title: Depth segregation and diet disparity revealed by stable isotope analyses in sympatric herbivorous cichlids in Lake Tanganyika.
abstract: BACKGROUND
  Lake Tanganyika in the African Great Rift Valley is known as a site of adaptive radiation in cichlid fishes. Diverse herbivorous fishes coexist on a rocky littoral of the lake. Herbivorous cichlids have acquired multiple feeding ecomorphs, including grazer, browser, scraper, and scooper, and are segregated by dietary niche. Within each ecomorph, however, multiple species apparently coexist sympatrically on a rocky slope. Previous observations of their behavior show that these cichlid species inhabit discrete depths separated by only a few meters. In this paper, using carbon (C) and nitrogen (N) stable isotope ratios as markers, we followed the nutritional uptake of cichlid fishes from periphyton in their feeding territories at various depths.
  
  
  RESULTS
  δ(15)N of fish muscles varied among cichlid ecomorphs; this was significantly lower in grazers than in browsers and scoopers, although δ(15)N levels in periphyton within territories did not differ among territorial species. This suggests that grazers depend more directly on primary production of periphyton, while others ingest animal matter from higher trophic levels. With respect to δ(13)C, only plankton eaters exhibited lower values, suggesting that these fishes depend on production of phytoplankton, while the others depend on production of periphyton. Irrespective of cichlid ecomorph, δ(13)C of periphyton correlated significantly with habitat depth, and decreased as habitat depth became deeper. δ(13)C in territorial fish muscles was significantly related to that of periphyton within their territories, regardless of cichlid ecomorph, which suggests that these herbivorous cichlids depend on primary production of periphyton within their territories.
  
  
  CONCLUSIONS
  Carbon and nitrogen stable isotope ratios varied among ecomorphs and among cichlid species in the same ecomorphs sympatrically inhabiting a littoral area of Lake Tanganyika, suggesting that these cichlids are segregated by nutrient source due to varying dependency on periphyton in different ecomorphs (especially between grazers and browsers), and due to segregation of species of the same ecomorph by feeding depth, grazers and browsers in particular.
authors.affiliation: Graduate School of Science and Engineering, Ehime University, 2-5 Bunkyo, Matsuyama, Ehime Japan.
authors.forename: Hiroki
authors.initials: H
authors.lastname: Hata
authors.affiliation: Center for Marine Environmental Studies (CMES), Ehime University, 2-5 Bunkyo, Matsuyama, Ehime Japan ; Environmental Research and Management Center, Hiroshima University, 1-4-4 Kagamiyama, Higashi-Hiroshima, Hiroshima Japan.
authors.forename: Jyunya
authors.initials: J
authors.lastname: Shibata
authors.affiliation: Center for Marine Environmental Studies (CMES), Ehime University, 2-5 Bunkyo, Matsuyama, Ehime Japan.
authors.forename: Koji
authors.initials: K
authors.lastname: Omori
authors.affiliation: Graduate School of Science, Osaka City University, Sumiyoshi-ku, Osaka Japan.
authors.forename: Masanori
authors.initials: M
authors.lastname: Kohda
authors.affiliation: Kyoto University, Yoshida-Honmachi, Sakyo, Kyoto Japan.
authors.forename: Michio
authors.initials: M
authors.lastname: Hori
keywords: Adaptive radiation; Ecomorph; Herbivore; Stable isotope; Tanganyikan cichlid