, Volume 16, Issue 3, pp 478–490 | Cite as

Cross-scale Habitat Structure Drives Fish Body Size Distributions on Coral Reefs

  • Kirsty L. NashEmail author
  • Nicholas A. J. Graham
  • Shaun K. Wilson
  • David R. Bellwood


Despite a large number of studies focusing on the complexity of coral reef habitats and the characteristics of associated fish assemblages, the relationship between reef structure and fish assemblages remains unclear. The textural discontinuity hypothesis, which proposes that multi-modal body size distributions of organisms are driven by discontinuous habitat structure, provides a theoretical basis that may explain the influence of habitat availability on associated organisms. In this study we use fractal techniques to characterize patterns of cross-scale habitat complexity, and examine how this relates to body-depth abundance distributions of associated fish assemblages over corresponding spatial scales. Our study demonstrates that: (1) Reefs formed from different underlying substrata exhibit distinct patterns of cross-scale habitat complexity; (2) The availability of potential refuges at different scales correlates with patterns in fish body depth distributions, but habitat structure is more strongly related to the relative abundance of fish in the body depth modes, rather than to the number of modes; (3) As reefs change from coral- to algal-dominated states, the complexity of the underlying reef substratum may change, presenting a more homogenous environment to associated assemblages; (4) Individual fish body depth distributions may be multi-modal, however, these distributions are not static characteristics of the fish assemblage and may change to uni-modal forms in response to changing habitat condition. In light of predicted anthropogenic changes, there is a clear need to improve our understanding of the scale of ecological relationships to anticipate future changes and vulnerabilities.


textural discontinuity hypothesis structural complexity rugosity reef degradation coral-dominated reef algal-dominated reef 



This study was funded by the Australian Research Council and the Queensland Smart Futures Fund. We thank Ross Barrett for making the measuring wheels, and two anonymous reviewers for their helpful comments.

Supplementary material

10021_2012_9625_MOESM1_ESM.doc (12 kb)
Supplementary material 1 (DOC 11 kb)


