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Evolution and Historical Biogeography of a Parasite-Host Assemblage: Alcataenia Spp

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Evolution and Historical Biogeography of a Parasite-Host Assemblage: Alcataenia Spp University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Faculty Publications from the Harold W. Manter Parasitology, Harold W. Manter Laboratory of Laboratory of Parasitology 1986 Evolution and Historical Biogeography of a Parasite-Host Assemblage: Alcataenia spp. (Cyclophyllidea: Dilepididae) in Alcidae (Charadriiformes) Eric P. Hoberg United States Department of Agriculture, Agricultural Research Service, [email protected] Follow this and additional works at: http://digitalcommons.unl.edu/parasitologyfacpubs Part of the Biodiversity Commons, Evolution Commons, and the Parasitology Commons Hoberg, Eric P., "Evolution and Historical Biogeography of a Parasite-Host Assemblage: Alcataenia spp. (Cyclophyllidea: Dilepididae) in Alcidae (Charadriiformes)" (1986). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 783. http://digitalcommons.unl.edu/parasitologyfacpubs/783 This Article is brought to you for free and open access by the Parasitology, Harold W. Manter Laboratory of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications from the Harold W. Manter Laboratory of Parasitology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Evolution and historical biogeography of a parasite-host assemblage: Alcataenia spp. (Cyclophyllidea: Dilepididae) in Alcidae (Charadriiformes) ERICP. HOBERG College of Veterinary Medicine, Oregon State University, Corvallis, OR, U.S.A. 97331-4802 Received April 25, 1986 HOBERG,E. P. 1986. Evolution and historical biogeography of a parasite-host assemblage: Alcataenia spp. (Cyclophyllidea: Dilepididae) in Alcidae (Charadriiformes) . Can. J. Zool . 64: 2576-2589. The methodology of phylogenetic systematics was used to develop hypotheses for the evolution of eight species of Alcataenia, a group of host-specific cestodes of the Alcidae and, to a lesser extent, the Laridae (Charadriiformes). Concurrently, aspects of the early biogeography of alcids were reevaluated making it possible to study the historical and distributional relationships of parasites and hosts. The most parsimonious hypothesis for the phylogeny of Ale-crtcrenicr suggested that sequential colonization or host switching with limited coevolution (coaccommodation) by parasites best explained the distributional patterns exhibited by Alcataenia spp. Morphological evolution of specific species of Alcataenia accompanied host switching, but was limited subsequent to the initial event of colonization. Thus evolution of these parasites following colonization lagged behind continuing diversification of the host group. Although alcids are an ancient group, as indicated by palaeontological and phylogenetic data, their cestode fauna is apparently not. It is postulated that Alcataenia spp. were acquired by their characteristic hosts in the late Pliocene and early Pleistocene, following diversification of the Alcidae at the generic level during the Miocene. Host distributions of respective Alcataenia spp. are more narrow than expected, considering the relatively young age of this assemblage. The current paradigm linking pronounced host specificity with coevolution of hosts and parasites in assemblages of great evolutionary age is not supported. HOBERG,E. P. 1986. Evolution and historical biogeography of a parasite-host assemblage: Alcataenia spp. (Cyclophyllidea: Dilepididae) in Alcidae (Charadriiforrnes). Can. J . Zool. 64: 2576-2589. La mCthodologie propre a la systCmatique phylogCnCtique a servi a Claborer des hypotheses sur 1'6volution de huit especes d'Alcataenia, un groupe de cestodes spkifiques aux Alcidae et, a un degr6 moindre, aux Laridae (Charadriiformes). En outre, certains aspects de la biogkographie primitive des alcidCs ont CtC rkexamines de fason a permettre 1'Ctude des relations historiques et geographiques des parasites et des hates. L'hypothese la plus simple sur la phylogenie d'Alcataenia suggere que la colonisation ~Cquentielle,ou changement d'h6te par les parasites et coCvolution limitCe (coaccomodation),est le phCnomkne qui peut 1e mieux expliquer la repartition actuelle des especes d'Alcatuenia. Certaines espkces particulieres d'Alcataenia ont subi une Cvolution morphologique en changeant d'hate, mais cette Cvolution a 6tC ralentie apres le debut de la colonisation. L'6volution de ces parasites aprks la colonisation a donc pris du retard par rapport a la diversification continue du groupe hate. Bien que les Alcidae constituent un groupe ancien, d'apres 1es donnCes pal6ontologiques et phylogCnCtiques, la faune de cestodes qui les parasite ne semble pas l'etre. I1 semble que les diverses especes d'Alcatuenia soient devenues les