Biodiversity and Conservation

, 18:3909 | Cite as

Richness gradients of stream invertebrates across the USA: taxonomy- and trait-based approaches

  • Leah A. Bêche
  • Bernhard Statzner
Original Paper


Large-scale diversity patterns in relationship to environmental factors at multiple spatial scales have been well-studied for many taxonomic groups; however, freshwater ecosystems remain understudied. Biodiversity is now widely recognized to encompass many more factors than just species numbers, particularly the inclusion of functional attributes. In this study, we examined richness patterns of stream invertebrate genera and their biological traits (“functional” richness) across 364 sites in the contiguous USA. In particular, we focused on the relationship between taxonomy- and trait-based richness to test for functional redundancy in stream communities. Further, we obtained environmental data to model the relative importance of local and watershed-scale environmental factors and residual spatial (latitude, longitude) influences on taxonomy- and trait-based richness. Trait richness increased linearly with genus richness (slope ≪ 1), although this appears to be an artifact of the restricted range of genus richness in our study (32 genera maximum). Furthermore, trait richness was significantly lower than expected under random community assembly. In contrast, the Ephemeroptera, Plecoptera, and Trichoptera (EPT) genera exhibited a saturating pattern between trait and genus richness and trait richness was no different from random. Our study indicates that there is functional redundancy among stream invertebrate genera, likely as a result of harsh habitat filters limiting trait diversity. Environmental factors (including spatially structured environmental factors) were always more important than spatial factors (latitude, longitude) in structuring richness despite strong longitudinal patterns of all richness measures (these differences were only significant for EPT genera). Finally, we found no significant difference in the relative importance of local and watershed scale environmental factors for taxonomy- and trait-based richness.


Diversity–function relationship Environmental filtering Functional diversity Land-use Redundancy Variance partitioning 



Aikake’s Information Criterion


Environmental Monitoring and Assessment Program


US Environmental Protection Agency


Ephemeroptera, Plectopera, Trichoptera


Geographic Information System


Standard deviation


Standard error


United States of America


Water Survey of America



We thank Alan Herlihy and Dave Peck for providing access to the WSA database. Two anonymous reviewers made valuable comments on the manuscript. This research was funded in part by a Marie Curie Incoming International Fellowship.


