, Volume 64, Issue 3, pp 306–313 | Cite as

The importance of predation, substrate and spatial refugia in determining lotic insect distributions

  • Alexander S. Flecker
  • J. David Allan
Original Papers


An experiment was conducted to evaluate the interaction between predation, substrate, and spatial refugia in the organization of a stream insect community (Reeds Creek, Pendleton Co., West Virginia). Patterns of insect colonization were compared between fish exclusion cages and open controls that allowed access to vertebrate predators. Each cage contained 4 different substrates that varied in the relative amount of spatial refugia. Fish had little influence on the diversity or abundance of any insect taxa, even when spatial refugia were limited. The only significant effect due to predation, was an increased diversity of large (>8 mm) invertebrates in the absence of predators. However, because these taxa were relatively rare, the overall role of fish predation on insect community structure was minimal.

In contrast, substrate had a marked effect on insect colonization. Insects were always more abundant (number/basket) on loose substrates containing large numbers of interstitial spaces, compared to cement-embedded substrates with few refuges available. In addition, invertebrates were more abundant on loose gravel compared to loose cobbles. Howver, when substrate “preferencesrd were examined according to insect density (number/m2), loose cobbles were generally the preferred substrate. The present experiment rejects the hypothesis that patterns of substrate colonization can be explained as differential insect mortality by predators, due to varying amounts of refugia. Alternative mechanisms such as differing amounts of trapped detritus and substrate surface area may account for substrate “preference”.


Detritus Cobble Fish Predation Insect Community Vertebrate Predator 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