parasites de leurs hates respectifs vers la fin du Pliocene et le dCbut du Pleistockne, a la suite de la diversification des Alcidae au niveau gCnCrique au cours du Miocene. La ripartition chez les hates des diverses espkces d'Alcataenia n'est pas aussi repandue que pourrait le laisser croire l'existence relativement rCcente de ces associations. Cette Ctude ne confirme pas le paradigme g6nCralement accept6 selon lequel une grande spCcificitC d'hbte est reliCe a la co6volution des hates et des parasites dans les associations trks anciennes. [Traduit par la revue] Introduction las) , Cyclorrhynchus psittacula (Pallas), Aethia cristatella Coevolution of parasite-host assemblages is a widely recog- (Pallas), A. pusilla (Pallas), and A. pygmaea (Gmelin));murre- nized concept (Brooks 1979a; Mitter and Brooks 1983; and lets are represented by two genera and six species (Brachyram- others). There have been relatively few studies objectively phus marmoratus (Gmelin), B . brevirostris (Vigors), Svnthli- assessing the extent and significance of this phenomenon boramphus antiquus (Gmelin), S. wumizuzume (Temminck), (Brooks 1977, 1978, 1979a, 19796; Brooks et al. 198 1 ; Platt S. craveri (Salvadori), and S. hypoleucus (Xantus de Vesey)); 1984; Glen and Brooks 1985) and none have dealt with guillemots are monogeneric with three species (Cepphus carbo cyclophyllidean cestodes in avian hosts. Krabbe (1869) and Pallas, C. columba Pallas, and C. grylle (Linnaeus)); dovkies Fuhrmann (1908) were among the first to critically recognize are monotypic (Alle alle (Linnaeus)); and the auks include the that specific taxa of cestodes were often characteristic of limited murres ( Uriu aalge (Pontoppidan), U. lomvia (Linnaeus)), and phylogenetically related taxa of birds. The analysis reported razorbills (Alca torda Linnaeus), and the recently extinct great here is the first attempt to develop hypotheses for the evolution auk (Pinguinus impennis (Linnaeus)).The family is restricted to and historical biogeography of Alcataenia spp. (Dilepidinae high latitudes in the northern hemisphere, and most taxa are Fuhrmann, 1907) and their charadriiform hosts. endemic to either the Pacific basin (16 species) or the Atlantic The principle hosts for Alcataenia spp., seabirds of the family basin (3 species, including the great auk). Only four genera and Alcidae, represent a monophyletic assemblage of genera and four species have distributions that are circumpolar (Udvardy species of great evolutionary age (Kozlova 1957; Storer 1960; 1963, 1979). Most members of the family have ranges centred Udvardy 1963; Olson 1985; Strauch 1985; and others). The in the region of the Bering Sea, including the Aleutian Islands, family contains 23 Recent species distributed among 12 genera: islands of the Bering Sea, and some areas of the surrounding puffins include two genera and four species (Cerorhinca coasts (for range maps, see Storer 1952; Udvardy 1963; Kozlova monocerata (PalIas) , Fratercula cirrhata (Pallas), F. arctica 1957). Consequently, the centre of origin for the family is (Linnaeus), and F. corniculata (Naumann)); auklets comprise considered to be the north Pacific (Udvardy 1963; Olson 1985). three genera and five species (Ptychoramphus aleuticus (Pal- Several works have dealt with biogeography and evolution Printed in Canada I lmprime au Canada HOBERG 2577 within the family (Kaftanovskii 195 1; Storer 1945a, 1945b, the study. The following were included in the ingroup: Alcataenia 1952, 1960; Kozlova 1957; Udvardy 1963; Bkdard 1969). campylacantha, A. armillaris, A. meinertzhageni, A. longicervica, A. Recent advances in interpretation of the phylogenetic (Strauch pygmaeus, A. fraterculae, A. cerorhincae, and A. larina (with A. I. 1985) and palaeontological (Olson 1985) relationships of the larina and A. I. pacifica considered as distinct). Two species, A. family have made it necessary to reevaluate the historical micracantha and A. dominicanus were excluded because of uncertain taxonomic relationships, particularly with reference to other nominal biogeography of the alcids. New hypotheses for the distribu- taxa that have been suppressed as their synonyms (see Joyeux and Baer tional history of alcids are presented in the current study. 1954; Odening 1982; Zdzitoweicki and Szelenbaum-Cielecka 1984). In the Holarctic, species of Alcataenia Spasskaya, 197 1 are Specimens of all Alcataenia spp. and representatives of several host-specific parasites of the Alcidae and, to a lesser extent, the prominent genera of the Dilepidinae were examined. A detailed listing Larinae (Charadriiformes) (Spasskaya 197 1 ; Hoberg 1984a, of specimens of A. campylacantha, A. armillaris, A. meinertzhageni, 1984b). The genus contains seven species in alcids: Alcataenia A. pygmaeus, A. fraterculae, A. cerorhincae, A. longicervica, and
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