  1. Ackerman JL, Bellwood DR, Brown J. 2004. The contribution of small individuals to density-body size relationships: examination of energetic equivalence in reef fishes. Oecologia 139:568-571.PubMedCrossRefGoogle Scholar
  2. Ackerman JL, Bellwood DR. 2000. Reef fish assemblages: a re-evaluation using enclosed rotenone stations. Mar Ecol Prog Ser 206:227-237.CrossRefGoogle Scholar
  3. Allen CR, Garmestani AS, Havlicek TD, Marquet PA, Peterson GD, Restrepo C, Stow CA, Weeks BE. 2006. Patterns in body mass distributions: sifting among alternative hypotheses. Ecol Lett 9:630-643.PubMedCrossRefGoogle Scholar
  4. Alvarez-Filip L, Dulvy NK, Gill JA, Cote IM, Watkinson AR. 2009. Flattening 465 of Caribbean coral reefs: region-wide declines in architectural complexity. Proc R Soc B 276:3019-3025.CrossRefGoogle Scholar
  5. Andres NG, Rodenhouse NL. 1993. Resilience of corals to hurricanes—a simulation-model. Coral Reefs 12:167–75.CrossRefGoogle Scholar
  6. Bartholomew A, Shine RL. 2008. Space size relative to prey width (Sp/Py) influences macrofaunal colonization of artificial structures. Mar Ecol-Prog Ser 358:95–102.CrossRefGoogle Scholar
  7. Bejarano S, Mumby P, Sotheran I. 2011. Predicting structural complexity of reefs and fish abundance using acoustic remote sensing (RoxAnn). Mar Biol 158:489–504.CrossRefGoogle Scholar
  8. Bellwood DR, Hoey AS, Ackerman JL, Depczynski M. 2006. Coral bleaching, reef fish community phase shifts and the resilience of coral reefs. Glob Change Biol 12:1587–94.CrossRefGoogle Scholar
  9. Benjamini Y, Hochberg Y. 1995. Controlling the false discovery rate: a practical and powerful approach to multiple testing. J R Stat Soc Ser B (Met) 57:289–300.Google Scholar
  10. Bergman KC, Ohman MC, Svensson S. 2000. Influence of habitat structure on Pomacentrus sulfureus, a western Indian Ocean reef fish. Environ Biol Fishes 59:243–52.CrossRefGoogle Scholar
  11. Blackburn TM, Gaston KJ. 1994. Animal body size distributions: patterns, mechanisms and implications. Trends Ecol Evol 9:471–4.PubMedCrossRefGoogle Scholar
  12. Bradbury RH, Reichelt RE, Green DG. 1984. Fractals in ecology: methods and interpretation. Mar Ecol Prog Ser 14:295–6.CrossRefGoogle Scholar
  13. Brown JH, Nicoletto PF. 1991. Spatial scaling of species composition: body masses of North American land mammals. Am Nat 138:1478–512.CrossRefGoogle Scholar
  14. Buckland ST, Magurran AE, Green RE, Fewster RM. 2005. Monitoring change in biodiversity through composite indices. Philos Trans R Soc B 360:243–54.CrossRefGoogle Scholar
  15. Cattaneo A. 1993. Size spectra of benthic communities in Laurentian streams. Can J Fish Aquat Sci 50:2659–66.CrossRefGoogle Scholar
  16. Clarke KR. 1993. Non-parametric multivariate analyses of changes in community structure. Aust J Ecol 18:117–43.CrossRefGoogle Scholar
  17. Cohen JE, Jonsson T, Carpenter SR. 2003. Ecological community description using the food web, species abundance, and body size. Proc Natl Acad Sci USA 100:1781–6.PubMedCrossRefGoogle Scholar
  18. Davenport J. 2004. Fractal dimension estimation in studies of epiphytal and epilithic communities: strengths and weaknesses. In: Seuront L, Strutton PG, Eds. Handbook of scaling methods in aquatic ecology. Boca Raton: CRC Press. p. 245–256.Google Scholar
  19. Dornelas M, Connolly SR, Hughes TP. 2006. Coral reef diversity refutes the neutral theory of biodiversity. Nature 440:80–2.PubMedCrossRefGoogle Scholar
  20. Emslie MJ, Cheal AJ, Sweatman H, Delean S. 2008. Recovery from disturbance of coral and reef fish communities on the Great Barrier Reef, Australia. Mar Ecol-Prog Ser 371:177–90.CrossRefGoogle Scholar
  21. Ernest SKM. 2005. Body size, energy use, and community structure of small mammals. Ecology 86:1407–13.CrossRefGoogle Scholar
