, Volume 61, Issue 1, pp 32–41 | Cite as

Species size distributions in marine benthic communities

  • R. M. Warwick
  • K. R. Clarke
Original Papers


Species body size distributions from eight temperate benthic communities show a highly conservative pattern with two separate lognormal distributions, corresponding to the traditional categories of meiofauna and macrofauna. The meiofaunal mode occurs at a dry body weight of 0.64 μg and the macrofaunal mode at 3.2 mg, with a trough between them at 45 μg. It is suggested that there is a particular body size at which meiofaunal life-history and feeding traits can be optimised, and another for macrofaunal traits. As size departs in either direction (larger or smaller) from these optima, fewer species of the same size are able to co-exist. The split occurs at 45 μg because many life history and feeding characteristics switch more or less abruptly at about this body size, compromise traits being either non-viable or disadvantageous. Meiofauna and macrofauna therefore comprise two separate evolutionary units each with an internally coherent set of biological characteristics.

The expression of this conservative pattern is modified by water depth: the proportion of macrofauna species increases from intertidal situations to deeper water, and it is suggested that mechanisms of resource partitioning and diversity maintainence in the meiofauna and macrofauna are affected differentially by sediment disturbance. Salinity does not affect this proportionality, and so does not differentially affect mechanisms for maintaining species diversity in any particular size category of animals. Meiofauna species size distributions may be modified in sandy sediments because of physical impositions on interstitial or burrowing lifestyles.

Brief discussion of some implications of these observations includes speculations on the larval ecology of macrofauna, on gigantism in Antarctic invertebrates, and on the benthic Sheldon spectrum.


