sc1. MAR.. 63 (3-4): 343-354 SCIENTIAMARINA 1999
BIOLOGY AND FlSHERY OF LWLPHlNFlSH AND RELATED SPECIES. E. MASSUTI~~II~B. MORA LES-NIN (rds.)
Parasitism of dolphinfishes, Coryphaena hippurus and Coryphaena equiselis, in the western Mediterranean (Balearie Islands) and central-eastern Atlantic (Canary Islandsj*
ENRIQUE CARBONELL', ENRIC MASSUTÍ~,JOSÉ JUAN CASTRO3 and ROSA MARÍA GARCÍA' 'Depanamento de Biología. Universidad de Valencia, Dr. Moliner, 59.46100 Burjassoi. Valencia (Spain) '1.E.O.- Centre Oceanografic de les Balears. Moll de Ponent s/n, P.O.Box 291.07080 Palma de Mallorca (Spain) 'Depanamenio de Biología, Universidad de Las Palmas de Gran Canarias. P.O. Box 550. 35 1 17 Las Palmas de Gran Canarias (Spain)
SUMMARY: A total of 648 doiphinfishes were exarnined for interna1 and exiernal parasties in western Mediterrane;in (Balearic Islands) and central-eastern Atlantic (Canary Islands) waters in order to iiiake a comparative study beiween thc two areas. The specirnens studied from the Mediterranean Sea was Coqphnrt~crIiip1n1vrr.i. wiih 62 large individuals coptured from May to Sepiember and 497 juveniles captured from August to December. The specimens studied from the cenrral-easi- ern Arlaniic were 39 adulr C. liippurrts and 49 adult Coryphaena equise1i.r. Parasites were found in 70%- of ihe fish exani- ined. and represented a ioial of riine endoparasitic taxa: six digeneans (Class Tremaioda. Subclass Digenea: Diiirrrrr.c ro171o- /irs. Dinrrrus hr
K<,\ wrds. Co~yhuc~irrIripprrus. Cor~phu INTRODUCTION water ternperarures exceed 20°C (Gibbs 2nd Col- lette, 1959). In the Atlantic, they are natural inhab- The dolphinfishes (Pisces: Coryphaenidae), itants and their spawning is probably year round. cornrnon dolphinfish (Coryphaena hlpp~rl-u.rL.) although it is rnost intense during the inonths with and pompano dolphin (Corvphaena equiseliz L.); hieher water surface ternperaturec (Shcherhachw. are epipelagic species distributed world-wide in 1973). In the Mediterranean Sea, C. hippur-rr.7 tropical and subtropical waters where surface appear seasonally and undergo a reproductive cycle during the surnrner rnonths, t'ollowing [he 'Receivcd June 15. 1998 Accepted May 8. 1999 appearance 0f adult specimens every yeilr ¡n May- June, when the water surface temperature reaches gastrointesrinal parasites. Both studies werr carried >18"C (Massuli and ,Morales-IYin, 199511. Juve- out along [he South-Eastem and Gulf coasts of the niles are captured from Iate August to early United States in rhe Western Atlantic. December and then, when water iemperatures fall Therefore the objective of this study was to carry [o lI°C, they possibly rriigra~e to thc: warnxr out. for the first time. a complete analysis of the par- waters of the Atlantic Ocean. asiric community of dolphinfishes in the eastem Dolphinfishes are top-leve1 prdators, but rhey A4tlanticand the Mediterranean, and to study thz reia- arz not very selective and fced on a wide rangc of iionshipj between diet and parasite recruitment in pelagic organisms. In adaition, they are ve-' agile order to identify possible pathways of parasitic infec- and capable of rakin; fasi-moving prey (Palko e! al., tion and life cycle. Special referente is rnade to the 1982j. C.hippurus grows rapidly throughout its life paasites which C. hippurus probably bnngs from he and has a maximum life span of about 4 yrars, Atlantic to the Mediterranean Sea and those which reaching lengths and weights in excess of onc mme this species acquires in the Meditsmnean anu ches and 10 kg respectively (Beardsle!, 1967; Rosz and to Atlantic waters. Our aim is to provide basic infor- Hassler, 1968). C. equiseiis is a relative11 Iirrle maiion that will allow parasites to be used as porer:- known species which does not