  22. Fewster RM, Buckland ST, Siriwardena GM, Baillie SR, Wilson JD. 2000. Analysis of population trends for farmland birds using generalized additive models. Ecology 81:1970–84.CrossRefGoogle Scholar
  23. Fischer J, Lindenmayer DB, Montague-Drake R. 2008. The role of landscape texture in conservation biogeography: a case study on birds in south-eastern Australia. Divers Distrib 14:38–46.CrossRefGoogle Scholar
  24. Friedlander AM, Brown EK, Jokiel PL, Smith WR, Rodgers KS. 2003. Effects of habitat, wave exposure, and marine protected area status on coral reef fish assemblages in the Hawaiian archipelago. Coral Reefs 22:291–305.CrossRefGoogle Scholar
  25. Friedlander AM, DeMartini EE. 2002. Contrasts in density, size, and biomass of reef fishes between the northwestern and the main Hawaiian Islands: the effects of fishing down apex predators. Mar Ecol-Prog Ser 230:253–64.CrossRefGoogle Scholar
  26. Friedlander AM, Parrish JD. 1998. Habitat characteristics affecting fish assemblages on a Hawaiian coral reef. J Exp Mar Biol Ecol 224:1–30.CrossRefGoogle Scholar
  27. Gagné SA, Proulx R, Fahrig L. 2008. Testing Holling’s textural-discontinuity hypothesis. J Biogeogr 35:2149–50.CrossRefGoogle Scholar
  28. Gotelli NJ, Ellison AM. 2004. A primer of ecological statistics. Sunderland, MA: Sinauer Associates Inc. p 510 p.Google Scholar
  29. Graham NAJ, Dulvy NK, Jennings S, Polunin NVC. 2005. Size-spectra as indicators of the effects of fishing on coral reef fish assemblages. Coral Reefs 24:118–24.CrossRefGoogle Scholar
  30. Graham NAJ, Wilson SK, Jennings S, Polunin NVC, Bijoux JP, Robinson J. 2006. Dynamic fragility of oceanic coral reef ecosystems. Proc Natl Acad Sci USA 103:8425–9.PubMedCrossRefGoogle Scholar
  31. Graham NAJ, Wilson SK, Jennings S, Polunin NVC, Robinson J, Bijoux JP, Daw TM. 2007. Lag effects in the impacts of mass coral bleaching on coral reef fish, fisheries, and ecosystems. Conserv Biol 21:1291–300.PubMedCrossRefGoogle Scholar
  32. Greenwood JJD, Gregory RD, Harris S, Morris PA, Yalden DW. 1996. Relations between abundance, body size and species number in British birds and mammals. Philos Trans R Soc Lond B Biol Sci 351:265–78.CrossRefGoogle Scholar
  33. Halley JM, Hartley S, Kallimanis AS, Kunin WE, Lennon JJ, Sgardelis SP. 2004. Uses and abuses of fractal methodology in ecology. Ecol Lett 7:254–71.CrossRefGoogle Scholar
  34. Hartley S, Kunin WE, Lennon JJ, Pocock MJO. 2004. Coherence and discontinuity in the scaling of species’ distribution patterns. Proc R Soc Lond 271:81–8.CrossRefGoogle Scholar
  35. Havlicek TD, Carpenter SR. 2001. Pelagic species size distributions in lakes: are they discontinuous? Limnol Oceanogr 46:1021–33.CrossRefGoogle Scholar
  36. Hixon MA, Beets JP. 1993. Predation, prey refuges, and the structure of coral-reef fish assemblages. Ecol Monogr 63:77–101.CrossRefGoogle Scholar
  37. Hoey AS, Bellwood DR. 2011. Suppression of herbivory by macroalgal density: a critical feedback on coral reefs? Ecol Lett 14:267–73.PubMedCrossRefGoogle Scholar
  38. Holling CS. 1992. Cross-scale morphology, geometry, and dynamics of ecosystems. Ecol Monogr 62:447–502.CrossRefGoogle Scholar
  39. Hughes TP, Rodrigues MJ, Bellwood DR, Ceccarelli D, Hoegh-Guldberg O, McCook L, Moltschaniwskyj N, Pratchett MS, Steneck RS, Willis B. 2007. Phase shifts, herbivory, and the resilience of coral reefs to climate change. Curr Biol 17:360–5.PubMedCrossRefGoogle Scholar
  40. Jennings S, Grandcourt EM, Polunin NVC. 1995. The effects of fishing on the diversity, biomass and trophic structure of Seychelles’ reef fish communities. Coral Reefs 14:225–35.Google Scholar
  41. Jennings S, Greenstreet SPR, Reynolds JD. 1999. Structural change in an exploited fish community: a consequence of differential fishing effects on species with contrasting life histories. J Anim Ecol 68:617–27.CrossRefGoogle Scholar