  1. Allan JD, Johnson LB (1997) Catchment-scale analysis of aquatic ecosystems. Freshw Biol 37:107–111CrossRefGoogle Scholar
  2. Barnes DKA, Griffiths HJ (2008) Biodiversity and biogeography of southern temperate and polar bryozoans. Glob Ecol Biogeogr 17:84–99Google Scholar
  3. Baselga A (2008) Determinants of species richness, endemism and turnover in European longhorn beetles. Ecography 31:263–271CrossRefGoogle Scholar
  4. Bêche LA, Resh VH (2007) Biological traits of benthic macroinvertebrates in California mediterranean-climate streams: long-term annual variability and trait diversity patterns. Fundam Appl Limnol 169:1–23CrossRefGoogle Scholar
  5. Bêche LA, McElravy EP, Resh VH (2006) Long-term seasonal variation in the biological traits of benthic-macroinvertebrates in two mediterranean-climate streams in California, USA. Freshw Biol 51:56–75CrossRefGoogle Scholar
  6. Blanchet FG, Legendre P, Borcard D (2008) Forward selection of explanatory variables. Ecology 89:2623–2632CrossRefPubMedGoogle Scholar
  7. Bonada N, Prat N, Resh VH, Statzner B (2006) Developments in aquatic insect biomonitoring: a comparative analysis of recent approaches. Annu Rev Entomol 51:495–523CrossRefPubMedGoogle Scholar
  8. Bonada N, Dolédec S, Statzner B (2007) Taxonomic and biological trait differences of stream macroinvertebrate communities between Mediterranean and temperate regions: implications for future climatic scenarios. Glob Chang Biol 13:1658–1671CrossRefGoogle Scholar
  9. Borcard D, Legendre P, Drapeau P (1992) Partialling out the spatial component of ecological variation. Ecology 73:1045–1055CrossRefGoogle Scholar
  10. Botta-Dukát Z (2005) Rao’s quadratic entropy as a measure of functional diversitybased on multiple traits. J Veg Sci 16:533–540CrossRefGoogle Scholar
  11. Boulton AJ, Boyero L, Covich AP (2008) Are tropical streams ecologically different from temperate streams? In: Dudgeon D et al (eds) Tropical stream ecology. Elsevier, AmsterdamGoogle Scholar
  12. Brosse S, Arbuckle CJ, Townsend CR (2003) Habitat scale and biodiversity: influence of catchment, stream reach and bedform scales on local invertebrate diversity. Biodivers Conserv 12:2057–2075CrossRefGoogle Scholar
  13. Carnicer J, Diaz-Delgado R (2008) Geographic differences between functional groups in patterns of bird species richness in North America. Acta Oecologica 33:253–264CrossRefGoogle Scholar
  14. Chapin FS III, Zavaleta ES, Eviner VT et al (2000) Consequences of changing biodiversity. Nature 405:234–242CrossRefPubMedGoogle Scholar
  15. Chessel D, Dufour A-B, Thioulouse J (2004) The ade4 package-I: one-table methods. R News 4:5–10Google Scholar
  16. Chevenet F, Dolédec S, Chessel D (1994) A fuzzy-coding approach for the analysis of long-term ecological data. Freshw Biol 31:295–309CrossRefGoogle Scholar
  17. Climate Atlas of the United States (2002) version 2.0 (online). Accessed 20 September 2008 at
  18. Compin A, Céréghino R (2007) Spatial patterns of macroinvertebrate functional feeding groups in streams in relation to physical variables and land-cover in southwestern France. Landsc Ecol 22:1215–1225CrossRefGoogle Scholar
  19. Díaz S, Cabido M (2001) Vive la différence: plant functional diversity matters to ecosystem processes. Trends Ecol Evol 16:646–655CrossRefGoogle Scholar
  20. Díaz S, Symstad AJ, Stuart Chapin F III et al (2003) Functional diversity revealed by removal experiments. Trends Ecol Evol 18:140–146CrossRefGoogle Scholar
  21. Didham RK, Watts CH, Norton DA (2005) Are systems with strong underlying abiotic regimes more likely to exhibit alternative stable states? Oikos 110:409–416CrossRefGoogle Scholar
  22. Dray S, Legendre P, Peres-Neto PR (2006) Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecol Modell 196:483–493CrossRefGoogle Scholar
  23. Frissell CA, Liss WJ, Warren CE et al (1986) A hierarchical framework for stream habitat classification: viewing streams in a watershed context. Environ Manage 10:199–214CrossRefGoogle Scholar
  24. Gentry AH (1988) Changes in plant community diversity and floristic composition on environmental and geographical gradients. Ann Mo Bot Gard 75:1–34CrossRefGoogle Scholar
  25. Giller PS, Hillebrand H, Berninger U-G et al (2004) Biodiversity effects on ecosystem functioning: emerging issues and their experimental test in aquatic environments. Oikos 104:423–436CrossRefGoogle Scholar
  26. Hagen EM, Webster JR, Benfield EF (2006) Are leaf breakdown rates a useful measure of stream integrity along an agricultural land-use gradient? J North Am Benthol Soc 25:330–343CrossRefGoogle Scholar
