© SociedadEspañola deMalacología Iberus, 19 (2): 87-95, 2001 Vexing question on fisheries research: the study ofcephalo- pods and their parásitas Un asunto embarazoso en investigación pesquera: el estudio de los cefalópodos y sus parásitos Santiago PASCUAL*^ andÁngel GUERRA** Recibidoel10-111-2001.Aceptadoel7-IX-2001 ABSTRACT For the beginning of this century it is evident that support for research will be increasingly dependent upon the results ofthot research having relevance to society's needs and public benefits. Within this web and coupled with the transfer of scientific knowledge is the opportunity to more effectively explain the society the benefits it receives for its investments in research. This viewpoint paper enlighten the general public on the scientific, industrial and commercial relevance of the research on cephalopods and their parasites. In a com- parative analysis with other commercially-important taxa, a historical negligible financial support for research on diseases in this important animal group was noted. Because of that policy-makers on Fisheries Research should balance this public debt in the future. RESUMEN En el comienzo de este siglo es evidente que el apoyo a la investigación se incrementará en función de la relevancia que para las necesidades de la sociedad y de los beneficios públicos se obtenga de los resultados de dicha investigación. En este contexto y paralela- mente a la transferencia de conocimientos científicos, surge la oportunidad de explicar más eficazmente a la sociedad los beneficios que recibe de su inversión en investigación. El punto de vista de este artículo ilustra al público en general sobre la relevancia cientí- fica, industrial y comercial de la investigación de los cefalópodos y sus parásitos, así como de la histórica e insignificante financiación destinada a la investigación de las enfermedades en este importante grupo animal. Un análisis comparativo con otros taxo- nes comercialmente importantes, sugiere que los gestores de la política de Investigación Pesquera deberían equilibrar esta deuda pública en el futuro. KEYWORDS: cephalopod, parasite,fisheriesresearch. PALABRAS CLAVE: cefalópodo,parásito, investigaciónpesquera. INTRODUCTION Cephalopods are fast-growing car- ecosystems (Clarke, 1996). Moreover, nivorous molluscs that play an impor- cephalopod stocks are of great interna- tant role in the trophic webs of marine tional importance in commercial fishe- * Área de Parasitología, Grupo PB2, Facultad de Ciencias del Mar. Universidad deVigo, Apdo. 874, 36200 Vigo. Spain.e-mail: [email protected] **InstitutodeInvestigacionesMarinas (C.S.I.C.). EduardoCabello6, 36208Vigo. Spain. 'Correspondingauthor 87 Iberus, 19 (2), 2001 Bivalves Cephalopods % 36.95 17.95% (1.77) Í0.86) Seacrustaceans % 5.85 (0.28) Marinefishi % 39.25 (1.88) Figure 1. Percentagevalúes andratios (in parentheses, ¥10-6) whichrepresenttherelativescientific effortbynominal catch unit foreachgroup ofspecies. Ratios arecalculated as the total numberof publications that address parasitology and associated-pathology (mean for the period 1992-98: source ofinformation in Biological Abstracts) and the nominal catch per group ofspecies (mean numberfortheperiod 1992-98: sourceofinformationinF.A.O., 2000). Figura 1. Porcentajesy ratios (enparéntesis ¥10-6) que representan elesfuerzo científico relativopor unidaddecaptura nominalparacadagrupo deespecies. Losratiossehan calculado como número total depublicacionessobreparasitologíaypatologíaasociada (mediadelperiodo 1992-98;fuente:Biological Abstracts)y la captura nominalpara cadagrupo de especies (mediapara elperiodo 1992-98;fuente: F.A.O., 2000). ríes (BoYLE, 1990; Guerra, 1992; Josu- how do they find food, escape from PEIT, 1995). World catch statistics record enemies, migrate, signal to one another atotal catchof3.5millionmetrictonsin and reproduce, how