Springer Nature is making SARS-CoV-2 and COVID-19 research free. View research | View latest news | Sign up for updates

Large-scale manipulation of mayfly recruitment affects population size


Recruitment establishes the initial size of populations and may influence subsequent population dynamics. Although strong inference can be made from empirical relationships between recruitment and population sizes, a definitive test of recruitment limitation requires manipulating recruitment at relevant spatial and temporal scales. We manipulated oviposition of the mayfly Baetis bicaudatus in multiple streams and measured the abundance of late-stage larvae at the end of the cohort. Based on fundamental knowledge of mayfly behavior, we increased, eliminated, or left unmodified preferred mayfly oviposition sites in 45-m reaches of streams (N = 4) of one high-altitude drainage basin in western Colorado, USA. We compared egg densities before (2001) and after the manipulation (2002) using paired t tests and compared larval densities before and after the manipulation among treatments using repeated measures analysis of variance. This manipulation altered not only egg densities, but also larval abundances 1 year later. Compared to the previous year, we experimentally increased egg densities at the addition sites by approximately fourfold, reduced egg densities to zero in the subtraction sites, and maintained egg densities in the control sites. After the manipulation, larval densities increased significantly by a factor of approximately 2.0 in the addition sites and decreased by a factor of approximately 2.5 in the subtraction sites. This outcome demonstrates that dramatic changes in recruitment can limit larval population size at the scale of a stream reach, potentially masking previously observed post-recruitment processes explaining the patterns of variation in abundance of a stream insect. Furthermore, our results emphasize the importance of preferred oviposition habitats to population sizes of organisms.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3


  1. Allan JD (1981) Determinants of diet of brook trout (Salvelinus fontinalis) in a mountain stream. Can J Fish Aquat Sci 38:184–192

  2. Allan JD (1982) The effects of reduction in trout density on the invertebrate community of a mountain stream. Ecology 63:1444–1455

  3. Allan JD, Russek E (1985) The quantification of stream drift. Can J Fish Aquat Sci 42:210–215

  4. Álvarez M, Peckarsky BL (2005) How do grazers affect periphyton heterogeneity in streams? Oecologia 142:576–587

  5. Brittain JE (1982) Biology of mayflies. Annu Rev Entomol 27:119–147

  6. Bunn SE, Hughes JM (1997) Dispersal and recruitment in streams: evidence from genetic studies. J N Am Benthol Soc 16:338–346

  7. Caley MJ, Carr MH, Hixon MA, Hughes TP, Jones GP, Menge BA (1996) Recruitment and the local dynamics of open marine populations. Annu Rev Ecol Syst 27:477–500

  8. Cole BJ, Wiernasz DC (2002) Recruitment limitation and population density in the harvester ant, Pogonomyrmex occidentalis. Ecology 83:1433–1442

  9. Connell JH (1985) The consequences of variation in initial settlement vs post-settlement mortality in rocky intertidal communities. J Exp Mar Biol Ecol 93:11–45

  10. Connell JH, Green PT (2000) Seedling dynamics over thirty-two years in a tropical rain forest tree. Ecology 81:568–584

  11. Cooper SD, Walde SJ, Peckarsky BL (1990) Prey exchange rates and the impact of predators on prey populations in streams. Ecology 71:1503–1514

  12. Dimond JB (1967) Evidence that drift of stream benthos is density related. Ecology 48:855–857

  13. Doherty PJ (1983) Tropical territorial damselfishes: is density limited by aggregation or recruitment? Ecology 64:176–190

  14. Downes BJ, Keough MJ (1998) Scaling of colonization processes in streams: parallels and lessons form marine hard substrata. Aust J Ecol 23:8–26

  15. Eaton AE (1888) A revisional monograph of the recent Ephemeridae or mayflies. Trans Linn Soc 3:1–352

  16. Encalada AC (2005) Patterns, mechanisms and consequences to population dynamics of selective oviposition behavior by Baetis bicaudatus (Ephemeroptera: Baetidae). PhD thesis. Cornell University, Ithaca

  17. Encalada AC, Peckarsky BL (2011) The influence of recruitment on within generation population dynamics of a mayfly. Ecosphere 2(10):107. doi:10.1890/ES11-00103.1

