail broadly rectangular with notched end, scattered gland cells. LUMINESCENCE: unknown DISTRIBUTION: world-wide in warm water Fig. 1-26 Fritilaria pellucida SPECIES: FAMILY: Fntillaridae '.t__ CLASS: Larvacea SIZE: trunk 1.5 mm, entire 3.0 mm DESCRIPTION: trunk elongate with enlarged A, anterior end, 2 conspicuous conical horns on posterior, tail broad with V notch in end. ....o.i LUMINESCENCE: unknown ~ø CD CD DISTRIBUTION: Atlantic, Pacific, Indian, Med. very common Fig. 1-27 SPECIES: Fritilaria venusta FAMILY: Fntilaridae CLASS: Larvacea SIZE: trunk 1.5 mm, entire 2.5 mm DESCRIPTION: trunk hourglass-shape from from above, flattened dorso-ventrally, with 2 large, flat horns posteriorly, mouth with large upper lip, tail lanceolate, notched. LUMINESCENCE: unknown DISTRIBUTION: world-wide 11 CD 116 Fig. T-28 SPECIES: Kowalevskia tenuis F AMIL V: Kowalevskiidae ..' CLASS: Larvacea SIZE: trunk 1.0 mm, entire 8.0 mm DESCRIPTION: trunk short, without endostyle, spiracles or heart, large rounded mouth, narrow, lanceolate tail, much longer than 'trunk. CD LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Antarctic, Med. Fig. 1-29 SPECIES: Megalocercus abyssorum FAMILV: Oikopleuridae CLASS: Larvacea SIZE: trunk 5 mm, entire 30 mm DESCRIPTION: ovoid trunk, with red-orange pigmentation, fairly small mouth with lower lip, tail broad, muscular with blunt end. LUMINESCENCE: Herring (1987) lists Oikopleura in this family as definite. o- - CD \. DISTRIBUTION: Atlantic, Pacific, Indian, Med. in deep water Fig. T-30 SPECIES: Oikopleura albicans FAMILV: Oikopleuridae CLASS: Larvacea SIZE: trunk 4 mm, entire 7 mm DESCRIPTION: trunk slender and elongate, conspicuous large white gonads in mature animals, tail slender and pointed, well developed muscles. LUMINESCENCE: Herring (1987) lists this genus as definite. House is also luminous (Galt, 1969). DISTRIBUTION: Atlantic, Pacific, Indian, Med. ~~.~~:.~:-~~-~, ~" - - -: " ,', "= ,',', ~:.., ,'- :.:-__ 117 Fig. 1..31 SPECIES: Oikopleura cophocerca FAMILV: Oikopleundae a CLASS: La rvace SIZE: trunk 0.7 mm, entire 2.6 mm Ii- ¡ !~ \ DESCRIPTION: trunk nearly rectangular, but tapered at antenor, fairly large mouth with prominent lower lip, tail muscular, with I \. .., .\ . tapered end. \\ 'I= ' LUMINESCENCE: Herring (1987) lists this "\ :i genus as definite. \ \ \'!i: DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. 1.32 SPECIES: Oikopleura dioica FAMIL v: Oikopleuridae CLAss: Larvacea SIZE: trunk 0.5 mm, entire 1.5 mm DESCRIPTION: small, globular trunk, separate sexes, terminal mouth with small lower lip, tail with narrow musculature and pointed tip. LUMINESCENCE: Herring (1987) lists this genus as definite. House is also luminous (Galt, 1969). DISTRIBUTION: world-wide except Antarctic Fig. 1.33 SPECIES: Oikopleura fusiformis FAMILV: Oikopleuridae CLAss: Larvacea SIZE: trunk 0.5 mm, entire 3.0 mm DESCRIPTION: trunk elongate, ovoid, flat dorsal surface, mouth opens obliquely upwards, tail long and slim, without subchordal cells. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide except Antarctic ~)-~ ~~-- -_...-- ---. 118 Fig. T-34 SPECIES: Oikopleura inter media FAMILY: Oikopleuridae CLASS: Larvacea SIZE: trunk 1.5 mm, entire 5.0 mm DESCRIPTION: ovoid trunk, tapered anteriorly, with convex dorsal surface, mouth opens obliquely upwards, tail with broad musculature, rounded tip. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. Oikopleura longicauda SPECIES: FAMILY: Oikopleuridae CLASs: Larvacea SIZE: trunk 0.7 mm, entire 3.5 mm DESCRIPTION: short, ovoid trunk with charactenstic membranous hood over posterior dorsal part, tail with broad musculature, rounded tip. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide, the commonest warm water species. Fig. T-36 SPECIES: Oikopleura parva FAMILY: Oikopleuridae CLASS: Larvacea SIZE: trunk 0.5 mm, entire 3.0 mm DESCRIPTION: trunk slender, ovoid, mouth opens anteriorly, with small lower lip, tail with narrow musculature, 4 subchordal cells near tip. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide in midwater 119 Fig. 1-37 SPECIES: Oikopleura rufescens FAMILY: Oikopleuridae CLAss: Larvacea . SIZE: trunk 1.5 mm, entire 5.0 mm DESCRIPTION: trunk short and ovoid, with l.. a A strongly convex' dorsal side, terminal mouth with small lower lip, tail broad with narrow musculature and 1 large subchordal celL. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. common G.PI Q -~e Fig. 1-38 SPECIES: Stegosoma magnum FAMILY:' Oikopleuridae CLASS: Larvacea SIZE: trunk 3.0 mm, entire 10 mm DESCRIPTION: trunk elongate and laterally compressed, with arched anterior dorsal surface, small terminal mouth, tail long with narrow musculature, 8 subchordal cells. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. 120 121 Crustaceans Crustaceans, especially copepods, are almost invariably the most abundant and often the most diverse constituent of the zooplankton. Some of the copepods and ostracods, and most of the euphausiids and decapods are known to be luminescent. Some possess discrete photophores and others discharge luminous secretions. A complete systematic coverage of the crustacean zooplankton of the western Mediterranean is well beyond the scope of this summary. Therefore this is not a comprehensive listing of the Mediterranean fauna, but those species of amphipods, euphausiids, mysids, ostracods, cope pods and decapods reported in recent zooplankton studies or from submersible observations in the western Mediterranean are summanzed in Table 9. Of those 88 species, 45, including most that are thought to be luminescent, are described and illustrated here. Some reports cited in Table 9 do not identify cope pods or ostracods to species; in cases where the genus is luminescent, a common species within it is given here as an example. "M" = male, "F" = female. Because of the diversity and complexity of crustaceans, identification to species, especially of copepods, can be difficult, and require expert familiarity with morphology of the body and appendages, and the accompanying descnptive terminology. Descriptions here refer where possible to general body shape and other characters that can be seen in live animals under a dissecting microscope. Identification of some groups may require the assistance of a specialist. Classification, descriptions and ilustrations for amphipods are compiled from Bowman and Gruner (1973), Shoemaker (1945), Stephensen (1925), Pillai, (1966a,b) and Tregouboff and Rose (1957). Information on copepods is principally from Rose (1933) with additional matenal from Owre and Foyo (1967), Tanaka (1956a,b, 1957, 1961, 1963, 1964) and Tregouboff and Rose. Ostracod descnptions are from Tregouboff and Rose. Descriptions and illustration of euphausiids are from Brinton (1975), Boden et al. (1955), Mauchline (1971), Wiebe (1976) and Tregouboff and Rose. Data on decapods is compiled from Crosnier and Forest (1973), Stephensen (1923) and Rice (1967). Terminology basal plate - the first segment of a pereopod, enlarged into a flat plate cephalothorax - the fused head and thorax of a copepod chelate - having a claw in which the 6th segment closes over the 5th furca - paired distal appendages on the urosome of copepods geniculate - having a grasping articulation at the end of the antenna pereopods - the thoracic legs rostrum - anterior projection of the carapace, out in front of the head t t ,r 122 simple - legs without claws subchelate - having a claw in which the 7th segment closes over the 6th uropods - the paired appendages of the urosome or tail urosome - the tail section consisting of last abdominal appendage, uropeds and telsen 123 Fig. CR-1 SPECIES: Brachysce/us cruscu/um FAMILY: Lycaeidae SUBORDER: Hyperiidea SIZE: to 17 mm DESCRIPTION: slender body with rounded head, large eyes, antenna 1 short, antenna 2 absent in F, long in M, pereopods 1 & 2 subchelate with teeth on margin, usually associated with medusae. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. CR-2 SPECIES: Phronima at/antica FAMILY: Phronimidae SUBORDER: Hyperiidea SIZE: to 40 mm DESCRIPTION: slender body with subconical head, elongate and narrowed ventrally, eyes have dorsal and lateral sections, pereopod 5 long with large claw, others simple, F in barrels made from salps. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. Fig. CR-3 SPECIES: Phronima sedentaria FAMILY: Phronimidae SUBORDER: Hyperiidea SIZE: to 40 mm DESCRIPTION: body and head similar to P. atlantica, pereopods 4,6,7 nearly as long as 5, narrow claw on 5, F in barrels made from salps. LUMINESCENCE: unknown DISTRIBUTION: world-wide 124 Fig. CR-4 SPECIES: Phronimella e/ongata FAMILY: Phronimidae SUBORDER: Hyperiidea SIZE: to 15 mm DESCRIPTION: very slender body with long abdomen, very long and thin pereopods, pereopod 5 with simple claw and toothed edge, F in short, round barrels. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. CR-5 SPECIES: Phrosina semi/unata FAMILY: Phrosinidae SUBORDER: Hyperiidea SIZE: to 20 mm DESCRIPTION: compact body, large head with anterior "horns", pereopods 1 & 2 Q) J" subchelate, 5 & 6 very large and subchelate, with toothed margins, pereopod 7 reduced to basal plate, free-swimming. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. Fig. CR-6 SPECIES: P/atysce/us ovoides F AMIL v: Platyscelidae SUBORDER: Hyperiidea SIZE: to 20 mm DESCRIPTION: body almost globular, rolls into ball, plate-like pereopods 5 & 6 cover ventral side, pereopods 1 & 2 chelate, pereopod 7 reduced, associated with siphonophores. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. 125 Fig. CR-7 SPECIES: Pseudo/yeaea paehypoda FAMILY: Lycaeidae SUBORDER: Hyperiidea SIZE: to 7 mm DESCRIPTION: body moderately plump, large round head, pereopods slender, without chelae, antenna 2 long and folded in M, absent in F, associated with medusae. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. Fig. CR-8 SPECIES: Scina crassicornis FAMILY: Scinidae SUBORDER: Hyperiidea SIZE: to 21 mm DESCRIPTION: elongate body, flattened dorso-ventrally, small head and eyes, long pointed antenna 1, long slender pereopods, long pointed uropods, body orange or red. LUMINESCENCE: Herring (1987) lists this genus as definite DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. CR-9 SPECIES: Streetsia challengeri FAMILY: Oycephalidae SUBORDER: Hyperiidea SIZE: to 40 mm DESCRIPTION: slender body with long pointed head, covered by compound eye, pereopods 1 & 2 chelate and spiny, other pereopods slender and simple. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. 126 Fig. CR-10 SPECIES: Euphausia krohnii FAMILY: Euphausiidae ORDER: Euphausiacea SIZE: to 25 mm DESCRIPTION: medium size round eye, 2 pairs of lateral teeth on carapace, pereopods 1-6 similar, 7 & 8 reduced. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. Fig. CR-11 SPECIES: Meganyctiphanes norvegica FAMILY: Euphausiidae ORDER: Euphausiacea SIZE: to 40 mm DESCRIPTION: elongate body, rostrum ends behind round eyes, pereopods 1-7 similar, 8 reduced, 1 pair of lateral teeth on carapace. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: N. Atlantic, Med. Fig. CR-12 SPECIES: Nematoscelis megalops FAMILY: Euphausiidae ORDER: Euphausiacea SIZE: to 20 mm DESCRIPTION: eyes divided into upper and lower lobes, 2nd pereopod extremely elongate, slender with apical bristles, no teeth on carapace. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. \ 1\ " '\ Il l,; "1" I~. '-if 127 Fig. CR-13 SPECIES: Stylocheiron maximum FAMILY: Euphausiidae ~" ORDER: Euphausiacea .:- .. DESCRIPTION: carapace with sharp rostrum ."