does theirnervous 1999 (JosuPEiT, 2000), with a rate ofin- system function and what is their fis- crease significantly greaterthanthatfor hery potential in terms of biomass. By finfish species for the period 1970-1992 comparison, in the last decade marine (BoYLE, 1990; Caddy, 1995). For the scientists have devoted very little time combined marine catch, all categories, and money to the study ofcephalopod between 1970-1992 there was anincrea- parasites and parasite-induced patho- se rate of 2%per year. In contrast, for logy in wild and cultivated popula- cephalopods, the rate of increase over tions. the same period averaged 8% per year. Because oftheirimportance as ahuman food resource (Pierce and Guerra, SCIENTIFICPRODUCTIVITY 1994) and because cephalopods have proved to be valuable as experimental Among 700,000 scientific papers a animáis forbiomedical andbehavioural year published around the world (see research (Gilbert, Adelman and Ar- Scientific Citation Index), alm.ost NOLD, 1990; Abbot, Williamson and nothing is written on cephalopod disea- Maddick, 1995; Hanlon and Messen- ses (Fig. 1). Based on current trends GER, 1996), scientists have spent consi- there also appears to be an overall derable time and money studying cep- decline in the number of recent papers halopods in the last two decades: how that treat taxonomy and basic research many species are there, how long do on parasites of other, non-commercial they live, how fast do they grow, how marine invertebrates. This may be due do they respond to changes in fishing in part to the loss of parasite workers intensity andenvironmental conditions. (specially taxonomists) through lack of Pascualand Guerra: Study ofcephalopods and theirparasites on fisheries research funding support or redirection of rese- how few researchers there are in the arch efforts to the study of human pat- field. hogens or ecological studies. Neverthe- less, despite the obvious scarcity of THE SPANISHPOLICY scientific effort, recent world hterature has stressed the important role of In this context, we may look at the squids, cuttlefishes and octopuses as Spanish situation which is comparable reservoirs for all taxa ofmarine eukario- with other fish-catching countries in tic parasites at both macro- and micro- Europe. Spain is the fifth largest cepha- geographic sampling levéis (Hochberg, lopod consuming country in the world, 1990; Pascual, Gestal, Estevez, Rodrí- the first in Europe. In Spain cephalo- guez, Soto, Abollo and Arias, 1996). pods represent animportant component Most large free-living, mature cephalo- of the diet (4 Kg./inhabitant/per year), pods carry some microscopic (viruses, yet no financing has been directed at bacteria, fungi, protists) and macrosco- research on cephalopod parasitic disea- pic parasites (metazoans) in almost all ses. Although over 4 million US $ per their tissues and organs (Hochberg, year is assigned by Plan Nacional de 1990). This author, in an extensive Investigación y Desarrollo (Comisión survey ofthe Hterature, record a total of Interministerialde CienciayTecnología) 225 parasite species in cephalopods to Research and Development (hereafter world-wide. Since the Hanlon and RandD) to finance research activities FoRSYTHE (1990) review on diseases and projects on Marine Science and caused by microorganisms and the Technology, nothingwas assigned to the Hochberg (1990) contribution on disea- investigation of cephalopod diseases in ses causedbyprotistans and metazoans, the last decade. In this context, itis rele- only 32 scientificpapershave dealtwith vant that cephalopod landings in Spain cephalopod pathology. Ofthese, only 16 (averaging 110,000 metric tons per year papers have been published in interna- for that period) contributed 308,000,000 tional scientificjournals, 4ofthesebeing US $ per year to our domestic economy. chapter reviews