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  1. Allan JD (1975) The distributional ecology and diversity of benthic insects in Cement Creek, Colorado. Ecology 56:1040–1053Google Scholar
  2. Allan JD (1982) The effects of reduction in trout density on the invertebrate community of a mountain stream. Ecology 63:1444–1455Google Scholar
  3. Brocksen RW, Davis GE, Warren CE (1968) Competition, food consumption, and production of sculpins and trout in laboratory stream communities. J Wild Manag 32:51–75Google Scholar
  4. Brusven MA, Rose ST (1981) Influence of substrate composition and suspended sediment on insect predation by the torrent sculpin, Cottus rhotheus. Can J Fish Aquat Sci 38:1444–1448Google Scholar
  5. Caswell H (1978) Predator-mediated coexistence: a nonequilibrium model. Amer Natur 112:127–154Google Scholar
  6. Charnov EL, Orians GH, Hyatt K (1976) Ecological implications of resource depression. Am Natur 110:247–259Google Scholar
  7. Coleman MJ, Hynes HBN (1970) The vertical distribution of the invertebrate fauna in the bed of a stream. Limnol Ocean 15:31–40Google Scholar
  8. Connell JH (1975) Some mechanisms producing structure in natural communities: a model and evidence from field studies. In: Cody ML, Diamond JM (eds) Ecology and evolution of communities. Harvard University Press, Cambridge MA, pp 460–490Google Scholar
  9. Cooper WE, Crowder LB (1979) Patterns of predation in simple and complex environments. In: Stroud RH, Clepper H (eds) Predator-prey systems in fisheries management. Sport Fishing Institute, Washington DC, pp 257–267Google Scholar
  10. Cummins KW, Lauff GH (1969) The influence of substrate particle size on the microdistribution of stream macrobenthos. Hydrobiologia 34:145–181Google Scholar
  11. Dethier MN (1980) Tidepools as refuges: predation and the limits of the Harpacticoid copepod Tigriopus californicus (Baker). J Exp Mar Biol Ecol 42:99–111Google Scholar
  12. Egglishaw HJ (1964) The distributional relationship between the bottom fauna and plant detritus in streams. J Anim Ecol 33:463–476Google Scholar
  13. Egglishaw HJ (1968) The quantitative relationship between bottom fauna and plant detritus in streams of different calcium concentrations. J Appl Ecol 5:731–740Google Scholar
  14. Fahy E (1975) Quantitative aspects of the distribution of invertebrates in the benthos of a small stream in Western Ireland. Freshwat Biol 5:167–182Google Scholar
  15. Flecker AS (1984) Experimental studies on the effects of predation and associated factors in structuring an Appalachian stream community. Masters Thesis. Univ. Maryland, College Park p 86Google Scholar
  16. Gause GF (1934) The struggle for existence. Williams and Wilkins, Baltimore MD, p 163Google Scholar
  17. Griffiths RW (1981) The effect of trout predation on the abundance and production of stream insects. Unpubl. Masters Thesis. Univ. British Columbia, Vancouver, p 106Google Scholar
  18. Hall DJ, Cooper WE, Werner EE (1970) An experimental approach to the production dynamics and structure of freshwater animal communities. Limnol Oceangr 15:829–928Google Scholar
  19. Hart DJ (1978) Diversity in stream insects: regulation by rock size and microspatial complexity. Verh Inter Verein Limnol 20:1376–1381Google Scholar
  20. Hildrew AG, Townsend CR (1977) The influence of substrate on the functional response of Plectrocnemia conspera (Curtis) larvae (Trichoptera: Polycentropidae). Oecologia (Berl) 31:21–26Google Scholar
  21. Holling CS (1965) The functional response of predators to prey density and its role in mimicry and population regulation. Mem Entomol Soc Canada 45:1–60Google Scholar
  22. Huffaker CB (1958) Experimental studies on predation: dispersion factors and predator-prey oscilations. Hilgardia 27:343–383Google Scholar
  23. Lubchenco J (1983) Littorina and Fucus: effects of herbivores, substratum heterogeneity, and plant escapes during succession. Ecology 64:1116–1123Google Scholar
  24. Macan TT (1977) The influence of predation on the composition of fresh-water animal communities. Biol Rev 52:45–70Google Scholar
  25. Menge BA (1976) Organization of the New England rocky community: role of predation, competition, and environmental heterogeneity. Ecol Monogr 46:355–393Google Scholar
  26. Menge BA, Sutherland JP (1976) Species diversity gradients: synthesis of the roles of predation, competition, and temporal heterogeneity. Am Natur 110:351–369Google Scholar
  27. Minshall GW, Minshall JN (1977) Microdistribution of benthic invertebrates in a Rocky Mountain (USA) stream. Hydrobiol 55:231–249Google Scholar
  28. Otto C (1982) Habitat, size and distribution of Scandinavian limnephilid caddisflies. Oikos 38:355–360Google Scholar
  29. Otto C, Svensson BS (1980) The significance of case material selection for the survival of caddis larvae. J Anim Ecol 49:855–865Google Scholar