  42. Kaufman LS. 1983. Effects of Hurricane Allen on reef fish assemblages near discovery Bay, Jamaica. Coral Reefs 2:43–7.CrossRefGoogle Scholar
  43. Klinkenberg B. 1994. A review of methods used to determine the fractal dimension of linear features. Math Geol 26:23–46.CrossRefGoogle Scholar
  44. Kovalenko K, Thomaz S, Warfe D. 2012. Habitat complexity: approaches and future directions. Hydrobiologia 685:1–17.CrossRefGoogle Scholar
  45. Leaper R, Raffaelli D, Emes C, Manly B. 2001. Constraints on body-size distributions: an experimental test of the habitat architecture hypothesis. J Anim Ecol 70:248–59.CrossRefGoogle Scholar
  46. Ledlie M, Graham NAJ, Bythell J, Wilson SK, Jennings S, Polunin NVC, Hardcastle J. 2007. Phase shifts and the role of herbivory in the resilience of coral reefs. Coral Reefs 26:641–53.CrossRefGoogle Scholar
  47. Levin PS, Hay ME. 1996. Responses of temperate reef fishes to alterations in algal structure and species composition. Mar Ecol-Prog Ser 134:37–47.CrossRefGoogle Scholar
  48. Levin SA. 1992. The problem of pattern and scale in ecology: the Robert H. MacArthur Award Lecture. Ecology 73:1943–67.CrossRefGoogle Scholar
  49. Mandelbrot BB. 1982. The fractal geometry of nature. New York: W.H. Freeman and Company. 468 p.Google Scholar
  50. Martin-Garin B, Lathuiliere B, Verrecchia EP, Geister J. 2007. Use of fractal dimensions to quantify coral shape. Coral Reefs 26:541–50.CrossRefGoogle Scholar
  51. McAbendroth L, Ramsay PM, Foggo A, Rundle SD, Bilton DT. 2005. Does macrophyte fractal complexity drive invertebrate diversity, biomass and body size distributions? Oikos 111:279–90.CrossRefGoogle Scholar
  52. Morse DR, Lawton JH, Dodson MM, Williamson MH. 1985. Fractal dimension of vegetation and the distribution of arthropod body lengths. Nature 314:731–3.CrossRefGoogle Scholar
  53. Munday PL, Jones GP. 1998. The ecological implications of small body size among coral-reef fishes. Oceanogr Mar Biol Annu Rev 36:373–411.Google Scholar
  54. O’Neill RV, DeAngelis DL, Waide JB, Allen TFH. 1986. A hierarchical concept of ecosystems. Princeton, NJ: Princeton University Press.Google Scholar
  55. Olden JD, Hogan ZS, Zanden MJV. 2007. Small fish, big fish, red fish, blue fish: size-biased extinction risk of the world’s freshwater and marine fishes. Glob Ecol Biogeogr 16:694–701.CrossRefGoogle Scholar
  56. Peterson GD, Allen CR, Holling CS. 1998. Ecological resilience, biodiversity, and scale. Ecosystems 1:6–18.CrossRefGoogle Scholar
  57. Pratchett M, Munday PL, Wilson SK, Graham NAJ, Cinner J, Bellwood DR, Jones GP, Polunin NVC, McClanahan T. 2008. Effects of climate-induced coral bleaching on coral-reef fishes: ecological and economic consequences. Oceanogr Mar Biol Annu Rev 46:251–96.CrossRefGoogle Scholar
  58. Purkis S, Kohler K. 2008. The role of topography in promoting fractal patchiness in a carbonate shelf landscape. Coral Reefs 27:977–89.CrossRefGoogle Scholar
  59. R Development Core Team. 2011. R: a language and environment for statistical computing. Vienna, Austria: R Project for Statistical Computing.Google Scholar
  60. Robson BJ, Barmuta LA, Fairweather PG. 2005. Methodological and conceptual issues in the search for a relationship between animal body-size distributions and benthic habitat architecture. Mar Freshw Res 56:1–11.CrossRefGoogle Scholar
  61. Silliman BR, van der Koppel J, Bertness MD, Stanton LE, Mendelssohn IA. 2005. Drought, snails, and large-scale die-off of southern U.S. salt marshes. Science 310:1803–6.PubMedCrossRefGoogle Scholar
  62. Stead TK, Schmid-Araya JM, Schmid PE, Hildrew AG. 2005. The distribution of body size in a stream community: one system, many patterns. J Anim Ecol 74:475–87.CrossRefGoogle Scholar