within a zoological It seems quite remarkable that at least a and/or fisheries biology context. small portion of the money derived Despite the fact that cephalopod lan- from the fishery and manufacture pro- dings have increased over the period cesses profits should be returned to 1992-1998, the number of people invol- support research on diseases of wild ved in this field and the number of and cultured cephalopods. The absence papers on cephalopod pathology is still of support for a technology transfer remarkably low. In fact, the scientific mechanismclearlyindicatesthattheres- effortdealingwithcephalopod parasites ponsibilities of the Spanish research and associated pathologyrelatedtorela- fundingagenciesdoesnotcloselyfollow tive nominal catch by group of species current fishing trends to support both account less than 54% of that on parasi- established and emerging activities as tic diseases of other commercially- the management of the ecological important invertebrates (bivalve impact ofparasitism inwildlife and cul- molluscs) and marine fish (Fig. 1). Why tured populations are (Grenfell and isthis so? Gulland, 1995). In other words, this Diversity of hosts versus number of indicates that in this field the Science- researchers is a key factor. Thus, there Technology-Industry Spanish System are approximately 700 species of cepha- (STISS) still has an imbalance between lopods and veryfewresearchers compa- the scientific andproductive spheres.As red with about 100,000 species of fishes regards STISS (involving parasitolo- and numerous researchers. Neverthe- gists), it appears to be particularly sti- less, the scientific productivity with mulated by the production of scientific regard to papers on cephalopod parasi- publications and thus, the most impor- tes actually is not too bad considering tant yardstick to promote researches is 89 Iberus, 19 (2), 2001 the number of publications within the diseases in the 1990's. Such a brief Scientific Citation Index (SCI). STISS synopsisisurgentlyrequiredinthelight lacks, however, of sufficient support to of the many advances which have been employ young trained scientists in rese- madeutilizingnewtechniques. arch activities and seems to be insuffi- Although older reports of infectious ciently motivated by the food technolo- diseases emphasized description and gical and sanitary aspects that imply a systematic classification of cephalopod suitable development of the research on parasites, considerable confusión exists. parasitic-caused cephalopod patholo- The identifications of the parasites and gies. In the case of Food technology, sometimes even the hosts are often in STISS have an important drawback doubt, with high synonymy rates (cióse which is the low level of interest of to 70%) for numerous parasitic nominal many prívate companies to deal with species identified by light microscopy the necessary innovation of its products (Pascual, Arias and Guerra, 1995; or processes. Consequently financial Mattiucci, Nascetti, Cianchi, Paggi, and human resources being assigned by Arduino, Margolis, Brattey, Webb, public policy and the prívate sector to Dámelio, OrecchiaandBullini, 1997). develop research activities aimed at a Researchers trained in modern techni- scientific assessment of the impact of ques, new trends in systematic, and infectious processes in cephalopod improved technologies for detecting stocks are patently insufficient. This and defining species have allowed us to situation is made worse when public elucídate and re-evaluate the taxonomic organizations or businessmen have to status and the host-parasite relations- solveseriousproblemsrelatedtothetre- hips of many already described species. atment and control ofcephalopod disea- For example, recent papers dealingwith ses at present day in industrial proces- scanning and transmission electrón ses. And the