--~ ~._~..... (~t.~~~:" ~'- SIZE: to 30 mm extending to end of large, elongate eyes, robust thorax, with reduced 1 st, 2nd, but extremely long 3d pereopod with chela. ~' '~i" c ---::.. ~~~.¡t.,-i ,',;l- LUMINESCENCE: Herring (1987) lists this genus as defi nite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. mesopelagic Fig. CR-14 SPECIES: Thysanopoda aequalis FAMILY: Euphausiidae ORDER: Euphausiacea SIZE: to 20 mm DESCRIPTION: carapace with dorsal trough, rostrum does not reach end of small, round eyes, very long antennae, pereopods uniformly short. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. CR-15 SPECIES: Acartia clausi FAMILY: Acartiidae SUBCLASS: Copepoda SIZE: to 1.2 mm DESCRIPTION: no rostrum, abdomen about 1/3 length of cephalothorax, short hairs on edges of thoracic segments. LUMINESCENCE: unknown DISTRIBUTION: world-wide 4t ~. ~ Psci 128 Fig. CR-16 SPECIES: Calanus helgolandicus FAMIL V: Calanidae SUBCLASS: Copepoda SIZE: to 3 mm DESCRIPTION: long, narrow body, antenna 1 longer than body and tail, 5 spines on each caudal furca, margin of basal segment of 5th pereopod toothed. LUMINESCENCE: Herring (1987) lists two genera in this family as uncertain. DISTRIBUTION: world-wide Fig. CR-17 SPECIES: Centropages chierchiae FAMIL V: Centropagidae SUBCLASS: Copepoda SIZE: 1.8 mm DESCRIPTION: body with tapered anterior, projections on posterior corners of last thoracic segment, antenna 1 shorter than body, long spines on urosome. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. Fig. CR-18 SPECIES: Centropages kroyeri F AMIL V: Centropagidae SUBCLASS: Copepoda SIZE: 1.3 mm DESCRIPTION: body tapered anteriorly, poterior projections on last thoracic segment, pereopod 5 chelate, with strong spines. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. 129 Fig. CR-19 SPECIES: Centropages typicus FAMIL V: Centropagidae SUBCLASS: Copepoda SIZE: to 2.0 mm DESCRIPTION: symmetrical postenor points on last thoracic segment in M, asymmetric in F, antenna 1 longer than cephalothorax. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Med. Fig. CR-20 SPECIES: Clausocalanus arcuicornis F AMIL V: Pseudocalanidae SUBCLASS: Copepoda SIZE: to 1.2 mm DESCRIPTION: short body, tapered antenorly, abdomen with 4 segments in ,5 in , pereopod 5 long and straight in M, short . Ù.~Ol ~\. ~ '\ '~ and curved in F. LUMINESCENCE: Unknown DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. Fig. CR-21 SPECIES: Corycaeus typicus F AMIL V: Corycaeidae SUBCLASS: Copepoda SIZE: 1.6 mm DESCRIPTION: cyclopoid copepods, body rounded anteriorly, with 2 large eyes, last (3rd) thoracic segment with postenor points, abdomen of 1 segment, long urosome. LUMINESCENCE: Herring (1987) lists this genus as uncertain. DISTRIBUTION: Atlantic, Pacific, Indian, Red Sea, Med. ~ l~ ~I ~ ¡~ ii (t \~I A~. lIpId ~ , ~.~ -rt~ i/A l ( . tid( tt~. V,' 130 Fig. CR-22 SPECIES: Eucalanus elongatus I\IC cd ",," FAMILY: Eucalanidae SUBCLASS: Copepoda SIZE: to 8.2 mm DESCRIPTION: elongate body, tapered anterior, very long antenna 1 with many spines and fan at ends, urosome with 1 long and several short terminal spines. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Med. Fig. CR-23 SPECIES: Haloptilis acutifrons FAMILY: Augaptilidae SUBCLASS: Copepoda SIZE: to 3.2 mm DESCRIPTION: cephalothorax with sharp anterior projection, antenna 1 much longer than body. LUMINESCENCE: Hernng (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Med. Fig. CR-24 SPECIES: Lucicutia f1avicornis FAMILY: Lucicutidae SUBCLASS: Copepoda SizE: 1.7 mm DESCRIPTION: oval body, numerous spines on antenna 1, slender abdomen with long terminal spines in F. LUMINESCENCE: Hernng (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. 