reach such a large tia1 biolopical tags for C. hippurus in rhe study of its size as C. hippurus, and its maximum known length migrato? routes between these two areas. is 74 cm (Herald, 1961). C. equiseiis is more pelag- ic and oceanic, and consequmtiy is rarely caught in coastal waters. Its distribution ranpe is more tropical MATERIALS AND METHODS and according LO Mather and Dtij (1954) it is not generaliy found in waters with surface water tem- The fishes were collected fiorn two areas, Cne peratures Iower than 24°C. C. eyuiselis does nor Balearic Islands jwestern Mediteii-anean) and the extend as fu beyond the rropics as C. hippurus. Canary Islands (central-eastem Atlanticj. In the lab- There are few reports of C. equiselis in the Mediter- oratory, the fish were measured ro thr nearest cm- ranean Sea. and hence its presence in these waters is tirrierre fork length (FL), weighed and sexed. Intc~u- no[ well known. ment, fins, natural openings and gilIs of every ficn Although severa1 authors have reponed parasites were exarnined. The gills were dissected in order to oi doiphinfishrs in every ocean. and án exhaus~ive survey al1 [he gil1 arches for parasites. In aII cases, iis; icin be found in Palko er al. (1982), only r few the parasites were collected and, in addition to their focus on rhe study of the parasire community of number, size, shape and location, any pathological #L--,. L.. ----:A--.-- A-., .L LIICX +JCL;G~ U? LUII~~UCIIIJ~ih~CLUIU~~ UI LLI~par- a:iZiaiiOliS .&sere recoided. A-,.-uir~c aiij pui---e.-:,-- U~~LGJ asites. Burnett-Herkes (1 974.1analysed [he ectopar- were found. they were fixtd in buffered glureralde- üsites on the gills and in the buccal iiavity cf C. hlp- hyde ( 10%) for larer identification, directly or aftzr pilrus, whereas Manooch e! al. (1984) studied its being cleared with lactophenol. T.4s.i l. - Nunber of specirnens o( C i:ii7purus and C. equk'is zapiurec! ir. Mrciicr:ariean and Atlaniic waicrs. bv kí1 Icngrh in~erva!~. )'ea; ;nd mx!h oi capture m3 th: surfase water tempcrature are also ;km. From the Mediterranean area, a total of 62 adult tode, two nernatodes and one acanthocephalan), of C. hippurus, ranging from 60 to 124 cm FL, were which nine were to species level. The change in sampled during May-September in 1990, 199 1 and prevalence, intensity and abundance of the main ir%. in iiie sañie years, 497 juveniies íi4-58 cm species in reiation to the size of the hosis is preseni- FL) were obtained from August to early December. ed in Table 2 and 3. From the Atlantic area, 25 adult (69- 102 cm FL) and 24 juvenile (38-53 cm FL) specimens of C. hippurus Class TREMATODA were sampled during June and October in 1994 and Subclass Digenea 1995 respectively. In this area, 49 C. equiselis Family Hemiuridae between 36 and 52 cm FL were also sampled during Dinurus spp. June in 1995. To establish differences in parasitism as a function The most frequent and most numerous endopar- of size, the fish exarnined were differentiated into six asites were four species of the genus Dinurus Looss, size intervals in order to obtain a sufficient number of '1907. These were D. tornatus (Rud. 18 19) Looss, sFciiíieiis pigicrUp. Sizr. iniervais, n"m'mr "f fjsh i9 i7, D. barbaius (Cuiin, i902j Looss, i907, D. analysed per size group, year and month of capture breviductus Looss, 1907 and D. longisinus Looss, and the surface water ternperature range are sum- 1907. D. tornatus was the dominant species. These rnarised in Table 1. For each parasite, the infection parasites were found in the stomach of specimens of leve1 by size group was analysed according to stan- C. hippurus and C. equiselis captured in both areas. dard methods (Margolis et al., 1982). For the prevalence and other ecological parameters there were marked differences depending on the annual cycles, but in general adult fish were the RESULTS most infected in both areas. Endoparasites Lecithocladium excisum (Rudolphi, 18 19) Looss. 