situation will be still worse microscopy (SEM and TEM) and atomic if, following the successful results in forcé microscopy (AFM) studies have experimental rearing of planktonic showed how much a combination of common octopus from hatching to set- increased depth of field, resolution and tlement (Villanueva, 1995) and on magnification is needed in the Identifi- growing of this species in floating cages cation and examination of the morpho- (Hebberecht, 1996; Moral-Rama, 1996; logy microtopography, topometry and Guerra, unp. data), the industrial cellbiology ofcephalopod parasites and culture of the octopuses become a busi- the host-parasite interface (Gestal, ness that rent good profits. In this Pascual, CorralandAzevedo, 1999). regard, a recent study by the Industrial Additionally, our understanding of Research and Development Advisory theepizootiology (whichinvolvesinves- Committee of the European Commis- tigations on the demographic infection sion (1994) warns ofthe obvious danger valúes, patterns of transmission, and ofeconomic stagnation, unless there is a disease control) of many parasitic greater coordination between the pro- species in cephalopods is severely ham- ductive system (extraction or produc- pered by morphological characters of tion, processing and marketing of the difficult interpretation The existence of resource) and the much-needed scienti- morphologically identical cryptic fic environment (comprising the RandD species and parasitic races or morphoty- groups). pes which can reflect selection pressure rather than taxonomic affiliation are problems faced by all taxonomists, but FUTURE RESEARCHDIRECTIONS present particular difficulties because of theplasticityofbody structures inendo- In this article we should not forget parasites. Moreover, when histological, some of the general trends emerging isolation and purification processes and from the study of cephalopod parasitic parasitic dissection techniques are all 90 Pascualand Guerra: Studyofcephalopods and their parasites on fisheries research needed to reveal diagnostic characters since it agrees with demographic para- of some protozoan and metazoan ecto- meters observed duringspecial cephalo- parasites, respectiyely, a high degree of pod disease surveys in the same área skill (and training) is required. A (Pascual, 1996). number of molecular techniques should Although an extensive literature be developed to overeóme these pro- dealing with diseases and defence blems and shouldbeappliedworldwide mechanisms is available for other com- as useful taxonomic tools for parasite mercially-important moUuscs, little detection andtheir speciesIdentification emphasis has been placed on the incephalopods. defence mechanisms of cephalopods. Diseases and pathology caused by Despite humoral and cellular defence microparasites on wild and cultured associated responses having been des- cephalopods have been reportad in a cribed for cephalopods maintained in few cases (Hanlon and Forsythe, closed sea-water systems forbiomedical 1990; PoYNTON, Reimschuessel and studies or fattening against potential Stoskoff, 1992;Gestal, 2000). However, bacterial pathogens (Hanlon and it should be noted that in Spain, during Forsythe, 1990; Ford, 1992), the effects the massive culture of paralarvae and of other microscopic and macroscopic juveniles insystem crowding, highmor- parasites on phagocyte capabilities, talities rates have been assigned to inflammation, wound healing and func- several environmental factors including tional morphology of cephalopod hae- diseases by bioagressors (Gestal, mocytes has not been investigated in Abollo and Pascual, 1998). Further- depth. The study of inmunobiology of more, despite cephalopod macroparasi- cephalopods is just starting (Malham, bteisonttyspi(caHlOlCyHhBaEvReG,be1e9n90c)o,nhsiisdteorpaadthsoylmo-- B1E9S9,6;1M99a7)l.hAambe,tDteurnuhnadmerastnadndSiencgomo-f