131 SPECIES: Oithona he/go/andica FAMILY: Oithonidae SUBCLASS: Copepoda SIZE: 0.7 mm DESCRIPTION: oval cephalothorax, tapered anteiorly and posteriorly, antenna 1 with long spines, conspicuous egg sacs on ~'¥ m n~ abdomen in F. LUMINESCENCE: Herring (1987) lists this genus as uncertain. ~ f M DISTRIBUTION: world-wide Fig. CR.26 SPECIES: Oncaea mediterranea FAMILY: Oncaeidae SUBCLASS: Copepoda SIZE: 1.3 mm DESCRIPTION: short, oval cephalothorax, 1 st abdomen segment much longer than all others, body orange-red. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: world-wide Fig. CR.27 SPECIES: Paraca/anus parvus FAMILY: Paracalanidae SUBCLASS: Copepoda SIZE: to 1.0 mm DESCRIPTION: short body, head rounded in lateral view, F with 3 free thoracic segments, 5 abdominal, M with 5 abdominal segments, strong antenna 1. LUMINESCENCE: unknown DISTRIBUTION: world-wide ~ 132 Fig. CR-28 SPECIES: Pleuromamma borealis F AMIL v: Metridiidae SUBCLASS: Copepoda SIZE: 2.25 mm DESCRIPTION: body with 4 thoracic segments, antenna 1 of F with hooks, pereopod 5 with 3 equal spines on each tip. w LUMINESCENCE: Herring (1987) lists this genus as definite DISTRIBUTION: Atlantic, Med. SPECIES: ,.",---~ ,// ~~" ~ Pleuromamma gracilis ;~ "'-'- SUBCLASS: Copepoda SIZE: 2.0 mm I)71: 'ìi..~~\ ':j ,ií!f prehensile on left side, short spines on DISTRIBUTION: Atlantic, Pacific, Indian, Med. '(,:! ~' 'r \j\t ~- '-'-t ~ , ;~, of 1 st thoracic segment, M antenna 1 genus as definite ~ /, .-~,~~,~ DESCRIPTION: dark brown spot on right side LUMINESCENCE: Herring (1987) lists this ,')'.. I." .. I / '-- FAMILV: Metridiidae ends of last articles of pereopod 5. Fig. CR-29 r~ t ,¡" /~".~'f . ~:T" ";."- \' ,~,;/ '\ \ Fig. CR-30 SPECIES: Rhinealanus nasutus F AMIL v: Eucalanidae SUBCLASS: Copepoda SIZE: to 5.0 mm DESCRIPTION: long body, triangular pointed head with concave sides, antenna 1 much longer than body, M pereopod 5 with c1aw- like segment. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med., often deep ~~' 133 Fig. CR-31 SPECIES: Sapphirina iris FAMILY: Sapphinnidae SUBCLASS: Copepoda SIZE: to 7.5 mm DESCRIPTION: body very flattened dorso- ventrally, iridescent, antennae very short, 2 closely-spaced frontal eyes, body elongate in F, ovoid in M. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. CR-32 SPECIES: Scolecithrix bradyi FAMILY: Scolecithncidae SUBCLASS: Copepoda SIZE: to 1.4 mm DESCRIPTION: short body, thoracic segments 4 & 5 nearly fused, antenna much shorter than body, pereopod reduced and asymmetnc. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Pacific, Indian, Med. Fig. CR-33 SPECIES: Temora longlcornis FAMILY: Temondae SUBCLASS: Copepoda SIZE: to 1.5 mm DESCRIPTION: short, oval body with mid- anterior eyespot, 4 thoracic segments, M antenna 1 geniculate on right, M pereopod 5 with clawlike end. LUMINESCENCE: unknown DISTRIBUTION: Atlantic, Indian, Med. Ps '" .~ 134 Fig. CR-34 SPECIES: Temora stylifera FAMILV: Temotidae SUBCLASS: Copepoda SIZE: to 1.9 mm DESCRIPTION: short, broad body with rounded head, prolonged back corners of 5th thoracic segment, M with geniculate antenna 1, grasping claw on pereopod 5. LUMINESCENCE: unknown j DISTRIBUTION: Atlantic, Indian, Med. Fig. CR-35 SPECIES: Conchoecia obtusata F AMIL v: Halocyptididae CLASS: Ostracoda SIZE: to 2.0 mm DESCRIPTION: valves with straight dorsal margin and nearly rectangular outline. LUMINESCENCE: Herting (1987) lists this genus as definite. DISTRIBUTION: Fig. CR-36 SPECIES: Cypridina castanea FAMILV: Cypridinidae CLASS: Ostracoda SIZE: to 7.0 mm DESCRIPTION: valves with strongly curved dorsal margin, nearly oval outline, antennae extend well beyond shell margin. LUMINESCENCE: Herting (1987) lists this genus as definite. DISTRIBUTION: 135 Fig. CR-37 Acanthephyra pelagica SPECIES: FAMILV: Oplophoridae ORDER: Decapoda SIZE: to 147 mm total length DESCRIPTION: orange-red