1907 Endoparasites were found in 390 of the speci- mens exarnined (70%). A total of eleven parasitic This other gastric hemiurid was only found in the taxa could be identified (seven digeneans, one ces- Mediterranean, from juvenile fish 40cm FL in the TABLE2. - Infeclion parameters of the helminth species (Dinurus spp.. LPcirllocladiunl exciston. Floriceps succarus. Rhadinorliychus pris- n ris and Merabrorie~nn magna) of Conphcrenu h~pprrruscaptured in Mediierranean waters. (FL. fork length in cm. firsi row. prevalence; E second row: mean intensity t SD; third row: mean abundance f SD; fourth row: range). a ,, n Dinurus spp. L. excisum F. saccarus R prisris M. magna o FL 1990 1991 i 995 1991 1990 1991 1995 1995 1995 3 3.6 2.8 3.6 3 2 I O. 1 +0.6 0.0620.3 0.03f0.2 O-? O-? O-: PARASITISM OF DOLPHINFISHES 345 CLASSN ELI .\TODA Order Spirurida Farnilq Cystidicolidae Metubronema (Qistidicoloides) magna (Taylor, 1925) Some cysts were collected from thr walls of the pyloric caeca and in the pancreatic rissue of C. hp- purus specimens captured in the Mediterranean. They contained several adult nematodes belongin; .A*. i YY i cycie. its prevaience, intensity anci abunuance to Chis species. The cysts measured 30-50 mm in were low. length and 20-30 mrn in width. and they were poor- 1y defined, solid, opaque and included in the iissues. Farnily Bathycotylidae Most of the cysts were calcified. The infection Bathycotyle brarzchialis Darr. 1 902 appeared in small fish, and the prevaience increased rapidlt, rernaining high at aII fish sizes, ahhough ir Collecred fron~gills of three large specirnens of was on1: possible to follow the chariges o? [he C. ni.ppurus captured in Mediterranean waters. iniection diiring the 1995 cycle. In rhz youn;esr fish, the cysts constituted a soiid mass of conjunc- Family Hiruciinellidae tive tissur which coniainrd a rnale aiid various Hirudinella sp. females encrusted irrzgulariy in the rnass. in the larser !ish, the calcification ufas constan1 and com- Coliected from the stomach of two Iarge speci- plete, with most of the nematodes being broken and mens of C. hipp~tr~iscaptured in [he Mediterranean fragmented. Class CESTODA Family Dracunculidae Grder Try panorhy ncha Philornerroides sp. Farnily Lacystorhynchidae Floriceps saccatus Cuvier, 16 17 Pleroczrci were fmnd in spe,-inizns of C. +i,?,v:¿- rus larser than 3V cn FL captured in both xeas. A ~oraiof 9i qsrs conrining 2 singie picrsczrioic were collected from the ci~domina!cavity anl tl-i? Phylium ACANTHCCEPHALA surfare srrosa of th? vissera. C~sisize \+as cririablz Po1 y morphida and ranged frorn 20 to 50 rnm. The shape oí thi Fsrni iy Rhadinorhynchidae cp~saiso showed a greut cririeiy of i'orrns, but they Rhadirzorlzynchus pristis (Rudo1pr.1,1802; always had a rerrninrtl globuix reg'ion, where [he Liihe, 191 i :xva occurred, anci a tail-like srruciure. The-smallesr cysts 1:4-7 rnm in Iength) showed an enlarged nr hoad-like scoIex with t\vo rounded exirnsions. Tne rnedium sized larvae measured 10- 12 rnm and ha3 :entades and tivi? leaflike bothridia. The larges: pie- roceícoiifs (15-25 in ieligti:; showcd iyqo deeply TABLE4. - lnfection parameters of the copepod species (Cdigus spp.. Pennellufilosa and Neohracliiella coryplioenidae) of Cotyliaena Iiip- pirrus captured in Mediterranean waters (FL: fork length in cm: first row: prevalence: second row: mean intensity ISD; third row: mean abundance k SD; fourth row: range). encapsulated in the caecal walls or deeply in pan- Table 5. - lnfestation paraeters of the copepod speciesuryphorus nynpltof Coryphaena hippurus and