title: Understanding trophic interactions in host-parasite associations using stable isotopes of carbon and nitrogen.
abstract: Stable isotope analysis of carbon and nitrogen can deliver insights into trophic interactions between organisms. While many studies on free-living organisms are available, the number of those focusing on trophic interactions between hosts and their associated parasites still remains scarce. In some cases information about taxa (e.g. acanthocephalans) is completely missing. Additionally, available data revealed different and occasionally contrasting patterns, depending on the parasite's taxonomic position and its degree of development, which is most probably determined by its feeding strategy (absorption of nutrients through the tegument versus active feeding) and its localization in the host.
  
  
  
  Using stable isotope analysis of carbon and nitrogen we provided first data on the trophic position of an acanthocephalan species with respect to its fish host. Barbels (Barbus barbus) infected only with adult acanthocephalans Pomphorhynchus laevis as well as fish co-infected with the larval (L4) nematodes Eustrongylides sp. from host body cavity were investigated in order to determine the factors shaping host-parasite trophic interactions. Fish were collected in different seasons, to study also potential isotopic shifts over time, whereas barbels with single infection were obtained in summer and co-infected ones in autumn.
  
  
  
  Acanthocephalans as absorptive feeders showed lower isotope discrimination values of δ15N than the fish host. Results obtained for the acanthocephalans were in line with other parasitic taxa (e.g. cestodes), which exhibit a similar feeding strategy. We assumed that they feed mainly on metabolites, which were reprocessed by the host and are therefore isotopically lighter. In contrast, the nematodes were enriched in the heavier isotope δ15N with respect to their host and the acanthocephalans, respectively. As active feeders they feed on tissues and blood in the body cavity of the host and thus showed isotope discrimination patterns resembling those of predators. We also observed seasonal differences in the isotope signatures of fish tissues and acanthocephalans, which were attributed to changes in food composition of the host and to seasonality in the transmission and development of acanthocephalans.
  
  
  
  This study provided first data on trophic interaction between an acanthocephalan species and its associated host, which support the tendency already described for other taxa with similar nutrition strategy (e.g. cestodes). Actively feeding taxa such as larval Eustrongylides sp., appear to act like predators as it can be seen from their isotope discrimination values. However, future research on additional host-parasite systems and especially on acanthocephalans is needed in order to corroborate these conclusions.
authors.affiliation: Aquatic Ecology, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany. milen.nachev@uni-due.de.|Centre for Water and Environmental Research, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany. milen.nachev@uni-due.de.
authors.forename: Milen
authors.initials: M
authors.lastname: Nachev
authors.affiliation: Centre for Water and Environmental Research, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.|Instrumental Analytical Chemistry, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.
authors.forename: Maik A
authors.initials: MA
authors.lastname: Jochmann
authors.affiliation: Aquatic Ecology, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.|Instrumental Analytical Chemistry, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.
authors.forename: Friederike
authors.initials: F
authors.lastname: Walter
authors.affiliation: Instrumental Analytical Chemistry, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.
authors.forename: J Benjamin
authors.initials: JB
authors.lastname: Wolbert
authors.affiliation: Instrumental Analytical Chemistry, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.
authors.forename: S Marcel
authors.initials: SM
authors.lastname: Schulte
authors.affiliation: Centre for Water and Environmental Research, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.|Instrumental Analytical Chemistry, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.
authors.forename: Torsten C
authors.initials: TC
authors.lastname: Schmidt
authors.affiliation: Aquatic Ecology, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.|Centre for Water and Environmental Research, University of Duisburg-Essen, Universitätsstr. 5, D-45141, Essen, Germany.|Department of Zoology, University of Johannesburg, PO Box 524, Auckland Park 2006, Johannesburg, South Africa.
authors.forename: Bernd
authors.initials: B
authors.lastname: Sures
mesh_terms: D000049:Acanthocephala / Q000502:physiology*; D000818:Animals; D017162:Ascaridida / Q000502:physiology; D017191:Ascaridida Infections / Q000469:parasitology / Q000662:veterinary; D002244:Carbon / Q000378:metabolism*; D002247:Carbon Isotopes; D003530:Cyprinidae / Q000469:parasitology*; D005393:Fish Diseases / Q000469:parasitology*; D020387:Food Chain; D006374:Helminthiasis, Animal / Q000469:parasitology*; D006790:Host-Parasite Interactions; D009584:Nitrogen / Q000378:metabolism*; D009587:Nitrogen Isotopes; D009752:Nutritional Status
chemical_list: D002247:Carbon Isotopes; D009587:Nitrogen Isotopes; D002244:Carbon; D009584:Nitrogen
keywords: Acanthocephala; Nematoda; Parasites; Stable isotopes; Trophic interactions