gical analysis on heavily parasitized host defence reactions in cephalopods cephalopods revealed the destruction of would also help to avoid or control out- vital organs and potential loss of their breaks of parasitic diseases in commer- functionality (Pascual, 1996; Abollo, cial mariculture conditions where Gestal, López, González, Guerraand animal densities, intensive husbandry Pascual, 1998). Unfortunately, in the and stress may increase occurrence of past although attentionhasbeenpaid to parasitic disease. Because cephalopods thepresenceofparasiticdiseasesinwild are afood source inmany regions ofthe cephalopods, most of the early studies world, the effects of parasitic infections can be classified as single or short-time on the biochemical composition and observations. Seasonal and continuous physiological characteristics including long-term parasite studies are missing, condition, growth rate, nutrient assimi- resulting in the current absence of relia- lation and protein/energy ratio of cep- ble data to be used in comparative halopods in nature and culture systems analysis. These data will improve our shouldalsobeevaluated. knowledge about whether present Most wildlife parasitic diseases have disease prevalence in wild exploited been investigated via pathological post- cephalopod stocks exceed natural pre- mortem examinations, or by producing valence, change with abiotic parameters lists of parasites identified in small and/or are influenced by host exploita- samples of hosts. There have been few tion rates and discarding practices. attempts to assess the impact of a These studies will be also very useful disease at the population rather than for mapping the existence of hot-spot individual level, or to describe the dis- áreasbyusing the grid systems ofInter- tribution of the disease agent in a national Fishery Organisations. To manner sufficient to understand its epi- obtain base-line data, cephalopod demiology. Pascual, Gestal and disease recording in standard stock- Abollo (1997) considered the statistical assessment surveys is potentially useful distribution of parasites throughout the 91 Iberus, 19 (2), 2001 host species population, and confirmed but when large-scale culture in high thenegative effectofgilí macroparasites density groups was initiated, fatal infec- on the condition of exploited ommas- tious diseases occurred (Hanlon, trephid stocks. That study clearly sug- Forsythe, Cooper, Dinuzzo, Folse gests the existence of causal relations- ANDKelly, 1984; Gestaletal., 1998). In hips, expressed in negative modifica- Spain, where cephalopod mariculture is tions of ecological potential, between changing from experimental to indus- parasiticinfection and cephalopod stock trialsphere, implementationofprocedu- productivity. Finally, aneconomiclossis res for detection and monitoring the present (Pascual, González and pathology and parasitic diseases on a Guerra, 1998). Although further bio- widescaleintheongrowingcephalopod chemical data on infected and parasite- industry should be common, together free cephalopod tissues should be recor- with other yet established diseases ded to ascertain the physiological inte- assessment policy (e.g. on cultured ractionsbetweencephalopods and para- fishes andbivalvemoUuscs). sitic infections, parasites may have a Furthermore, parasitoses in cephalo- considerable effect on infected stocks or pods appears not only as an important individuáis, as has been recently noted problem in the management of infected by Gestal (2000). Less obvious to most stocks, but also a zoonotic problem fisheries scientists is the important role during food-processing. Larval stages of of parasites in regulating the general macroparasites are found in many "well-beingor fitness" ofthehostpopu- species of squids, cutlefishes and octo- lation (i.e. in regulating