  27. Hawkins BA (2004) Summer vegetation, deglaciation and the anomalous bird diversity gradient in eastern North America. Ecography 13:321–325Google Scholar
  28. Hawkins BA, Diniz-Filho JAF (2004) ‘Latitude’ and geographic patterns in species richness. Ecography 27:268–272CrossRefGoogle Scholar
  29. Hawkins BA, Porter EE, Diniz-Filho JAF (2003) Productivity and history as descriptors of the latitudinal diversity gradient for terrestrial birds. Ecology 84:1608–1623CrossRefGoogle Scholar
  30. Heino J (2005) Functional biodiversity of macroinvertebrate assemblages along major ecological gradients of boreal headwater streams. Freshw Biol 50:1578–1587CrossRefGoogle Scholar
  31. Heino J, Mykrä H (2008) Control of stream insect assemblages: roles of spatial configuration and local environmental factors. Ecol Entomol 33:614–622CrossRefGoogle Scholar
  32. Heino J, Muotka T, Paavola R (2003) Determinants of macroinvertebrate diversity in headwater streams: regional and local influences. J Anim Ecol 72:425–434CrossRefGoogle Scholar
  33. Heino J, Mykrä H, Kotanen J, Muotka T (2007) Ecological filters and variability in stream macroinvertebrate communities: do taxonomic and functional structure follow the same path? Ecography 30:217–230Google Scholar
  34. Heino J, Mykrä H, Kotanen J (2008) Weak relationships between landscape characteristics and multiple facets of stream macroinvertebrate biodiversity in a boreal drainage system. Landsc Ecol 23:417–426CrossRefGoogle Scholar
  35. Hérault B (2007) Reconciling niche and neutrality through the Emergent Group approach. Perspect Plant Ecol Evol Syst 9:71–78CrossRefGoogle Scholar
  36. Hof C, Braendle M, Brandl R (2008) Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types. Glob Ecol Biogeogr 17:539–546CrossRefGoogle Scholar
  37. Hooper DU, Chapin FS, Ewel JJ et al (2005) Effects of biodiversity on ecosystem functioning: a consensus of current knowledge. Ecol Monogr 75:3–35CrossRefGoogle Scholar
  38. Hubbell SP (2001) The unified neutral theory of biodiversity and biogeography. Princeton University Press, PrincetonGoogle Scholar
  39. Hurlbert SH (1971) The nonconcept of species diversity: a critique and alternative parameters. Ecology 52:577–586CrossRefGoogle Scholar
  40. Jetz W, Rahbek C (2001) Geometric constraints explain much of the species richness pattern in African birds. Proc Natl Acad Sci USA 98:5661–5666CrossRefPubMedGoogle Scholar
  41. Kaufman DM, Willig MR (1998) Latitudinal patterns of mammalian species richness in the New World: the effects of sampling method and faunal group. J Biogeogr 25:795–805CrossRefGoogle Scholar
  42. Kearns FR, Kelly NM, Carter JL et al (2005) A method for the use of landscape metrics in freshwater research and management. Landsc Ecol 20:113–125CrossRefGoogle Scholar
  43. Klemm DJ, Blocksom KA, Fulk FA et al (2003) Development and evaluation of a macroinvertebrate biotic integrity index (MBII) for regionally assessing mid-Atlantic highlands streams. Environ Manage 31:656–669CrossRefPubMedGoogle Scholar
  44. Lavorel S, Grigulis K, McIntyre S et al (2008) Assessing functional diversity in the field–methodology matters! Funct Ecol 22:134–147Google Scholar
  45. Legendre P, Legendre L (1998) Numerical ecology, 2nd edn. Elsevier, AmsterdamGoogle Scholar
  46. Lemmon PE (1957) A new instrument for measuring forest overstory density. J For 55:667–669Google Scholar
  47. Lenat DR, Resh VH (2001) Taxonomy and stream ecology—the benefits of genus- and species-level identifications. J North Am Benthol Soc 20:287–298CrossRefGoogle Scholar
  48. Loreau M (2000) Biodiversity and ecosystem functioning: recent theoretical advances. Oikos 91:3–17CrossRefGoogle Scholar
  49. Loreau M (2004) Does functional redundancy exist? Oikos 104:606–611CrossRefGoogle Scholar
  50. Maloney KO, Feminella JW, Mitchell RM et al (2008) Land-use legacies and small streams: identifying relationships between historical land-use and contemporary stream conditions. J North Am Benthol Soc 27:280–294CrossRefGoogle Scholar
  51. McCreadie JW, Adler PH, Hamada N (2005) Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions. Ecol Entomol 30:201–209CrossRefGoogle Scholar
  52. McGill BJ, Enquist BJ, Weiher E et al (2006) Rebuilding community ecology from functional traits. Trends Ecol Evol 21:178–185CrossRefPubMedGoogle Scholar
  53. Mendez PK (2007) Life history of benthic macroinvertebrates: studies and applications to freshwater ecology. Ph.D. Thesis, University of California, BerkeleyGoogle Scholar
  54. Merritt RW, Cummins KW, Berg MB (eds) (2008) An introduction to the aquatic insects of North America, 4th edn. Kendall Hunt Publishers, DubuqueGoogle Scholar
