Abstract
The more individuals hypothesis (MIH) postulates that productivity increases species richness by increasing mean equilibrium population size, thereby reducing the probability of local extinction. We tested the MIH for invertebrates colonizing microcosms that simulated tree holes by manipulating productivity through additions of leaf or animal detritus and subsequently determining the relationships among richness, total abundance, abundance per species, and measures of productivity. We quantified productivity as the rate of microorganism protein synthesis, microorganism metabolic rate, nutrient ion concentration, and type and amount of detritus. Microcosms with animal detritus attracted more species, more individuals per species, and more total individuals than did microcosms with similar amounts of leaf detritus. Relationships between richness or abundance and productivity varied with date. Richness in June increased as a linear function of productivity, whereas the power function predicted by the MIH fit best in July. Abundance in June and July was best described by a power function of productivity, but the linear function predicted by the MIH fit best in September. Abundance per species was best described by a power function of productivity in June and July. Path analysis showed that the indirect effect of productivity through abundance on richness that is predicted by MIH was important in all months, and that direct links between productivity and richness were unnecessary. Our results support many of the predictions of the MIH, but they also suggest that the effects of abundance on richness may be more complex than expected.
Similar content being viewed by others
References
Abrams PA (1995) Monotonic or unimodal diversity-productivity gradients: what does competition theory predict? Ecology 76:2019–2027
Anderson JM, Macfayden A (eds) (1976) The role of terrestrial and aquatic organisms in decomposition processes. Blackwell, Oxford
Barrera R (1988) Multiple factors and their interactions on structuring the community of aquatic insects of treeholes. PhD thesis, The Pennsylvania State University, University Park, Pa.
Carpenter SR (1983) Resource limitation of larval treehole mosquitoes subsisting on beech detritus. Ecology 64:219–223
Chase JM, Leibold MA (2002) Spatial scale dictates the productivity–biodiversity relationship. Science 416:427–430
Clements AN (1992) The biology of mosquitoes, vol 1. Chapman & Hall, London
Cloe WW III, Garman GC (1996) The energetic importance of terrestrial arthropod inputs to three warm-water streams. Freshw Biol 36:105–114
Evans KL, Warren PH, Gaston KJ (2005) Species–energy relationships at the macroecological scale: a review of the mechanisms. Biol Rev 80:1–25
Garman GC (1991) Use of terrestrial arthropod prey by a stream-dwelling cyprinid fish. Environ Biol Fishes 30:325–331
Garman GC (1992) Fate and potential significance of postspawning anadromous fish carcasses in an Atlantic costal river. Trans Am Fish Soc 121:390–394
Groner E, Novoplansky A (2003) Reconsidering diversity–productivity relationship: directness of productivity estimate matters. Ecol Lett 6:695–699
Hatcher L (1994) A step-by-step approach to using the SAS system for factor analysis and structural equation modeling. SAS Institute, Cary, N.C.
Hatcher L, Stepanski EJ (1994) A step-by-step approach to using the SAS system for univariate and multivariate statistics. SAS Institute, Cary, N.C.
Henschel JR, Mahsberg D, Stumpf H (2001) Allochthonous aquatic insects increase predation and decrease herbivory in river shore food webs. Oikos 93:429–438
Hurlbert AH (2004) Species–energy relationship and habitat complexity in bird communities. Ecol Lett 7:714–720
Hurlbert AH (2006) Linking species–area and species–energy relationships in Drosophila microcosms. Ecol Lett 9:287–294
Jenkins B, Kitching RL, Pimm SL (1992) Productivity, disturbance and food web structure at a local spatial scale in experimental container habitats. Oikos 65:249–255
Kaspari M, Yuan M, Alonso L (2003) Spatial grain and the causes of regional diversity gradients in ants. Am Nat 161:459–477
Kirchman DL (1993) Leucine incorporation as a measure of biomass production by heterotrophic bacteria. In: Kemp PL, Sherr BF, Sherr EB, Cole JJ (eds) Handbook of methods in aquatic microbiology. Lewis, Boca Raton, pp 509–512
Kitching RL (1971) An ecological study of water-filled tree-holes and their position in the woodland ecosystem. J Anim Ecol 40:281–302
Kitching RL (2000) Food webs and container habitats. The natural history and ecology of phytotelmata. Cambridge University Press, England
Kitching RL (2001) Food webs in phytotelmata: “bottom-up” and “top-down” explanations for community structure. Annu Rev Entomol 46:729–760
Kneitel JM, Miller TE (2002) Resource and top-predator regulation in the pitcher plant Sarracenia purpurea inquiline community. Ecology 83:680–688