Con.phaenu equiselis captured creatic tissue. This species was also found in C. in Atlantic waters (FL: fork length in cm; first row: prevalence; sec- equiselis specimens smaller than 40 cm ond row: mean intensity ISD; third row: mean abundance f SD; FL. founh row: range). Ectoparasites Seven species were found, al1 of which were crustacean copepods. Their infection parameters are given in Table 4 and 5. Class COPEPODA Order Shiponostomatoida. Family Caligidae Caligus spp. Collected in the gills of C. hippurus captured in higher water surface temperature. Parasite females !he Medi?errme~r!2nd thre~ghe~!a!! !he sizr inter- WPK EKXX ahmdan! thar! ma!es, u/i!h !he sex-rctb vals considered. Caligus quadratus Shiino, 1957 being 7: 1. was the dominant species. Other species of this genus, such as C. coryphaenae Steenstrup and Family Euryphoridae S n/ r Lütken, i 86 i, C. bonito Wiison, i 905 ana C. pro- Curypizorus nymphae Sreenstrup ana Lüiken, I ao i ducrus Müller, 1785 were also found. They were located in large masses of mucus surrounding the Collected from both species in specimens bigger gills (60%) and on the inner surface of the opercu- than 40 cm FL captured in Atlantic waters. The pro- lum (40%). C. hippurus and C. equiselis from the portion between parasite females and males was 2: 1. Atlantic area were not infected by these copepods. This species, as well as Caligus spp., produced an The prevalence in juvenile fish was higher than in abundan1 mucus hypersecretion in the gills, and the adults. It was also higher during the months with parasites were found in this mucus mass. They were also present on [he inner surface of the operculum. body of smali size (45~-14rnm in lengtti; ranpe 22- Though the examined fish were dead, the parrtsites í3 mmj. The rest were composed of pregravid aná uere still highly active and they could be found in gravid females with three wrli-developed horns, an íiie ora; cavicy, skin on rne head and oiher skin ioca- inienseiy keriiiinizeá boáy anli a dark brown coiour. tions. The sizt: has 41-13 mm (59115 nim). The gilis consisted ~7frhe main fiiarnenr irxn wnicr! prima-, Family Lemaeopodidar secondary and even tenia? hranches arose, hut thzy .Veobrachiella coryphaenidae Pearse, 1 952 were always withoui anastamosis. Collected from C. hippurus juvenile and adult specimens captured in the Mediienanean. The infec- DISCUSSION tion took place in juvenile fish during [he months with high water temperarures, and in every case par- The results obtained allowed the little existing asices were attached to the gil1 filaments by rheir infomation concernin; the parasite fauna of dol- appendages. 0~1~.a few parasites carried dwad phinfishes in the Mediterranean and Centrai-Eastem males. Mucus hypersecretion or any other iinaromi- .4rlantic to be extended. None of the species found cal alterat~onwere not observed. in the hlediterranean had been reported in this area until now. The Mediterranean records were limited Family Pennellidae to Dollfus (1927), who reported a metacercaria of Pennellu filosa Linnaeus, 1 758 the hemiurid trematode Dinurus notarus. Lozano- Cabo ( 196 1 ) who found the isopod Anilocro Collected only from large specimens oi C. hip- ph~sodes,and Delamare-Deboutteviiie and Nunes- purus captured in Mediterranean waters during Riiivo (195s) who reported the two parasitic gil1 May-Seprember in 1990, 1991 and 1995. Young fish copepods Brachielia LO-phaenae and Caligus caughr in this sea, and the ones from the Canaq befone. By contrast, al1 the species found in the Islands, were not infected by this parasite. Although Canary Islands had been reported ir1 Atiantic ivaters, the ecological paramerers were different during the although most of these records correspond to the three years studied, in