  55. Micheli F, Halpern BS (2005) Low functional redundancy in coastal marine assemblages. Ecol Lett 8:391–400CrossRefGoogle Scholar
  56. Mykrä H, Heino J, Muotka T (2007) Scale-related patterns in the spatial and environmental components of stream macroinvertebrate assemblage variation. Glob Ecol Biogeogr 16:149–159CrossRefGoogle Scholar
  57. Naeem S (1998) Species redundancy and ecosystem reliability. Conserv Biol 12:39–45CrossRefGoogle Scholar
  58. Naeem S, Wright JP (2003) Disentangling biodiversity effects on ecosystem functioning: deriving solutions to a seemingly insurmountable problem. Ecol Lett 6:567–579CrossRefGoogle Scholar
  59. Ni J (2003) Plant functional types and climate along a precipitation gradient in temperate grasslands, north-east China and south-east Mongolia. J Arid Environ 53:501–516CrossRefGoogle Scholar
  60. Oberdorff T, Guegan J-F, Hugueny B (1995) Global scale patterns of fish species richness in rivers. Ecography 18:345–352CrossRefGoogle Scholar
  61. Oksanen J, Kindt R, Legendre P et al (2008) Vegan: community ecology package. R package version 1.13-1Google Scholar
  62. Peck DV, Herlihy AT, Hill BH et al (2006) Environmental Monitoring and Assessment Program—Surface Waters Western Pilot Study: field operations manual for wadeable streams. EPA/620/R-06/003. Office of Research and Development, USA Environmental Protection Agency, Washington, DCGoogle Scholar
  63. Peres-Neto PR, Legendre P, Dray S et al (2006) Variation partitioning of species data matrices: estimation and comparison of fractions. Ecology 87:2614–2625CrossRefPubMedGoogle Scholar
  64. Petchey OL, Gaston KJ (2006) Functional diversity: back to basics and looking forward. Ecol Lett 9:741–758CrossRefPubMedGoogle Scholar
  65. Petchey OL, Evans KL, Fishburn IS et al (2007) Low functional diversity and no redundancy in British avian assemblages. J Anim Ecol 76:977–985CrossRefPubMedGoogle Scholar
  66. Poff NL (1997) Landscape filters and species traits: towards mechanistic understanding and prediction in stream ecology. J North Am Benthol Soc 16:391–409CrossRefGoogle Scholar
  67. Qian H (1998) Large-scale biogeographic patterns of vascular plant richness in North America: an analysis at the generic level. J Biogeogr 25:829–836CrossRefGoogle Scholar
  68. Qian H (1999) Spatial pattern of vascular plant diversity in North America north of Mexico and its floristic relationship with Eurasia. Ann Bot 83:271–283CrossRefGoogle Scholar
  69. Qian H, Klinka K, Kayahara GJ (1998) Longitudinal patterns of plant diversity in the North American boreal forest. Plant Ecol 138:161–178CrossRefGoogle Scholar
  70. Qian H, Wang S, He J-S et al (2006) Phytogeographic analysis of seed plant genera in China. Ann Bot 98:1073–1084CrossRefPubMedGoogle Scholar
  71. R Development Core Team (2008) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, URL
  72. Rahbek C (2005) The role of spatial scale and the perception of large-scale species-richness patterns. Ecol Lett 8:224–239CrossRefGoogle Scholar
  73. Resetarits WJ Jr, Chalcraft DR (2007) Functional diversity within a morphologically conservative genus of predators: implications for functional equivalence and redundancy in ecological communities. Funct Ecol 21:793–804CrossRefGoogle Scholar
  74. Reyjol Y, Hugueny B, Pont D et al (2007) Patterns in species richness and endemism of European freshwater fish. Glob Ecol Biogeogr 16:65–75CrossRefGoogle Scholar
  75. Ribera I, Foster GN, Vogler AP (2003) Does habitat use explain large scale species richness patterns of aquatic beetles in Europe? Ecography 26:145–152CrossRefGoogle Scholar
  76. Ricklefs RE (2004) A comprehensive framework for global patterns in biodiversity. Ecol Lett 7:1–15CrossRefGoogle Scholar
  77. Ricotta C (2005) A note on functional diversity measures. Basic Appl Ecol 6:479–486CrossRefGoogle Scholar
  78. Sandin L, Johnson RK (2004) Local, landscape and regional factors structuring benthic macroinvertebrate assemblages in Swedish streams. Landsc Ecol 19:501–514CrossRefGoogle Scholar
  79. Scheffer M, van Nes EH (2006) Self-organized similarity, the evolutionary emergence of groups of similar species. Proc Nat Acad Sci USA 103:6230–6235CrossRefPubMedGoogle Scholar
  80. Schwartz MW, Brigham CA, Hoeksema JD et al (2000) Linking biodiversity to ecological function: implications for conservation ecology. Oecologia 122:297–305CrossRefGoogle Scholar
  81. Sponseller RA, Benfield EF, Valett HM (2001) Relationships between land-use, spatial scale and stream macroinvertebrate communities. Freshw Biol 46:1409–1424CrossRefGoogle Scholar
  82. Statzner B, Moss B (2004) Linking ecological function, biodiversity and habitat: a mini-review focusing on older ecological literature. Basic Appl Ecol 5:97–106CrossRefGoogle Scholar