Léonard PM, Juliano SA (1995) Effect of leaf litter and density on fitness and populations performance of the tree hole mosquito Aedes triseriatus. Ecol Entomol 20:125–136
Mason CF, MacDonald SM (1982) The input of terrestrial invertebrates from tree canopies to a stream. Freshw Biol 12:305–311
MacArthur RH, Wilson EO (1967) The theory of island biogeography. Harvard University Press, Cambridge
Merritt RW, Dadd RH, Walker ED (1992) Feeding behavior, natural food, and nutritional relationships of larval mosquitoes. Annu Rev Entomol 37:349–376
Mitchell RJ (2001) Path analysis: pollination. In: Schiner SM, Gurevitch J (eds) Design and analysis of ecological experiments. Oxford University Press, Oxford, pp 211–231
Mittelbach GG, Steiner CF, Scheiner SM, Gross KL, Reynolds HL, Waide RB, Willig MR, Dodson SI, Gough L (2001) What is the observed relationship between species richness and productivity? Ecology 82:2381–2396
Moore JC, Berlow EL, Coleman DC, Ruiter PC, Dong Q, Hastings A, Johnson NC, McCann KS, Melville K, Morin PJ, Nadelhoffer K, Rosemond AD, Post DM, Sabo JL, Scow KM, Vanni MJ, Wall DH (2004) Detritus, trophic dynamics and biodiversity. Ecol Lett 7:584–600
Nakano S, Miyasaka H, Kuhara N (1999) Terrestrial–aquatic linkages: riparian arthropod inputs alter trophic cascades in a stream food web. Ecology 80:2435–2441
O’Neill RV, Reichle DA (1980) Dimensions of ecosystem theory. In: Waring RH (ed) Forests: fresh perspectives from ecosystem analysis. Oregon State University Press, Corvallis, pp 11–26
Paradise CJ (2004) Relationship of water and leaf litter variability to insect inhabiting treeholes. J North Am Bentho Soc 23:793–805
Preston FW (1962) The canonical distribution of commonness and rarity. Ecology 43:185–215
Rosenzweig ML (1971) Paradox of enrichment: destabilization of exploitation ecosystems in ecological time. Science 171:385–387
Rosenzweig ML (1995) Species diversity in space and time. Cambridge University Press, Cambridge
Rosenzweig ML, Abramsky Z (1993) Species diversity gradients: we know more or less than we thought. J Mammal 73:715–730
SAS Institute (1990) SAS/STAT users guide, version 6, 4th edn, vol 1 and 2. SAS Institute, Cary, N.C.
Srivastava DS, Lawton JH (1998) Why more productive sites have more species: an experimental test of theory using tree-hole communities. Am Nat 152:510–529
Waide RB, Willig MR, Steiner CF, Mittelbach GG, Gough L, Dodson SI, Juday P, Parmenter R (1999) The relationship between productivity and species diversity. Annu Rev Ecol Syst 30:257–300
Walker ED, Lawson DL, Merritt RW, Morgan WT, Klug MJ (1991) Nutrient dynamics, bacterial populations, and mosquito productivity in tree hole ecosystems and microcosms. Ecology 72:1529–1546
Walker ED, Kaufman MG, Ayres MP, Riede MH, Merritt RW (1997) Effects of variation in quality of leaf detritus on growth of the eastern tree-hole mosquito, Aedes triseriatus (Diptera: Culicidae). Can J Zool 75:706–718
Weiher E (1999) The combined effects of scale and productivity on species richness. J Ecol 87:1005–1011
Wright DH (1983) Species–energy theory: an extension of species–area theory. Oikos 41:496–506
Yanoviak SP (2001) Predation, resource availability, and community structure in Neotropical water-filled tree holes. Oecologia 126:125–133
Yee DA (2006) Effects of species interactions and productivity on aquatic macroinvertebrate diversity and community composition in tree holes: patterns and mechanisms. PhD thesis, Illinois State University, Ill.
Yee DA, Juliano SA (2006) Consequences of detritus type in an aquatic microsystems: assessing water quality, micro-organisms, and the performance of the dominant consumer. Freshw Biol 51:448–459
Acknowledgments
Discussions with M.R. Willig regarding the MIH improved this study. We thank M. Kaufman for guidance with leucine methodologies. We thank S. Hohm, S. Harrell Yee, H. Les, K. Costanzo, B. Kesavaraju, and C. Villanueva who provided assistance in the field, three anonymous referees who provided insightful and helpful suggestions on an earlier version of this manuscript, The ParkLands Foundation for access to their property to conduct this experiment, and the Sakaluk lab, ISU, for providing us with crickets. This work was supported by grants to D.A. Yee from the ISU Department of Biological Sciences and the Phi Sigma Biological Society, and to S. A. Juliano and D. A. Yee from the National Institute of Allergy and Infectious Disease (R15 AI-051374).
Author information
Authors and Affiliations
Corresponding author
Additional information
Communicated by Nathan Sanders.
Rights and permissions
About this article
Cite this article
Yee, D.A., Juliano, S.A. Abundance matters: a field experiment testing the more individuals hypothesis for richness–productivity relationships. Oecologia 153, 153–162 (2007). https://doi.org/10.1007/s00442-007-0707-1
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00442-007-0707-1