evep case they showed an western toas[ (e.g Cable and Linderoth, 1963; Ho, i~~~ta~iFLotenri~m~\~ fiSh h~~a~~~eOjdc~-The !363). The on!i avui!ah!e inf~rmxi~r!from the cm- Q L"'U"'-: parasites ufere mainly attached to the dorsal and anal tral-western Atlantic were studiec on trematodes fins (50%j. to dorsolareral muscular tissue (25%) frorri fishes off Ghana and Senegal (Fischthal, 1972; and to the abdominal cavity (2W¿). Mised parasite Fischthal and Thomas, 1 W2b). locations were frequent in fish larger [han 100 cm FL (40%). The depth at which the cephalothorax or Endoparasi tism radicular apparatus \vas anchored varird according to parasite location. Thus, it was deeper in the The presence o! gastric herniurid digenean para- abdorninai cavity and rnuscularure [han in fin rays C!i- sirrs in juvconik and adult specimens of C. hippurru subcutaneous tissur. Some parasites ivere hyperpar- suggesis tnat infection takes place ;n Mediterranean asitized by tne cirriped Conckxierm~:au,ritut,;. Skin 2s ii~l!ss in Ailanri: iv3cers. The iife ¿yci,~of ihrae . . *.,A ,..,,.. 1,- ,,,,,. :r. ...e,, ,l...,.,.- rrlirli,, ..m- tL.3 nllu iiiu~~uiuiii~~iu>i> WGIL aiwuya picscii., L::~L iii~ rfiJspa;as;:ej 1s re:;iijt.ij.v ci,ííipii:<, i:v,pi:al of many head and horns oi he parasice were immtrsed ir) ii i-:elmintn parasites. with set,eral iniermediate hoats. bioody and infiammatory esudate mass. The Ir may foiiow the typical modd drscribed by Koie cephalothorax was surrounded b) a fibrous conjunc- (1 579, i940b, 1590c), who reponed benthic gas- tive tissue capsuie and the adjacenr tissues showed rropods as first intermediate hosrs where [he ~*rcar~--O different ratec of inflan~maroryalterations wirlh iis- a:: deveiop, whereas meiasercariae occur in [he tulizations. haemocoe! oi copepo2s and rhaeto_gnarns. This 30% of the materia! iourid corresponded to author also sho?ifed ihat sume ciecaood ianlae may premetamorphii forms that lacked cephaiii homs be infected by eating the cercariae and that thz and showed simple branrhial filaments, wirhcw! de5nitive hosts wcre clupzids, scomhrids and oiher ramifications or ar most with prirnary branchcs rhai plankton-feeding Mes. .As al\ these 0rpa.i.isrns are always arose frorn the main branch. Thev uere present in the Me~iterranean2nd .4ílzr,tic waiers, ir premetamorphic females with a fragiie, whitish ap?ears thai C. izi,-.;.urits couid dcqiiire rne parasites either through the ingestion of copepods and other copepod crustaceans. Metacercariae have also been crustaceans with metacercariae or through eating described in jellyfish, in the ctenophores P~ELI~O- small infected fish, which act as second intermediate brachia, in diverse species of Sagitta and in the hosts (Gibson and Bray, 1986). These authors seem coelorn of the polychaete Totnopteris. These non- to conclude that non clupeid fish could acquire the crustacean invertebrates are certainly infected when hemiurids by feeding upon clupeids or other plank- eating copepods with metacercariae, and should ton-feeders. The infection of metacercariae herni- therefore be considered as paratenic hosts. In our urid digeneans from clupeid fish and crustaceans has case( the infection by this trematode is early, which been recorded in C. hippurus by severa1 authors suggests that it took place when C. hippurus juve- (Dollfuss, 1927; Yamaguti, 197 1 ; Manooch et al., niles preyed on pelagic crustaceans, although the 1984). possible infection route by paratenic hosts such as Sirnilarly to other areas of the Pacific and Sagitta and other invertebrate components of the Atlantic, fish. crustaceans and cephalopods have pelagic system cannot be ruled out. been reponed as main components of the diet of C. The other two digeneans Barhycoryle branchialis lzippurus