  30. Peckarsky BL (1979) Biological interactions as determinants of distributions of benthic invertebrates within the substrate of stony streams. Limnol Oceangr 24:59–68Google Scholar
  31. Peckarsky BL (1983) Biotic interactions or abiotic limitations? A model of lotic community structure. In: Fontaine TD III, Bartell SM (eds) Dynamics of lotic ecosystems. Ann Arbor Science Publ, Ann Arbor MI. pp 303–323Google Scholar
  32. Peckarsky BL, Dodson SI (1980) An experimental analysis of biological factors contributing to stream community structure. Ecology 61:1283–1290Google Scholar
  33. Rabeni CF, Minshall GW (1977) Factors affecting the microdistribution of stream benthic insects. Oikos 29:33–43Google Scholar
  34. Reice SR (1980) The role of substratum in benthic macro-invertebrate microdistribution and litter decomposition in a woodland stream. Ecology 61:580–590Google Scholar
  35. Reice SR (1983) Predation and substratum: factors in lotic community structure. In: Fontaine TD III, Bartell SM (eds) Dynamics of lotic ecosystems. Ann Arbor Science Publ, Ann Arbor MI, pp 325–345Google Scholar
  36. Shannon CE, Weaver W (1963) The mathematical theory of communication. Univ Illinois Press, Urbana IL. 117 ppGoogle Scholar
  37. Sheldon AL (1977) Colonization curves: application to stream insects on semi-natural substrates. Oikos 28:256–261Google Scholar
  38. Shelly TD (1979) The effect of rock size upon the distribution of species of Orthocladiinae (Chironomidae: Diptera) and Baetis intercalaris McDunnough (Baetidae: Ephemeroptera). Ecol Entomol 4:95–100Google Scholar
  39. Smith FE (1972) Spatial heterogeneity, stability, and diversity in ecosystems. In: Deevey ES (ed) Growth by Intussusception: Ecological essays in honor of G.E. Hutchinson. Florida State Museum, Gainesville FL, pp 307–336Google Scholar
  40. Stein RA (1977) Selective predation, optimal foraging, and the predator-prey interaction between fish and crayfish. Ecology 58:1237–1253Google Scholar
  41. Stein RA (1979) Behavioral response of prey to fish predators. In: Stroud RH, Clepper H (eds) Predator-prey systems in fisheries management. Sport Fishing Institue, Washington DC. pp 343–353Google Scholar
  42. Stein RA, Magnuson JJ (1976) Behavioral response of crayfish to a fish predator. Ecology 57:751–761Google Scholar
  43. Stout J, Vandermeer J (1975) Comparison of species richness for stream-inhabiting insects in tropical and mid-latitude streams. Am Natur 109:263–280Google Scholar
  44. Ulfstrand S (1968) Benthic animal communities in Lapland streams. Oikos Suppl 10:1–116Google Scholar
  45. Walton OE Jr (1980) Invertebrate drift from predator-prey associations. Ecology 61:1486–1497Google Scholar
  46. Walton OE Jr, Reice SR, Andrews RW (1977) The effects of density, sediment particle size, and velocity on drift of Acroneuria abnormis (Plecoptera). Oikos 28:291–298Google Scholar
  47. Ware DM (1972) Predation by rainbow trout (Salmo gairdneri): the influence of hunger, prey density, and prey size. J Fish Res Bd Can 29:1193–1201Google Scholar
  48. Waters TF (1969) Invertebrate drift-ecology and significance to stream fishes. In: Northcote T (ed) Symposium on Salmon and Trout in Streams, University of British Columbia Vancouver, pp 121–134Google Scholar
  49. Waters TF (1972) The drift of stream insects. Ann Rev Entomol 17:253–272Google Scholar
  50. Wiens JA (1977) On competition and variable environments. Am Sci 65:590–597Google Scholar
  51. Wiley MJ (1981) Interacting influences of density and preference on the emigration rates of some lotic chironomid larvae (Diptera: Chironomidae). Ecology 62:426–438Google Scholar
  52. Williams DD (1980) Some relationships between stream benthos and substrate heterogeneity. Limnol Oceangr 25:166–172Google Scholar
  53. Williams DD, Mundie JH (1978) Substrate size selection by stream invertebrates and the influence of sand. Limnol Oceangr 23:1030–1033Google Scholar
  54. Wise DH, Molles MC Jr (1979) Colonization of artificial substrates by stream insects: influence of substrate size and diversity. Hydrobiol 65:69–74Google Scholar
  55. Woodin SA (1978) Refuges, disturbance, and community structure: a marine soft-bottom example. Ecology 59:274–284Google Scholar
  56. Zelinka M (1974) Die Eintagsfliegen (Ephemeroptera) in Forellenbächen der Beskiden. III. Der Einfluß des verschiedenen Fisch-bestandes. Vestnik Cs spol zool 38:76–80Google Scholar
  57. Zelinka M (1976) Mayflies (Ephemeroptera) in the drift of trout streams in the Beskydy Mountains. Acta Bohemoslov 73:94–101Google Scholar

Copyright information

© Springer-Verlag 1984

Authors and Affiliations

  • Alexander S. Flecker
    • 1
  • J. David Allan
    • 1
  1. 1.Department of ZoologyUniversity of MarylandCollege ParkUSA

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