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The birds and the seas: body size reconciles differences in the
abundance-occupancy relationship across marine and terrestrial
vertebrates
Thomas J. Webb, Nicholas K. Dulvy, Simon Jennings and Nicholas V.C. Polunin
T. J. Webb (t.j.webb@sheffield.ac.uk), Dept of Animal and Plant Sciences, Univ. of Sheffield, Sheffield, S10 2TN, UK. – N. K. Dulvy,
Biological Sciences, Simon Fraser Univ., Burnaby, BC, V5A 1S6, Canada. – S. Jennings, Centre for Environment, Fisheries and Aquaculture
Science, Pakefield Road, Lowestoft, NR33 0HT, UK and School of Environmental Sciences, Univ. of East Anglia, Norwich, NR4 7TJ, UK. –
N. V. C. Polunin, School of Marine Science and Technology, Newcastle Univ., NE1 7RU, UK.
Despite some fundamental differences in production processes and the ecology of consumer species on land and in the
sea, further understanding of pattern and process in both biomes might be gained by applying common methods of mac-
roecological analysis. We develop methods that reconcile apparent differences in abundance and occupancy for marine
and terrestrial vertebrates, as exemplified by fish and birds. hese recognize and take account of those aspects of the life
history and ecology of marine and terrestrial animals that influence their abundance, distribution and trophic role. When
abundance and occupancy are averaged within species over time we show that variation within a region is less for birds than
fish, but when abundance and occupancy are averaged over space, the difference between birds and fish disappears. Further,
we develop size rather than species-structured abundance-occupancy relationships for fish assemblages and demonstrate
that patterns of intra-size class variation that are very similar to intraspecific variation in bird species, over both time and
space. We argue that this result reflects the relative importance of body size and species identity respectively in determining
trophic roles in marine and terrestrial environments. Selection of the appropriate analytical unit on land (species) and in the
sea (size) helps to reconcile apparently divergent macroecological patterns, especially when these are driven by contrasting
patterns of energy acquisition and use.
‘I argue that we should attempt to address the question
of [ecological] generalizations capable of crossing the
land-to-sea boundary’ (Steele 1991a, p. 425).
‘…a strong conceptual framework for comparing life
in the marine realm with life on land has not been
developed’ (Dawson and Hamner 2008, p. 137).
he exchange of ideas between marine and terrestrial ecol-
ogy has largely been limited to a few targeted efforts (Steele
1991a, Beddington et al. 1994, Stergiou and Browman
2005). his reflects the ‘wet’ and ‘dry’ division of many facil-
ities, journals, meetings and funding streams (Raffaelli et al.
2005, Menge et al. 2009, Webb 2009), a possible response
by scientists to the physical and biological differences
between systems. Although biophysical differences between
marine and terrestrial systems can be overstated (Dawson and
Hamner 2008), in terms of primary production and the ecology
of consumer species these systems do differ fundamentally.
he dominant primary producers in the sea are unicellular,
motile, short lived and very responsive to short term changes
in the physical environment while the dominant primary
producers on land often provide structure, can grow in
mass by many orders of magnitude and are often relatively
unresponsive to short term environmental fluctuations.
Further, major groups of marine consumers (especially fish)
tend to grow over many orders of magnitude in mass, change
habitats and prey types with size, release large numbers of
small pelagic eggs and provide no parental care while ter-
restrial vertebrate consumers have limited scope for growth,
form long-term associations with habitat and provide their
larger young with significant parental care.
One consequence of the differences between marine and
terrestrial systems is that size rather than species identity
accounts for most of the variation in the trophic roles of
marine animals. Recognition of the significance of size led
to the emergence of theory and applications that describe
marine food web and system properties in terms of size
composition (Kerr and Dickie 2001). Conversely, terrestrial
food web theory focuses on species and on groups of spe-
cies with similar trophic roles. he marine approach reflects
the dominance of small primary producers in the marine
environment, the relatively consistent relationships between
predator and prey body sizes and the considerable scope for
growth in many marine species, often exceeding five orders
of magnitude in mass (Cushing 1975). Since many mac-
roecological patterns and processes are a consequence of
the energy demands and life histories of the species in a
community (Brown et al. 2004) the development of theory
Oikos 000: 001–013, 2011
doi: 10.1111/j.1600-0706.2011.18870.x
© 2011 he Authors. Oikos © 2011 Nordic Society Oikos
Subject Editor: Andrea Belgrano. Accepted 18 November 2010
OIKO_A_018870.indd 1 OIKO_A_018870.indd 1 1/6/2011 5:22:08 PM 1/6/2011 5:22:08 PM