  18. Encalada AC, Peckarsky BL (2006) Selective oviposition behavior of the mayfly Baetis bicaudatus. Oecologia 148:526–537

  19. Encalada AC, Peckarsky BL (2007) A comparative study of the costs of alternative mayfly oviposition behavior. Behav Ecol Sociobiol 61:1437–1448

  20. Flecker A, Allan JD (1988) Flight direction in some Rocky Mountain mayflies (Ephemeroptera) with observation of parasitism. Aquat Insect 10:33–42

  21. Fonseca DM, Hart DD (2001) Colonization history masks habitat preferences in local distributions of stream insects. Ecology 82:2897–2910

  22. Forrester GE (1995) Strong density dependent survival and recruitment regulate the abundance of a coral-reef fish. Oecologia 103:275–282

  23. Hairston NG, Braner M, Twombly S (1987) Perspective on prospective methods for obtaining life table data. Limnol Oceanogr 32:517–520

  24. Hildrew AG, Woodward G, Winterbottom JH, Orton S (2004) Strong density dependence in a predatory insect: large-scale experiment in a stream. J Anim Ecol 73:448–458

  25. Hixon MA, Pacala SW, Sandin SA (2002) Population regulation: Historical context and contemporary challenges of open vs. closed systems. Ecology 83:1490–1508

  26. Holbrook SJ, Forrester GE, Schmitt RJ (2000) Spatial patterns in abundance of a damselfish reflect availability of suitable habitat. Oecologia 122:109–120

  27. Hughes TP (1990) Recruitment limitation, mortality, and population regulation in open systems: a case study. Ecology 71:12–20

  28. Humphries S (2002) Dispersal in drift-prone macroinvertebrates: a case for density-independence. Freshw Biol 47:921–929

  29. Jones GP (1990) The importance of recruitment to the dynamics of a coral reef fish population. Ecology 71:1691–1698

  30. Kohler SL (1992) Competition and the structure of a Benthic stream community. Ecol Monogr 62:165–188

  31. Kohler SL, McPeek MA (1989) Predation risk and the foraging behavior of competing stream insects. Ecology 70:1811–1825

  32. Lancaster J, Downes B, Arnold A (2010) Environmental constraints on oviposition limit egg supply of a stream insect at multiple scales. Oecologia 163:373–384

  33. McIntosh AR, Peckarsky BL, Taylor BW (2002) The influence of predatory fish on mayfly drift: extrapolating from experiments to nature. Freshw Biol 47:1497–1513

  34. McKnight D (2001) Freshwater ecosystems and climate change: recent assessments and recommendations. Limnol Oceanogr Bull 10:61–65

  35. Nakano S (1999) Terrestrial-aquatic linkages: riparian arthropod inputs alter trophic cascades in a stream food web. Ecology 80:2435–2441

  36. Palmer MA, Allan JD, Butman CA (1996) Dispersal as a regional process affecting the local dynamics of marine and stream benthic invertebrates. Trends Ecol Evol 11:322–326

  37. Peckarsky BL, Cowan CA, Penton MA, Anderson C (1993) Sub-lethal consequences of stream dwelling predatory stoneflies on mayfly growth and fecundity. Ecology 74:1836–1846

  38. Peckarsky BL, Taylor BW, Caudill CC (2000) Hydrologic and behavioral constraints on oviposition of stream insects: implications for adult dispersal. Oecologia 125:186–200

  39. Peckarsky BL, Taylor BW, McIntosh AR, McPeek MA, Lytle DA (2001) Variation in mayfly size at metamorphosis as a developmental response to risk of predation. Ecology 82:740–757

  40. Peckarsky BL, McIntosh AR, Taylor BW, Dahl J (2002) Predator chemicals induce changes in mayfly life history traits: a whole-stream manipulation. Ecology 83:612–618

  41. Peckarsky BL, Kerans BL, Taylor BW, McIntosh AR (2008) Predator effects on prey population dynamics in open systems. Oecologia 156:431–440