host abundance puses, which are of commercial impor- or fecundity). To this end, we should tance. The appearance of parasites attempt to blend mathematical models makes cephalopods unsightly and for host-parasite relationships with unappealing to consumers. Moreover, those used by fisheries biologists to although several species of anisakid determine how parasites can affect the nematodes and trypanorhynchcestodes, dynamics of exploited cephalopod at the larval stage, canbe pathogenic if populations, following the seminal arti- consumed in raw or improperly cooked cles of Crofton (1971), Anderson and cephalopod dishes, few cases of illness May (1978, 1979), Sindermann (1987), byhelminths inmanhavebeenreported DoBSON AND May (1987) and Grenfell inSpain (Abolloetal., 1998). andGulland (1995). Thecomplications Cephalopod ecologists may also introduced by the presence of parasitic benefit by applying new ideas such as disease will in general further increase the study of the host-parasite systems. the levéis of uncertainty that cephalo- Pascualand Hochberg (1996) revised pod fisheries managers have to contend the use of parasites as non-intrusive with (Beddington, 1984), this mostly in natural tags of cephalopod hosts in fis- relation to cephalopod condition and its heries science. Piotozoan and metazoan potential fecundity (i.e., its recruitment parasites have been used to assess the dynamics). status of current stocks of several com- Roughly 10% of the known species mercially-exploited cephalopod species. ofliving cephalopods (i.e. over 80 ofthe Few examples of the trophic status of 700 known species) have been either cephalopodswithinfood webs and their maintained, reared or cultured in capti- parasitecommunitystructureareavaila- vity (BOLETZSKY AND Hanlon, 1983). ble from the literature. The quantifica- The vast majority of these 82 species tion of genetic variation obtained from (representing 30 genera) have mainly allozyme frequencies among or within been maintained or reared, while 12 populations of larval anisakid nemato- species (7 genera) have been cultured des (i.e., the parasite most frequently through their entire life cycle (Hanlon, employed as tags for marine organisms 1987). No signs of diseases were ever and the most commonly encountered observed during small-scale production. macroparasite in cephalopods) can 92 Pascualand Guerra: Studyofcephalopods and theirparasites on fisheries research provide valuable data on trophic rela- fits they receive for their investments in tionships and stock identity of most research (Murrell, 1996). Bearing in wild cephalopod stocks. mind all the comments above, we feel Through workshops and symposia that few cephalopod parasitologists CIAC (Cephalopod International Advi- have succeeded well in communicating sory Council) aims to present current many aspects of scientific and technical research and to stimulate and promote knowledge, but they have been less future research. Among the almost 40 effective in enlightening the general symposia and workshops on cephalo- public and prívate partnerships on the pods held by fisheries biologists from societal valué and economic relevance 1973, the 1996 workshop on cephalopod of their research contributions. To over- parasites developed on behalf of Com- eóme this it is imperative to encourage mission Internationale pour l'Explora- public and prívate managers, fisheries tionScientifique dela MerMéditerranée scientists and parasitologists to contact (CIESM) by Boletzsky and Hochberg each other to go one step further in the (Laboratory Arago, Banuyls-sur-mer, 2ithcentury. France) was the first attempt to provide on the parasite diseases processes of wild and reared cephalopods. The con- ACKNOWLEDGEMENTS tents ofthis workshop established itself as the main introductory handbook of We thank Comisión Interministerial working