  83. Statzner B, Hildrew AG, Resh VH (2001) Species traits and environmental constraints: entomological research and the history of ecological theory. Annu Rev Entomol 46:291–316CrossRefPubMedGoogle Scholar
  84. Statzner B, Dolédec S, Hugueny B (2004) Biological trait composition of European stream invertebrate communities: assessing the effects of various trait filter types. Ecography 27:470–488CrossRefGoogle Scholar
  85. Statzner B, Bonada N, Dolédec S (2007) Conservation of taxonomic and biological trait diversity of European stream macroinvertebrate communities: a case for a collective public database. Biodivers Conserv 16:3609–3632CrossRefGoogle Scholar
  86. Stevens RD, Cox SB, Strauss RE et al (2003) Patterns of functional diversity across an extensive environmental gradient: vertebrate consumers, hidden treatments and latitudinal trends. Ecol Lett 6:1099–1108CrossRefGoogle Scholar
  87. Stewart KW, Stark BP (2002) Nymphs of North American stonefly genera, 2nd edn. The Caddis Press, ColumbusGoogle Scholar
  88. Suding KN, Lavorel S, Chapin FSIII et al (2008) Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants. Glob Chang Biol 14:1125–1140CrossRefGoogle Scholar
  89. Thompson R, Townsend C (2006) A truce with neutral theory: local deterministic factors, species traits and dispersal limitation together determine patterns of diversity in stream invertebrates. J Anim Ecol 75:476–484CrossRefPubMedGoogle Scholar
  90. Thorp JH, Covich AP (eds) (2001) Ecology and classification of North American freshwater invertebrates, 2nd edn. Academic Press, LondonGoogle Scholar
  91. USEPA (2004) Wadeable Streams Assessment: field operations manual. EPA841-B-04-004. U.S. Environmental Protection Agency, Office of Water and Office of Research and Development, Washington, DCGoogle Scholar
  92. USEPA (2006) Wadeable Streams Assessment: a collaborative survey of the nation’s streams, EPA 841-B-06-002. U.S. Environmental Protection Agency, Office of Water and Office of Research and Development, Washington, DCGoogle Scholar
  93. Usseglio-Polatera P, Bournaud M, Richoux P et al (2000) Biological and ecological traits of benthic freshwater macroinvertebrates: relationships and definition of groups with similar traits. Freshw Biol 43:175–205CrossRefGoogle Scholar
  94. Vieira NKM, Poff NL, Carlisle DM et al (2006) A database of lotic invertebrate traits for North America. USGS, Reston Virginia. Available at
  95. Vinson MR, Hawkins CP (1998) Biodiversity of stream insects: variation at local, basin, and regional scales. Annu Rev Entomol 43:271–293CrossRefPubMedGoogle Scholar
  96. Vinson MR, Hawkins CP (2003) Broad-scale geographical patterns in local stream insect genera richness. Ecography 26:751–767CrossRefGoogle Scholar
  97. Waite IR, Herlihy AT, Larsen DP et al (2000) Comparing strengths of geographic and nongeographic classifications of stream benthic macroinvertebrates in the Mid-Atlantic Highlands, USA. J N Am Benth Soc 19:429–441CrossRefGoogle Scholar
  98. Walker BH (1992) Biodiversity and ecological redundancy. Conserv Biol 6:18–23CrossRefGoogle Scholar
  99. Walker BH, Kinzig A, Langridge J (1992) Plant attribute diversity, resilience, and ecosystem function: the nature and significance of dominant and minor species. Ecosystems 2:95–113CrossRefGoogle Scholar
  100. Wallace JB, Webster JR (1996) The role of macroinvertebrates in stream ecosystem function. Annu Rev Entomol 41:115–139CrossRefPubMedGoogle Scholar
  101. Waters TF (1995) Sediment in streams: sources, biological effects, and controls. American Fisheries Society Monograph 7, Bethesda, MarylandGoogle Scholar
  102. Weiher E, Keddy PA (1995) Assembly rules, null models, and trait dispersion: new questions from old patterns. Oikos 74:159–164CrossRefGoogle Scholar
  103. Wiggins GB (2004) Caddisflies: the underwater architects. University of Toronto Press, TorontoGoogle Scholar
  104. Willig MR, Kaufman DM, Stevens RD (2003) Latitudinal gradients of biodiversity: pattern, process, scale, and synthesis. Annu Rev Ecol Syst 34:273–309CrossRefGoogle Scholar
  105. Woodward G, Papantoniou G, Edwards F et al (2008) Trophic trickles and cascades in a complex food web: impacts of a keystone predator on stream community structure and ecosystem processes. Oikos 117:683–692CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2009

Authors and Affiliations

  1. 1.CNRS-UMR 5023 Écologie des Hydrosystèmes FluviauxUniversité de LyonVilleurbanne CedexFrance
  2. 2.CNRS—Biodiversité des Écosystemes LotiquesParcieuxFrance

Personalised recommendations