color overall, toothed rostrum extends well forward of small eyes, all legs simple, 7-11 pairs of spines on telson. LUMINESCENCE: Herring (1987) lists this definite. genus as DISTRIBUTION: Atlantic, Pacific, Indian, Med. meso pelagic Fig. CR-38 SPECIES: Gennadas elegans FAMILV: Penaeidae ORDER: Decapoda SIZE: to 40 mm DESCRIPTION: body red with blue spots, very long first antennae, no rostral projection LUMINESCENCE: Herring (1987) lists this genus as uncertain DISTRIBUTION: Atlantic, Med. Fig. CR-39 SPECIES: Pasiphaea multidentata F AMIL v: Pasiphaeidae ORDER: Decapoda SIZE: to 100 mm DESCRIPTION: carapace shorter than abdomen, rostrum short, pereopods 4,5 elongate and chelate, telson forked. LUMINESCENCE: Herring (1987) lists one genus in this family as definite and one as uncertain DISTRIBUTION: Atlantic, Med. 136 Fig. CR-40 SPECIES: Pasiphaea sivado FAMILV: Pasiphaeidae ORDER: Decapoda ~J)1~r-~ SIZE: to 100 mm DESCRIPTION: like P. multidentata, but telson not forked, with 2 longer lateral and 6 shorter medial spines. LUMINESCENCE: Herring (1987) lists one genus in this family as definite and one as uncertain ~ ..", ~ ,, '... ..,'.:: ",' ~" .='... ~ . . , ~,".,..""_'.,"'.~,...,',..'n".~ DISTRIBUTION: Atlantic, Med. Fig. CR-41 SPECIES: Sergestes arcticus F AMIL v: Sergestidae ORDER: Decapoda SIZE: 50 mm DESCRIPTION: body half red, 3rd maxilliped subequal with 3rd pereopod, setae on uropod outer margins end in tooth, 1 st segment of antenna longer than 3rd. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Indian, Pacific, Med. Fig. CR-42 SPECIES: Sergestes robustus FAMILV: Sergestidae ORDER: Decapoda SIZE: to 94 mm total length DESCRIPTION: body red allover, photophores without lenses on uropods and antennal scale only LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Pacific, Indian, Med. 137 Fig. CR-43 SPECIES: Sergestes sargassi FAMILY: Sergestidae ORDER: Decapoda SIZE: 30 mm DESCRIPTION: body half-red, 3rd maxillped longer than 3rd pereopod, its distal segment divided into 5 parts with irregular spines. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. SPECIES: Sergestes vigilax FAMILY: Sergestidae ORDER: Decapoda SIZE: 30 mm DESCRIPTION: body half red, 3rd maxiliped longer than 3rd peropod, its distal segment divided in 4 parts, rostrum blunt except apical spine. LUMINESCENCE: Herring (1987) lists this genus as definite. DISTRIBUTION: Atlantic, Med. ~ ,. 138 139 Acknowledgments i must initially thank Dr. Edith Widder for commissioning this project in the first place, and express my hope here that it meets her needs. Thanks also to N. Copley, R. Harbison, M. Omori and P. Wiebe for the loan of source material, and especially to K. Madin for invaluable and timely assistal1ce in all stages of the project. Preparation of this report was supported by a subcontract from the Harbor Branch Oceanographic Institution to the Woods Hole Oceanographic Institution, with principal contract support to E. Widder at Harbor Branch (Grant No. NOOO 14-91C6007) from the Naval Oceanic and Atmospheric Research Laboratory. References Allain, C. 1960. Topographie dynamique et courants gene raux dans Ie bassin occidentale de la Mediterranee. Rev. Trav. Inst. Peche Marit. 24:121-145. Alvariño, A 1957. Estudio del zooplancton del Mediterraneo occidental (Campana del "Xauen") in el verano de 1954. Bol. Inst. Esp. Oceanog. 81 :1-26. Alvariño, A. 1971. Siphonophores of the Pacific, with a review of the world distribution. BulL. Scripps Inst. Oceanogr. 16:1-432. Benovic, A. and A. Bender 1987. Seasonal distribution of medusae in the Adriatic Sea. In: Modern Trends in the Systematics, Ecology, and Evolution of Hydroids and Hydromedusae. J. 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Bioi' 15:139-149. DOCUMENT LIBRAY March 11. 