techniques on cephalopod de Ciencia y Tecnología (CICYT) for parasitology. their financial support (CICYTMAR95- Today is well-established that 1919-C05). Thanks also are extended to support for research is increasingly an anonymous review and all members dependent upon the results of that rese- of the research groups University of arch having relevance to society's needs Vigo-PB2 and Instituto de Investigacio- and public benefits. Coupled with the nes Marinas (ECOBIOMAR-CSIC) for transfer of knowledge is the opportu- aid and numerous helpful suggestions nity to more effectively explain to fishe- for a decade of fruitful joint collabora- ries managers and companies the bene- tion. BIBLIOGRAPHY Abollo,E.,Gestal,C,López,A.,González, Boletzsky,S.v.andHanlon,R.T.,1983.Are- A. F., Guerra, A and Pascual, S., 1998. viewofthelaboratorymaintenance,rearing Squids as trophicbridges forparasite flow andcultureofcephalopodmolluscs.Memoirs withinmarine ecosystems: the case ofAni- ofthe National Museum ofVictoria, 44: 147- sakissimplexorwhenthewrongwaycanbe 187. right.SouthAfricanJournalofMarineScience, Boyle, P.R., 1990. Cephalopod biology inthe 20:223-232. fisheriescontext.FisheriesResearch,8:303-321. Abbot,N.J.,Willlamson,L.andMaddick,L., Caddy,J. F., 1995. Cephalopod anddemersal 1995. Cephalopod neurobiology. Neuroscience finfishstocks:somestatisticaltrendsandbio- studiesinsqiiid,octopusandciittlefish.Oxford logicalinteractions. Squid'94 VeniceInterna- UniversityPress,Oxford,542pp. tionalCephalopodTradeConferenceProceedings. Anderson, R. M. and May, R. M., 1978. Re- AgráEuropeLondonLtd,25pp. gulation and stability of host-parasite po- Clarke,M.R.,1996.Roleofcephalopodsinthe pulationinteractions.I.Regulationprocesses. world'soceans. Philosophical Transactions of JournalofAnimalEcology,47:219-247. theRoyalSocietyofLondonB,351 (1343):977- Anderson, R. M. andMay, R. M., 1979. Po- 1112. pulationbiologyofinfectiousdiseases:Part Crofton, M. D., 1971. Amodelofhost-para- I.Nature,280:361-367. siterelationships.Parasitology,63:343-364. Beddington,J.R.,1984.Managementxinderim- DoBSON, A. P. andMay, R. M., 1987. Theef- certainty.InMay,R.M.(Ed.):Exploitationof fectsofparasitesonfishpopulations-theo- MarineCommunities.DahlemWorkshop32, reticalaspects.InternationalJournalforPara- SpringerVerlag,Berlín,pp227-244. sitology, 17:363-370. 93 Iberus, 19 (2), 2001 F.A.O.,2000.Yearbook-FisheryStatistics-Cap- IRDAC(IndustrialResearchandDevelopment tureproduction.Vol. 86/1. Food andAgri- AdvisoryCommitteeoftheEuropeanCom- cultureOrganisationoftheUnitedNations, mission),1994.QualityandRelevance:thecha- Rome2000:713pp. llengertoEuropeanEducation.UnlockingEu- Ford,L.A.,1992.Hostdefencemechanismsof rope'sHumanPotential.E.C.March:156pp. cephalopods.AnnualRevieivofFishDiseases, Josupeit,H.,1995.Worldsupplyandmarkets. pp.25-41. In:Squid94Venice.The3rdInternationalCep- Gestal,C,2000.Epidemiologíaypatologíadelas halopodTradeConference.Agra-Europe(Lon- coccidiosis en cefalópodos. Ph.D. Thesis. Uni- don).Ltd: 13pp. versidaddeVigo,Vigo.España. Josupeit,H.,2000.Losmercadosmundialesde Gestal,C,Abollo,E.andPascual,S.,1998. cefalópodos.ProductosdelMar,Noviembre- Rickettsiales-likeorganismsinthegillsofre- Diciembre:43-48. ared Octopiis vulgaris (Mollusca, Cephalo- Malham,S.K., 1996.InmunobiologyofEledone poda). Bulletin oftheEuropeanAssociation of cirrhosa(Lamarck)Ph.D.Thesis,University FishPathologists, 18(1): 13-14. ofWales,Bangor,U.K. Gestal,C.,Pascual,S.,Corral,L.andAze- Malham,S.K.,Dunham,N.W.,Secombes,C. VEDO,C.,1999.Ultrastructuralaspectsofthe J. 1997. Phagocytosis by haemocytes from sporogony ofAggregataoctopiana (Apicom- the lesser octopus Eledone cirrhosa. Iberus, plexa,Aggregatidae),acoccidianparasiteof 15(2):1-11. Octopus vulgaris (Mollusca, Cephalopoda) Mattiucci,S.,Nascetti,G.,Clanchi,R.,Paggi, fromNE Atlanticcoast. EuropeanJournal of