in the Mediterranean (Massutí et al., and Hirudinella sp. only appeared in a few cases in 1998). Nevertheless, an important change takes Mediterranean waters, and can be considered rare as place during the ontogenic development of the secondary parasitic species. B. branchialis had been species. Whereas amphipod and crustacean larvae previously reported in the gills of large C. hippurus make up more than 50% of the diet of juvenile spec- specimens frorn the Straits of Florida (Burnett- imens smaller than 30 cm, cephalopods and mainly Herkes, 1974), although it is not proved whether this fish are the most important preys for bigger fish. parasite infects the gills or is vomited from the stom- Taking into account that the Dinurus species ach, as we have found in Dinurus spp. Two species appeared in juveniles (>40 cm FL) and adult speci- of Bathycotyle have been described, B. branchialis mens (>60 cm FL), [he transrnission mechanism in scornbrids and B. conphaenae in C. hippurus. from invertebrates does not seem probable and their The only data to differentiate them is the presence of infection is likely to be produced frorn ingestion of an external opening of the Laurer's canal in the first clupeid or other plankton-feeder fishes such as species (Gibson and Bray. 1979). Although it is not carangids, which are an irnportant component of the clear whether the Laurer's canal has an external C. hippurus diet. opening or not, we have seen a dorsal opening in our The only data thai we possess on the biological specimens and in consequence we have considered cycle of the species of Dinurus are due to Dollfus thern as B. branchiaiis. The life history of this . . (! 927) 2nd Szidx ( !950). The f;,rs! autho: mggests spec:es :S m! kmvín. The gp,i,u ,~irudf?t,~!!cis 8 that cenain metacercariae found in the body cavity typical stornach parasite of rnarine teleosts (mainly of the crustacean decapod Cerataspis monstrosa scombrids and tunas), although coryphaenids are belong to Dinurus notarus, a stornach parasite of C. habitual hosts (Yamaguti. 197 1 ). Due to their great liippurus, for which two fornis of transmission are morphological variability, many studies have quoted as being possible: through invertebrates described up to fourteen different species of this (copepods, decapods and chaetognaths) or through genus, although Yarnaguti ( 197 1 ) stated that a wide- vertical migratory fish (clupeids: carangids and ranging revision is required. The life cycle of this scombrids). Szidat (1950). on the other hand, con- endoparasite is also unknown. siders that the clupeids are intermediate hosts for Fforiceps saccatus plerocerci have been recorded Dinurus breviductus. whose rnetacercariae encyst or in teleosts, whereas adult cestodes are gut parasites -C 1 -1 L -,.-,. l.,. r--- .L- A +l*-+:.-. -..A ertcapsüiaie iñ the skiñ O¡' Snrdinü piichordu~and VI >CVCI~I CI~~IIIUUIPIILII~I IUIII LIIC MLIL ! ,, Ywo-host model for Rhodlnorhynchus pristfi 1 ;+C,MOBWC~~~ L - Regarding the ectopxasites (Fig. 2). PPUP?PI!C ate hasts. The rrmainin; copepod parasites nave -fiiosa is a copepod that needs two hosts, a fish o: a copepodite and pelagic chalimus phaszs which iorm cetacrirn as a definitive hcist and cephalopod moi- part oi the planktonic community, here re !he ii?urrn iuscc as intermediate tiosts. These last ones, are chalirnus stagc difierzntiates into ~readut~miles anO infec~edby copepodite larvae thar develop four c hai- females thar ar? rhe infxtive iormc ior [he silis ei : a.n"..- :# *t.,. -.Al¡ .%..A *L.,. l,.c, -4- ~\,L:*L..,-A A:; &.c. .-.A,-.,nb.fl,-..;A.- IIIIU~-.,.aiasc3 iii LIIL iiiuiiu.