  1. Arnaud PM (1977) Adaptations within the Antarctic marine benthic ecosystem. In: Llano GA (ed) Adaptations within Antarctic ecosystems. Smithsonian Institution, Washington DC, pp 135–157Google Scholar
  2. Banse K (1982) Mass-scaled rates of respiration and intrinsic growth in very small invertebrates. Mar Ecol Prog Ser 9:281–297Google Scholar
  3. Banse K, Mosher S (1980) Adult body mass and annual production/biomass relationships of field populations. Ecol Monogr 50:355–379Google Scholar
  4. Buchanan JB, Warwick RM (1974) An estimate of benthic macrofauna production in the offshore mud of the Northumberland coast. J mar biol Ass UK 54:197–222Google Scholar
  5. Caswell H (1981) The evolution of “mixed” life histories in marine invertebrates and elsewhere. Am Nat 117:529–536Google Scholar
  6. De Broyer C (1977) Analysis of the gigantism and dwarfness of Antarctic and Subantarctic gammaridean Amphipoda. In: Llano GA (ed) Adaptations within Antarctic ecosystems. Smithsonian Institution, Washington DC, pp 327–334Google Scholar
  7. Desbruyères D (1977) Benthic bionomy of the Continental Shelf of the Kerguelen Archipelago. Macrofaune 2. Diversity of benthic annelid population in a fjord close to the Morbihan Gulf. In: Llano GA (ed) Adaptations within Antarctic ecosystems. Smithsonian Institution. Washington DC, pp 227–238Google Scholar
  8. Gerlach SA (1977) Means of meiofauna dispersal. Microfauna Meeresboden 61:89–103Google Scholar
  9. Gray JS (1981) The ecology of marine sediments. Cambridge, Cambridge University PressGoogle Scholar
  10. Holme NA, McIntyre AD (in press) Methods for the study of marine benthos (2nd edition). Blackwell, OxfordGoogle Scholar
  11. Hutchinson GE (1959) Homage to Santa Rosalia, or why are there so many kinds of animals. Am Nat 93:145–159Google Scholar
  12. Hutchinson GE, MacArthur RH (1959) A theoretical ecological model of size distributions among species of animals. Am Nat 93:117–125Google Scholar
  13. Joint IR, Gee JM, Warwick RM (1982) Determination of fine-scale vertical distribution of microbes and meiofauna in an intertidal sediment. Mar Biol 72:157–164Google Scholar
  14. Kerr SR (1974) Theory of size distribution in ecological communities. J Fish Res Bd Can 31:1859–1862Google Scholar
  15. Kerr SR (1979) Prey availability, metaphoetesis, and the size structure of lake trout stocks. Inv Pesq 43:187–198Google Scholar
  16. May RM (1975) Patterns of species abundance and diversity. In: Cody ML, Diamond JM (eds) Ecology and evolution of communities. Belknap, Cambridge, Mass, pp 81–120Google Scholar
  17. Morris AW, Bale AJ, Howland RJM (1982) Chemical variability in the Tamar estuary. Est cstl Shelf Sci 14:649–661Google Scholar
  18. Pechenik JA (1979) Role of encapsulation in invertabrate life histories. Am Nat 114:859–870Google Scholar
  19. Picken GB (1980) Reproductive adaptations of Antarctic benthic invertebrates. Biol J Linn Soc 14:67–75Google Scholar
  20. Platt T, Denman K (1977) Organization in the pelagic ecosystem. Helgolander wiss Meeresunters 30:575–581Google Scholar
  21. Preston FW (1948) The commonness and rarity of species. Ecology 29:254–283Google Scholar
  22. Price R, Warwick RM (1980) Temporal variations in annual production and biomass in estuarine populations of two polychaetes, Nephtys hombergi and Ampharete acutifrons. J mar biol Ass UK 60:481–487Google Scholar
  23. Richardson MD, Hedgpeth JW (1977) Antarctic soft-bottom, macrobenthic community adaptations to a cold, stable, highly productive, glacially affected environment. In: Llano GA (ed) Adaptations within Antarctic ecosystems. Smithsonian Institution, Washington DC, pp 181–196Google Scholar
  24. Schoener TW (1974) Resource partitioning in ecological communities. Science 185:27–39Google Scholar
  25. Schwinghamer P (1981) Characteristic size distributions of integral benthic cummunities. Can J Fish Aquatic Sci 38:1255–1263Google Scholar
  26. Sheldon RW, Prakash A, Sutcliffe WH (1972) The size distribution of particles in the Ocean. Limnol Oceanogr 17:327–340Google Scholar
  27. Sheldon RW, Sutcliffe WH, Paranjape MA (1977) Structure of pelagic food chain and relationship between plankton and fish production. J Fish Res Bd Can 34:2344–2355Google Scholar
  28. Teare MJ (1978) An energy budget for Tachidius discipes (Copepoda, Harpacticoida) from an estuarine mud-flat. PhD Thesis, University of ExeterGoogle Scholar
  29. Theil H (1975) The size structure of the deep-sea benthos. Int Rev Ges Hydrobiol 60:575–606Google Scholar
  30. Thiery RG (1982) Environmental instability and community diversity. Biol Rev 57:671–710Google Scholar
  31. Thorson G (1946) Reproduction and larval development of Danish marine bottom invertebrates. Medd Komm Danm Fisk-og Havunders, ser Plankton 4:1–523Google Scholar
  32. Warwick RM (1981) Survival strategies of meiofauna. In: Jones NV, Wolff WJ (eds) Feeding and survival strategies of estuarine organisms Plenum, New York, pp 39–52Google Scholar
  33. Warwick RM (1982) The partitioning of secondary production among species in benthic communities. Neth J Sea Res 16:1–16Google Scholar
  34. Warwick RM, Buchanan JB (1970) The meiofauna off the coast of Northumberland. I. The structure of the nematode population. J mar biol Ass UK 50:129–146Google Scholar
  35. Warwick, Buchanan JB (1971) The meiofauna off the coast of Northumberland. II. Seasonal stability of the nematode population. J mar biol Ass UK 51:355–362Google Scholar
  36. Warwick RM, Price R (1975) Macrofauna production in an estuarine mud-flat. J mar biol Ass UK 55:1–18Google Scholar
  37. Warwick RM, Price R (1979) Ecological and metabolic studies on free-living nematodes from an estuarine mud-flat. Est cstl mar Sci 9:257–271Google Scholar
  38. Warwick RM, George CL, Davies JR (1978) Annual macrofauna production in a Venus community. Est cstl mar Sci 7:215–241Google Scholar
  39. Warwick RM, Joint IR, Radford PJ (1979) Secondary production of the benthos in an estuarine environment. In: Jefferies RL, Davey AJ (eds) Ecological processes in coastal environments. Blackwell, Oxford, pp 429–450Google Scholar
  40. Whitlatch RB (1980) Patterns of resource utilization and coexistence in marine intertidal deposit-feeding communities. J mar Res 38:743–765Google Scholar
  41. Whittaker RH (1970) Communities and ecosystems. MacMillan, New YorkGoogle Scholar
  42. Wieser W (1960) Benthic studies in Buzzards Bay. II. The meiofauna. Limnol Oceanogr 5:121–137Google Scholar


  1. Cox DR, Hinkley DV (1974) Theoretical statistics. Chapman and Hall, London, p 511Google Scholar
  2. Nash JC (1979) Compact numerical methods for computers. Adam Hilger, Bristol, p 227Google Scholar
  3. Silvey SD (1975) Statistical inference. Chapman and Hall, London, p 192Google Scholar

Copyright information

© Springer-Verlag 1984

Authors and Affiliations

  • R. M. Warwick
    • 1
  • K. R. Clarke
    • 1
  1. 1.Institute for Marine Environmental ResearchNERCPlymouthUK

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