1991 Distrbution Listfor Technical Report Exchange Attn: Stella Sanchez-Wade Documents Section Scripps Institution of Oceanography Library, Mail Code C-075C Pell Marine Science Library University of Rhode Island Hancock Library of Biology & Texas A&M University Dartmouth, NS, B2Y 4A2, CANADA University of Washington Seattle, WA 98195 Library La Jolla, CA 92093 Narragansett Bay Campus Narragansett, RI 02882 Working Collecton Oceanography Alan Hancock Laboratory University of Southern California University Park Los Angeles, CA 90089-0371 Gifts & Exchanges Library Bedford Institute of Oceanography P.O. Box 1006 Dept. of Oceanography College Station, TX 77843 Library Virginia Institute of Marine Science Gloucester Point, VA 23062 Fisheries-Oceanography Library Office of the International Ice Patrol c/o Coast Guard R & D Center R.S.M.A.S. Avery Point Groton, CT 06340 NOAA/EDIS Miami Library Center 4301 Rickenbacker Causeway Miami, FL 33149 Library Skidaway Institute of Oceanography P.O. Box 13687 Savannah, GA 31416 Institute of Geophysics University of Hawaii Library Room 252 2525 Correa Road Honolulu, HI 96822 Marine Resources Information Center 151 Oceanography Teaching Bldg. University of Miami 4600 Rickenbacker Causeway Miami, FL 33149 Maury Oceanographic Library Naval Oceanographic Office Stennis Space Center NSTL, MS 39522-5001 Marine Sciences Collection Mayaguez Campus Library University of Puerto Rico Mayagues, Puerto Rico 00708 Library Institute of Oceanographic Sciences Deacon Laboratory Wormley, Godalming Surrey GU8 5UB UNITED KINGDOM Building E38-320 MIT Cambridge, MA 02139 Library The Librarian CSIRO Marine Laboratories G.P.O. Box 1538 Colombia University Palisades, NY 10964 Library Proudman Oceanographic Laboratory Library Serials Department Oregon State University Merseyside L43 7 RA Lamont-Dohert Geological Observatory Corvalls, OR 97331-5503 Hobart, Tasmania AUSTRALIA 7001 Bidston Observatory Birkenhead UNITED KINGDOM Mac 90-32 50272-101 PAGE WHOI-91-26 REPORT DOCUMENTATION 11. REPORT NO. 2. 4, Title and Subtitle Distribution and Taxonomy of Zooplankton in the Alboran Sea and Adjacent Western Mediterranean - A Literature Survey and Field Guide. 3. Recipient's Accession No. 5, Repor Date SePtember, 1991 6. 7. Author(s) Laurnce P. Madin 8. Performing Organization Repl No. 9. Performing Organiztion Name and Address 10. ProJectaskMork Un" No. WHOI-91-26 I I Woo Hole Ocogrphic Institution Woo Hole, Masachusett 02543 11. Contract(C) or Grant(G) No. (C) NOOI4-91-C607 (G) 13. Type of Repo & Period Covered 12. Sponsoring Organization Name and Address Technica Report Naval Ocogrhic and Atmospheric Resch Laboratory 14. 15. Supplementary Notes This report should be cited as: Woos Hole Oceaog. lost Tech. Rept, WHOI-91-26. 16. Abstract (Llm": 200 words) Th is a surey of literature rerd for ocurence and taonomy of zooplan in the Western Medterrea, with parcula emphais on the Albora Sea It is intended to give a genera background on the fauna, and facilitate identication of spimens collecte or observed. A description of the hydrgrphy of the Albora Sea is followed by a genera account of zoplan biomass ditrbution, and more detaed lists of the ocurence of 361 spies of medusae, siphonophores, ctenophores, worms, tucate and crustaceas in 7 regions of the Western Medterrea. Bioluminescent propertes of the orgamsms are indicate where known. An ilustrated taonomic guide provides capsule descptions and ilustrtions of 254 of the listed spies. 17. Document Analysis L Descriptors zoplanon Albora Sea biolwnescence b. Idantiflerslpen-Ended Terms c. COSATI Field/Group 18. Availabilty Statement Approved for public relea; ditrbution unlimited. See ANSI-Z39.18) 19. Secur"y Class (Tis Report) 21. No. of Pages 20. Security Class (This Page) 22. Price 150 See Instructions on Reverse OPTIONAL FORM 272 (4-77 (Fonerly NTIS-35) Department of Commerce ,:€rr , " l., :f