Journal of Animal
Ecology 2003
72, 873 – 887
© 2003 British
Ecological Society
Blackwell Publishing Ltd.
Nutrient exchanges between marine and terrestrial
ecosystems: the case of the Galapagos sea lion
Zalophus wollebaecki
J. M. FARIÑA*†, S. SALAZAR‡, K. P. WALLEM*, J. D. WITMAN† and
J. C. ELLIS†
*Center for Advanced Studies in Ecology and Biodiversity (CASEB), Pontificia Universidad Católica de Chile,
Santiago, Chile; †Department of Ecology and Evolutionary Biology, Brown University, Providence, USA; and
‡Marine Research and Conservation Department, Charles Darwin Research Station (CDRS), Puerto Ayora,
Galápagos, Ecuador
Summary
1. The movement of materials and organisms between ecosystems is a common process
in nature.
2. In the present study we investigate the hypothesis that the transport of nutrients by
low-mobility species and their effect on terrestrial ecosystems depends on habitat
topography. Specifically, we hypothesized that the influence of a marine organism with
low mobility on terrestrial environments would be spatially restricted.
3. To address this hypothesis we analysed the distribution (both geographical and local
scales) of Galapagos sea lion colonies, and quantified the spatial extent of their influ-
ence on terrestrial ecosystems (soil and plants).
4. Our results showed that the influence of Z. wollebaecki on Galapagos terrestrial
habitats is restricted to shorelines with low elevations, but that it is it is geographically
ubiquitous across the Archipelago.
5. Our study demonstrated that Z. wollebaecki is an effective vector for the transport of
marine nutrients to terrestrial ecosystems. Transported nutrients occur in high concen-
trations in the soils and are used by shoreline plants. These effects are spatially restricted
to the areas where seals occur and the most parsimonious explanatory variable for these
patterns is the islands’ topography (or elevation).
Key-words: Cryptocarpus, isotopic enrichment, sea lion, Sesuvium, soil ammonia and
nitrate, topography.
Journal of Animal Ecology (2003) 72, 873 – 887
Introduction
The movement of materials and organisms between
ecosystems is a common process in nature (e.g. Likens
& Borman 1975; Nakano & Murakami 2001). Cross-
ecosystem exchanges can be categorized into three
major groups: (1) transport of nutrients and materials
by physical agents such as water or wind; (2) transport
of nutrients and materials by biotic agents such as
vertebrates; and (3) movement of prey and consumers
between habitats (Polis, Anderson & Holt 1997). Since
Elton (1927), this process has been noted in both ter-
restrial and aquatic communities. However, a theoret-
ical framework for it has only recently been generated,
mainly based on food web theory (DeAngelis 1980;
Polis et al . 1997; Huxel & McCann 1998; Fagan,
Cantrell & Cosner 1999; Huxel, McCann & Polis
2002). In this framework it is recognized that nutrients
and materials are transported from more to less pro-
ductive ecosystems, and that these spatial subsidies
depend on the permeability of adjacent habitats, or on
specific characteristics of physical–biotic vectors. For
example, in the case of physical transport, the flow,
speed and volume of water and /or wind could deter-
mine the amount and kind of transported materials (Gosz
1991; Witman, Ellis & Anderson, in press). Mobility,
behaviour and physiology are important characteristics
Correspondence: J. M. Fariña, Center for Advanced Studies
in Ecology and Biodiversity (CASEB), Pontificia Universidad
Católica de Chile, Alameda 340, C.P. 6513677, Santiago,
Chile. Tel. +56 2 6862610; Fax: +56 2 6862621; E-mail: jmfa-
rina@bio.puc.cl