Patterns of diversity, depth range and body size among pelagic fishes along a gradient of depth

Studies of geographical patterns of diversity have focused largely on compiling and analysing data to evaluate alternative hypotheses for the near-universal decrease in species richness from the equator to the poles. Valuable insights into the mechanisms that promote diversity can come from studies of other patterns, such as variation in species distributions with elevation in terrestrial systems or with depth in marine systems. To obtain such insights, we analysed and interpreted data on species diversity, depth of occurrence and body size of pelagic fishes along an oceanic depth gradient. We used a database on pelagic marine fishes native to the north-east Pacific Ocean between 40°N and 50°N. We used data from the Pacific Rim Fisheries Program that were obtained from commercial, management and scientific surveys between 1999 and 2000. Depth of occurrence and maximum body length were used to assess the distributions of 409 species of pelagic fishes along a depth gradient from 0 to 8000 m. A presence‐absence matrix was used to classify the depth range of each species into 100-m intervals. Atmar & Patterson’s (1995) software was used to quantify the degree of nestedness of species distributions. Pelagic fish species diversity decreased steeply with increasing depth; diversity peaked at less than 200 m and more than half of the species had mean depths of occurrence between 0 and 300 m. The distribution of species showed a very strong nested subset pattern along the depth gradient. Whereas species with narrow ranges were generally restricted to shallow waters, wide-ranging species occurred from near the surface to great depths. The relationship between maximum body size and mean depth range differed between teleost and elasmobranch fishes: being positive for teleosts, but negative for elasmobranches. Results support hypotheses that some combination of high productivity and warm temperature promote high species diversity, and reject those that would attribute the pattern of species richness to the mid-domain effect, habitat area, or environmental constancy. The data provided a clear example of Rapoport’s rule, a negative correlation between average depth range and species diversity.

[1]  V. Sánchez‐Cordero Elevation gradients of diversity for rodents and bats in Oaxaca, Mexico , 2001 .

[2]  M. Rex,et al.  Global-scale latitudinal patterns of species diversity in the deep-sea benthos , 1993, Nature.

[3]  K. Gaston,et al.  Animal body size distributions: patterns, mechanisms and implications. , 1994, Trends in ecology & evolution.

[4]  Mark V. Lomolino,et al.  Frontispiece: Diversity patterns of small mammals along elevational gradients , 2001 .

[5]  D. Currie Energy and Large-Scale Patterns of Animal- and Plant-Species Richness , 1991, The American Naturalist.

[6]  E. Leigh Community diversity and environmental stability: A re-examination. , 1990, Trends in ecology & evolution.

[7]  M. Rex,et al.  Marine Biodiversity: Large-scale patterns of species diversity in the deep-sea benthos , 1997 .

[8]  L. Heaney Small mammal diversity along elevational gradients in the Philippines: an assessment of patterns and hypotheses , 2001 .

[9]  H. L. Sanders Benthic marine diversity and the stability-time hypothesis. , 1969, Brookhaven symposia in biology.

[10]  K. Rohde,et al.  Latitudinal Ranges of Teleost Fish in the Atlantic and Indo-Pacific Oceans , 1996, The American Naturalist.

[11]  George L. Pickard,et al.  Descriptive Physical Oceanography: An Introduction , 1963 .

[12]  Jon C. Lovett,et al.  Biodiversity and environmental stability , 1997, Biodiversity & Conservation.

[13]  H. L. Sanders,et al.  Marine Benthic Diversity: A Comparative Study , 1968, The American Naturalist.

[14]  Wirt Atmar,et al.  Nested subsets and the structure of insular mammalian faunas and archipelagos , 1986 .

[15]  W. Pearcy,et al.  Species Associations of Benthic Fishes on the Continental Shelf and Slope off Oregon , 1968 .

[16]  K. Roy,et al.  Latitudinal distribution of body size in north‐eastern Pacific marine bivalves , 2001 .

[17]  P. Jensen Nematode assemblages in the deep-sea benthos of the Norwegian Sea , 1988 .

[18]  C. Duarte,et al.  Patterns in species richness, size, and latitudinal range of East Atlantic fishes , 1994 .

[19]  G. C. Stevens Extending Rapoport's rule to Pacific marine fishes , 1996 .

[20]  M. P. M. Reddy,et al.  Descriptive physical oceanography , 2001 .

[21]  M. Rosenzweig Species Diversity Gradients: We Know More and Less Than We Thought , 1992 .

[22]  J. Terborgh On the Notion of Favorableness in Plant Ecology , 1973, The American Naturalist.

[23]  W. Prell,et al.  Environmental controls on the geographic distribution of zooplankton diversity , 1999, Nature.

[24]  O. Pfannkuche and abyssal plain (NE Atlantic): population structure, distribution, standing stocks , 1985 .

[25]  G. C. Stevens The Elevational Gradient in Altitudinal Range: An Extension of Rapoport's Latitudinal Rule to Altitude , 1992, The American Naturalist.

[26]  Timothy R. Parsons,et al.  Biological oceanography: an introduction. Second edition , 1997 .

[27]  A. Longhurst Ecological Geography of the Sea , 1998 .

[28]  K. Rohde Latitudinal gradients in species diversity: the search for the primary cause , 1992 .

[29]  D. Jablonski,et al.  Invariant size-frequency distributions along a latitudinal gradient in marine bivalves. , 2000, Proceedings of the National Academy of Sciences of the United States of America.

[30]  Bruce D. Patterson,et al.  Distribution of bats along an elevational gradient in the Andes of south‐eastern Peru , 1996 .

[31]  G. C. Stevens The Latitudinal Gradient in Geographical Range: How so Many Species Coexist in the Tropics , 1989, The American Naturalist.

[32]  Mark V. Lomolino,et al.  Elevation gradients of species‐density: historical and prospective views , 2001 .

[33]  George C. Hurtt,et al.  Nonbiological Gradients in Species Richness and a Spurious Rapoport Effect , 1994, The American Naturalist.

[34]  James H. Brown Mammals on mountainsides : elevational patterns of diversity , 2001 .

[35]  Colwell,et al.  The mid-domain effect: geometric constraints on the geography of species richness. , 2000, Trends in ecology & evolution.

[36]  Robert M. May,et al.  The Search for Patterns in the Balance of Nature: Advances and Retreats , 1986 .

[37]  T. Fujita,et al.  Size structure of dense populations of the brittle star Ophiura sarsii (Ophiuroidea: Echinodermata) in the bathyal zone around Japan , 1990 .

[38]  M. Rex,et al.  Bathymetric patterns of body size: implications for deep-sea biodiversity , 1998 .