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A Contribution to the Knowledge on The ACONTRIBUTIONTO THEKNOWLEDGE ON THEMORPHOMETR Y ANDTHE ANA TOMICALCHARACTERS OF PENNELLABALAENOPTERAE (COPEPODA,SIPHONOSTOMA TOIDA,PENNELLIDAE), WITH SPECIAL REFERENCETO THEBUCCAL COMPLEX BY P. ABAUNZA1,4/,N.L.ARROYO 2/ andI. PRECIADO 3/ 1/ InstitutoEspañ ol deOceanografía (IEO),Apdo. 240, E-39080 Santander, Spain 2/ UniversidadComplutense de Madrid(U.C.M.), Facultad de Biologí a, Departamentode Biologí a AnimalI, E-28040Madrid, Spain 3/ Asociación Cientí cade Estudios Marinos (A.C.E.M.), Apdo. 1061, E-39080 Santander, Spain ABSTRACT Pennellabalaenopterae ,thelargest known copepod, still presents many questions regarding its morphologicalfeatures and its mode of life. T welvespecimens of P.balaenopterae werecollected from Balaenopteraphysalus inthe northeastern Atlantic Ocean. Their morphological characteristics havebeen analysed by scanningelectron microscopy and a detailedmorphometric study has been accomplished.The buccal complex is describedfor the rsttime, providing details on themouth, rst maxillae,second maxillae, and buccal stylets, together with a new,trilobedstructure not previously describedfor the genus. Other appendages are also presented, including the rstantenna, second antenna,swimming legs, and other external and internal structures. The results are discussed and comparedwith those found in theliterature, together offering a morecomplete picture of the anatomy ofthisspecies. RÉSUMÉ Pennellabalaenopterae ,leplus grand copé pode connu, pose encore des questions quant à ses caractères morphologiques et son mode de vie. Douze spé cimens de P.balaenopterae ont été ré- coltés sur Balaenopteraphysalus dansl’ océan Atlantique nord-oriental. Leurs caracté ristiques mor- phologiquesont é téanalysé es enmicroscopieé lectroniqueà balayageet une é tudemorphomé trique détaillé e aétéré alisé e. Lecomplexe buccal est dé crit pour la premiè re fois,fournissant des dé tails surla bouche,les maxillules, les maxilles et les stylets buccaux, avec une nouvelle structure trilobé e, quin’ avaitpas é tédé crite auparavant pour le genre. D’ autresappendices sont aussi pré senté s, inclu- antles antennules, les antennes, les pattes natatoires ainsi que d’ autres structures externes et internes. Lesré sultats sont discuté s etcomparé s avecceux trouvé s dansla littérature, l’ ensemble offrant une imageplus complè te de l’ anatomiede cetteespè ce. 4/ Fax:+34. 942 275072; e-mail: [email protected]. es c KoninklijkeBrill NV ,Leiden,2001 Crustaceana 74(2):193-210 ° 194 P.ABAUNZA, N.L.ARROYO &I.PRECIADO INTRODUCTION Copepodsare oneof the most important groupsof ectoparasites living on shes andother marine animals. Thenumber of parasitic species ofcopepods has beenestimated to bearound 1600 to 1800,75% of which belong to the order Siphonostomatoida(cf. Kabata, 1992). Themembers ofthe family Pennellidae,order Siphonostomatoida, are typically mesoparasitic anddiffer fromother families ofthat order,because in their life cycle theyhave intermediate hosts (Kabata,1979). The genus Pennella Oken, 1816,is amongthe most representative ofthis family,includingthe onlycopepods that parasitize marine mammals, nextto shes andother groups of invertebrates. Inspite ofthis peculiarity andits zoologicalimportance, Pennella is one of the least knowngenera in its family. Thehigh degree of metamorphosis shown bythe adult females, their phenotypicplasticity ,andthe difculty involvedin obtaininga sufcient numberof specimens fordescription, might well explainthe currentsystematic confusionwith regardto the identication ofthe variousspecies (Kabata,1979, 1992). Especially also the buccalarea ofthe pennelidcopepods is still inadequatelyknown, because it is verysmall anddif cult to studyin detail (Castro &Baeza,1991). Pennellabalaenopterae Koren& Danielsen,1877, the largest knowncopepod, is noexception, and there are still manyuncertainties regardingits morphological characters andbiological cycle.The literature onthis species consists ofonly a few papers; most ofthem are outdated,and some are denitely misleading. Anthony &Calvet (1905)provided a generaldescription ofthe species’morphology . Amoredetailed studywas accomplishedby Turner (1905), who analysed the parasite’s internal anatomy.Somemorphometric data canbe found in Brian (1944),who analysed two specimens, andin Raga& Sanpera(1986), who studied specimens collected from nwhales, Balaenopteraphysalus L.,1758, in waters offthe northwesternIberian Peninsula. Other reviewers ofthe genus Pennella havetried to create some orderin the criteria forthe systematics ofthe group, with varyingsuccess. Leigh-Sharpe(1928) and Delamare Deboutteville &Nunes- Ruivo(1951) reviewed the genus,the formerauthor describing the morphology of P.balaenopterae rather inaccurately.Kabata(1979), gave a complete reviewof the species that parasitize sh,whence P.balaenopterae was notincluded. More recently,Hogans(1987) explicitly distinguished between Pennella losa (L., 1758) and P.balaenopterae ,usingexternal morphologicalcharacters. Henextconcluded that P.balaenopterae is the onlyspecies in the genusthat parasitizes marine mammals. Papers ondistribution andreports onthe presenceof P.balaenopterae are notabundant either, the informationbeing con ned to Wilson (1920),Heegard (1943),Margolis &Dailey (1972),Raga & Sanpera(1986), Dailey &Vogelbein.
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