~,iii~ iaJL vi niiici. ulr. uii- LiiL cUiJ piLdLIliU3. ferentiatec sexiidly arid are fenilised. Tne males The presrnr siudy shows that [he Iife qcle arid rhen disa~pearand the knilised females :nfm C. brna\iour- of C. Aippurus 2nd C. eauiseíi.; Are of ,i:+purus where they unaergo extensive metamor- ;real imponanx in Irte rzxaitrnen:, 3evzis?mr,t, phosis. This explains ths great imporrarice of ihe and transrnlssiar. 3i parasites in the Aririnti< Ocean physicai contacts betweeri definitive ara in~ermeai- and in [he Mediterranean Sea. Ir1 borh species, the recruitment of parasites is qualitatively similar to that of the large fish whose diet is based almost exclusively on teleost fish. Therefore, the feeding habits, surface water temperature, and length and body weight of fishes are related to parasite recruit- ments (Fig. 3). This study shows that there are severa1 parasites of Cj. hhippurus that are potentially useful as biologi- cal tags for studying its migratory movements and stock differentiation within the Mediterranean, and between this area and adjacent Atlantic waters. Within the endoparasites, the trypanorhynch species PLnhrruc CIULWS( Ch-l ro CM) F: saccatus, widely recognised as a long-lived para- site, and the acanthocephalan R. pristis, despite its low prevalence, probably have the greatest potential as biological tags. These species have a sufficient life span and remain in an identifiable form in C. hippurus long enough to cover the time scale of the investigation. For the same reason, the usually short P(uihmc NAUPUUS life spans (<1 year) of adult digeneans in the ali- mentary tract of fish, tends to limit the use of these species as bioiogicai tags. Of t'ne ectoparasite cope- pods found, /? jlosa has a particular advantage as a tag because it is a large, very easily seen ectopara- site that leaves a prominent scar after its death, Fic. 2. - Proposed life cycles for the ectoparasites. Left: Cycle for Pennellajiloso. Right: Caligid cycle for Caligus spp and Eurypho- thereby extending its usefulness as a tag beyond its rus nyrnphae. Lemeopodid cycle for Neobrachiella coryphaenae. actual life span. REFERENCES Krioata. Z - !Yo?. Copr!~nd; p:rrii.r,:!:. oi<,'i.\i~r.\ SynapG of Eriiisn Saum. 47. Linnran Swiriy. London. Koir. M. - 1979. Or. [he morphoiogy ano iik-rii~r?.of Lk-ropne.~ Aridrrhon, R.C. - 1992. bi~'e~iiu~odrP~ir,iciit.:; of i ertehrures. Fsrn- vcirirus ('vlüllsr. 1784) Loas. 190 1 (Trrniaioda: Hrrniuridaei. ham Koyal Buckh CAB lnrrrnarional. Wallingford. 578 pp. Spi!scnrifrfirr Purriri~e~rhu~ide.59: 6?-65. Beardsley. G.L. - 1967. Age. grourh and rcproduciion of [he dol- Koie hl. - 1990b. Redejc:ipiior. o:' [he cercar¡;: o- 2;-i!iiochir;unt phin Co~pkxníinrppurus trom [he slraiis of Florida. Ccpeiiii. rufiviride (Rudsiphi. LS 19) Luhe. 130 i (Lh-nea, Herniuri- 2: 441451. dar). Olnheiic. ? 1 : 85-95. Benz. G.W iind W.E. Hogans. - !993. Pet~e/lo]i/n~a (L. 1758) Koie. hl. - 1990~.On :he morpiioiogy and iiie-nistory of He~riiccrti.s (Copepoda: Stphonoston.iatoidai ironi thz escolar Lcpido-Ci- Iizehei Oahnsr, 1905 !Digene¿: Wrrniuridse~.J. Heirnini.. 61: urn j7nvobrunnrum (Srnith. 1849) in [he nortii-wesr .4tiantii. 193-202. Sp! Pnrasi:.. S6 (3): ! 27- i 3 l. K&, hl. - !991. Af,pec:.s ui thz wiorphorogy 2nd iiie cycle of Biaci, G..k and N.W. Lankesier. - 1930. Miprarlon and develop- kL-iri:<,c.lntiiirit;ii~~r tV.rc.irun:(Dipea: Hcmiuridae). L parasite 01' rnenr oi swin-~bladder nemarode Cwtidicoirr sp. (hletabronz- Scornber spp. 1r:r. J. Porirsi:.. 2 l(5;. 592403. rnaridaej in ihrir definiiive hos:. Cciri. J Um.. 58: I997-?005. Lozano-Cabo. F. - 1061. Biorneiris, bioiogia y prscd de la lampii- Burnerr-Herkes, J. - 1973. Parasiirs o!' the giiis 2nd buccril cavity of ga (Coq)iiiien[i h:ppuru.sj ds las lsiix Baleares. A~miernic~u'e [he dolphin. Cortl>/icirrrr; hr/?r>urrt.v.lroni ;he Straiis of Florida. Cie~rkisErtrcitis. Fi'iSiciu y .V~~luruie.riic .&í:r