L., Arduino, P., Margolis, L, Brattey, J., Protistology,35:417-425. Webb,S.,D'Amelio,S.,Orecchl\,P.andBu- GiLBERT,D.L.,Adelman,W.J.andArnold, LLINI,L.,1997.Geneticandecologicaldataon J. M., 1990. Squid as Experimental Animáis. theAnisakissimplexcomplex,withevidencefor PlenumPress, NewYorkandLondon, 516 anewspecies(Nematoda,Ascaridoidea,Ani- pp. sakidae).JournalofFarasitology,83(3):401-416. Grenfell,B.T.andGulland,F.M.D.,1995. Moral-Rama, A., 1996. Estudios bioqin'micose Introduction:ecologicalimpactofparasitism histológicos de cefalópodos relacionados con la on wildlife host populations. Farasitology, aplicación detecnologíasconvencionalesynue- 111 (Suppl),S3-S14. vasyconelcontroldecalidad.Informefinaldel Guerra, A., 1992. Cephalopod resources of proyectoTS3CT93-0109. the world: a present day view. Squid'91 Murrell, K., 1996. Communications: techno- MadridInternatioiialCephalopodTradeConfe- logytransferinthedevelopedworld. Vete- rence Proceedings. Agrá Europe London rinaryFarasitology,64: 107-120. Ltd,1-15. Pascual,S.,1996.Lossistemashospedador-parási- Hanlon,R.T., 1987.Mariculture.InBoyle,P. toenlapesqueríadeommastréfidosdeGalicia.Ph.D. R. (Ed.): Cephalopod Ufe cycles. Vol. 11. Aca- Thesis,UniversidaddeVigo,Vigo,España. demicPressLondon,291-305. Pascual, S. and Hochberg, F. G., 1996. Ma- Hanlon,R.T.andForsythe,J.W., 1990.Di- rineparasites asbiologicaltagsofcephalo- seases ofMollusca: Cephalopoda. Diseases podhosts.FarasitologyToday,12(8):324-327. causedbymicroorganisms.InKinne,O(Ed.): Pascual,S.,Arias,C.andGuerra,A., 1995. DiseasesofMarineAnimáis.BiologischesAns- ElectrophoreticIdentificationofL3larvaeof taltHelgoland,Hamburg.Vol.lU,23-46. Anisakis simplex (Ascaridida: Anisakidae), HanlonR.T.andMessenger,J.B.,1996.Cep- parasitesofsquidsinNEAtlantic. Research halopod Behaviour. Cambridge University andReviewsinFarasitology,55(4):239-241. Press,232pp. Pascual,S.,Gestal,C.andAbollo,E.,1997. Hanlon, R. T., Forsythe, J. W., Cooper, K. EffectofPennellasp.(Copepoda,Pennellidae) M.,DiNuzzo,A.R.,FoLSE,D.S.and Kelly, on the conditionofIllex coindetii and Toda- M.T., 1984.Fatalpenetratingskinulcersin ropsiseblanae(Cephalopoda,Ommastrephi- laboratory reared octopuses. Journal ofIn- dae). Bulletin ofthe European Association of vertébratePathology,44:67-83. FishPathologists, 17(3/4):91-95.. Hebberecht,C,1996.Experienciadecultivoin- Pascual,S.,González,A.F.andGuerra,A. tensivodepulpo(Octopusvulgaris)ensufasede 1998.Effectofparasitismontheproductivity engorde en artefacto flotante. Memoria. Con- oftheommastrephidstocksinGalicianwa- selleríadePesca, MarisqueoeAcuicultura. ters (NW Spain): economic loss. Iberus, 16 Xunta de Galicia. Santiago de Compostela. (2):95-98. Spain,5pp. Pascual, S., Gestal, C, Estévez, J., Rodrí- Hochberg, F. G., 1990. Diseases ofMollusca: guez,H.,Soto,M., Abollo,E.andArias, Cephalopoda. Diseases caused by protistans C,1996.Parasitesincommercially-exploited andmetazoans. InKimie,O(Ed.): Diseasesof cephalopods (Mollusca, Cephalopoda) in MarineAnimáis.BiologischesAnstaltHelgo- Spain:anupdatedperspectíve.Aquaculture, land,Hamburg.Vol.IH,47-227. 142: 1-10. 94 Pascualand Guerra: Study ofcephalopods and their parasites on fisheries research PiERCE, G. AND Guerra, A., 1994. Stock as- Sindermann, C. ]., 1987. Effects of parasites sessmentmethodsusedforcephalopodfis- onfishpopulations:practicalconsiderations. heries.FisheriesResearch,21:255-285. InternationalJournalforParasitology, 17:371- POYNTON,S.L.,REIMSCHUESSEL, R. ANDStOS- 382. KOFF,M.K.,1992.Aggregatadobellin.sp.and Villanueva, R., 1995. Experimental rearing Aggregata milleroruin n. sp. (Apicomplexa: and growth of planktonic Octopus vulgaris Aggregatidae) fromtwo species ofOctopiis fromhatchingtosettlement. CanadianJour- (Mollusca: Octopodidae) from the Eastern nalofFisheriesandAquaticSciences,52:2639- NorthPacificOcean.]ournalofProtozoology, 2650. 39(1):248-256. 95