  42. Pfister C (1996) The role and importance of recruitment variability to a guild of tide pool fishes. Ecology 77:1928–1941

  43. Poff NL, Tokar S, Johnson P (1996) Stream hydrological and ecological responses to climate change assessed with an artificial neural network. Limnol Oceanogr 41:857–863

  44. Price PW, Ohgushi T (1995) Preference and performance linkage in a Phyllocolpa sawfly on the willow, Salix miyabeana, on Hokkaido. Res Popul Ecol 37:23–28

  45. Reich P, Downes BJ (2004) Relating larval distributions to patterns of oviposition: evidence from lotic hydrobiosid caddisflies. Freshw Biol 49:1423–1436

  46. Resetarits WJ, Bernardo J (eds) (1998) Experimental ecology. Oxford University Press, Oxford

  47. SAS Institute (2003) User’s guide for SAS software navigator. SAS Institute, Cary

  48. Schmitt RJ, Holbrook SJ (2000) Habitat-limited recruitment of coral reef damselfish. Ecology 81:3479–3494

  49. Spencer M, Blaustein L, Cohen JE (2002) Oviposition habitat selection by mosquitoes (Culiseta longiareolata) and consequences for population size. Ecology 83:669–679

  50. Taylor BW, McIntosh AR, Peckarsky BL (2001) Sampling stream invertebrates using electroshocking techniques: implications for basic and applied research. Can J Fish Aquat Sci 58:437–445

  51. Taylor BW, McIntosh AR, Peckarsky BL (2002) Reach-scale manipulations show invertebrate grazers depress algal resources in streams. Limnol Oceanogr 47:893–899

  52. Underwood AJ (1991) Beyond BACI: experimental designs for detecting human environmental impacts on temporal variations in natural populations. Aust J Mar Freshw Res 42:569–587

  53. Underwood AJ, Fairweather PG (1989) Supply side ecology and benthic marine assemblages. Trends Ecol Evol 4:16–20

  54. Vance SA, Peckarsky BL (1997) The effect of mermithid parasitism on predation of nymphal Baetis bicaudatus (Ephemeroptera) by invertebrates. Oecologia 110:147–152

  55. Wallace JB, Eggert SJ, Meyer JL, Webster JR (1997) Multiple trophic levels of a forest stream linked to terrestrial litter inputs. Science 277:102–104

  56. Wilcox A, Peckarsky BL, Taylor BW, Encalada AC (2008) Hydraulic and geomorphic effects on mayfly drift in high-gradient streams at moderate discharges. Ecohydrology 1:176–186

  57. Wilson JA, Osenberg CW (2002) Experimental and observational patterns of density-dependent settlement and survival in marine fish Gobiosoma. Oecologia 130:205–215

Download references


Special thanks for help in the field from Maruxa Álvarez, Ben Koch, Matt Harper, Wendy Brown, Brad Taylor, Tracy Smith, Juan Esteban Suárez-Encalada, Bryan Horn, Mark Wallin, and Alison Horn. We are grateful to Chris Fishel, Anthony Panzera, Elizabeth Kramer, Nathan Sanders, and Summer Rayne-Oakes for helping sort insect samples in the laboratory. This paper was improved by discussions with Nelson Hairston, Alex Flecker, Maruxa Álvarez, Brad Taylor, Angus McIntosh, Cole Gilbert, and Esteban Suárez, and by the insightful reviews of Barbara Downes and two anonymous reviewers. This research was supported by funds from National Science Foundation (NSF) Doctoral Dissertation Improvement Grant DEB-0206095 to ACE and from NSF grant DEB-0089863 and HATCH funds to BLP. ACE was also supported by the Ecuadorian Foundation for Science and Technology (FUNDACYT).

Author information

Correspondence to Andrea C. Encalada.

Additional information

Communicated by Barbara Downes.

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Encalada, A.C., Peckarsky, B.L. Large-scale manipulation of mayfly recruitment affects population size. Oecologia 168, 967–976 (2012). https://doi.org/10.1007/s00442-011-2147-1

Download citation


  • Ephemeroptera
  • Large-scale experiment
  • Life histories
  • Oviposition behavior
  • Recruitment limitation
  • Streams