  63. Sugihara G, May R. 1990. Applications of fractals in ecology. Trends Ecol Evol 5:79–86.PubMedCrossRefGoogle Scholar
  64. Thibault KM, White EP, Hurlbert AH, Ernest SKM. 2011. Multimodality in the individual size distributions of bird communities. Glob Ecol Biogeogr 20:145–53.CrossRefGoogle Scholar
  65. Tilman D, Fargione J, Wolff B, D’Antonio C, Dobson A, Howarth R, Schindler D, Schlesinger WH, Simberloff D, Swackhamer D. 2001. Forecasting agriculturally driven global environmental change. Science 292:281–4.PubMedCrossRefGoogle Scholar
  66. Tokeshi M, Arakaki S. 2012. Habitat complexity in aquatic systems: fractals and beyond. Hydrobiologia 685:27–47.CrossRefGoogle Scholar
  67. Vergnon R, Dulvy NK, Freckleton RP. 2009. Niches versus neutrality: uncovering the drivers of diversity in a species-rich community. Ecol Lett 12:1079–90.PubMedCrossRefGoogle Scholar
  68. Watson D, Harvey E, Fitzpatrick B, Langlois T, Shedrawi G. 2010. Assessing reef fish assemblage structure: how do different stereo-video techniques compare? Mar Biol 157:1237–50.CrossRefGoogle Scholar
  69. White EP, Ernest SKM, Kerkhoff AJ, Enquist BJ. 2007. Relationships between body size and abundance in ecology. Trends Ecol Evol 22:323–30.PubMedCrossRefGoogle Scholar
  70. Wiens JA. 1989. Spatial scaling in ecology. Funct Ecol 3:385–97.CrossRefGoogle Scholar
  71. Wilding TA, Rose CA, Downie MJ. 2007. A novel approach to measuring subtidal habitat complexity. J Exp Mar Biol Ecol 353:279–86.CrossRefGoogle Scholar
  72. Wilkinson CR. 2000. World-wide coral reef bleaching and mortality during 1998: a global climate change warning for the new millenium? In: Sheppard CRC, Ed. Seas at the millennium: an environmental evaluation. Amsterdam: Elsevier Science. p. 43–57.Google Scholar
  73. Wilson SK, Fisher R, Pratchett MS, Graham NAJ, Dulvy NK, Turner RA, Cakacaka A, Polunin NVC. 2010. Habitat degradation and fishing effects on the size structure of coral reef fish communities. Ecol Appl 20:442–51.PubMedCrossRefGoogle Scholar
  74. Wilson SK, Fisher R, Pratchett MS, Graham NAJ, Dulvy NK, Turner RA, Cakacaka A, Polunin NVC, Rushton SP. 2008. Exploitation and habitat degradation as agents of change within coral reef fish communities. Glob Change Biol 14:2796–809.CrossRefGoogle Scholar
  75. Wilson SK, Graham NAJ, Polunin NVC. 2007. Appraisal of visual assessments of habitat complexity and benthic composition on coral reefs. Mar Biol 151:1069–76.CrossRefGoogle Scholar
  76. Wilson SK, Graham NAJ, Robinson J, Nash KL, Chong-Seng K, Polunin NVC, Aumeeruddy R, Quatre R. 2012. Macroalgal expansion and marine protected areas drive coral recovery following climatic disturbances. Conserv Biol 26:995–1004.Google Scholar
  77. Wood SN. 2006. Generalized additive models: an introduction with R. Boca Raton: Chapman & Hall/CRC. p. 410.Google Scholar
  78. Yamanaka T, White PCL, Spencer M, Raffaelli D. 2012. Patterns and processes in abundance-body size relationships for marine benthic invertebrates. J Anim Ecol 81:463–71.PubMedCrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media New York 2012

Authors and Affiliations

  • Kirsty L. Nash
    • 1
    Email author
  • Nicholas A. J. Graham
    • 1
  • Shaun K. Wilson
    • 2
    • 3
  • David R. Bellwood
    • 1
    • 4
  1. 1.Australian Research Council Centre of Excellence for Coral Reef Studies, James Cook UniversityTownsvilleAustralia
  2. 2.Department of Environment and ConservationMarine Science ProgramKensingtonAustralia
  3. 3.The Oceans Institute, University of Western AustraliaCrawleyAustralia
  4. 4.School of Marine and Tropical Biology, James Cook UniversityTownsvilleAustralia

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