Fidelity or love the one you're with? Biotic complexity and tradeoffs
can drive strategy and specificity in by-product mutualisms
Abstract
By-product mutualisms are ubiquitous yet seldom considered in models of
mutualism. Most models represent conditional mutualisms that shift
between mutualism and antagonism in response to shifts in costs and
benefits resulting from changes in environmental quality. However, in
by-product mutualisms, benefits arise as a part of normal life processes
and do not incur additional costs and do not have antagonistic alternate
states. Recognizing that mutualisms are diverse, new, or more flexible,
models are needed. We present a conceptual model that differs from
traditional models in three ways: (1) partners exchange by-product
benefits, (2) interactions do not have alternate antagonistic states,
and (3) tradeoffs are allowed among factors that influence environmental
quality. We applied this model to bark and ambrosia beetles that
associate with fungi and that have developed two distinct pathways to
by-product mutualism. We used independent axes for each major factor
influencing environmental quality including those that exhibit
tradeoffs. For these beetles, tradeoffs in nutrition and defense were
key to which mutualism pathway was taken and the degree of fidelity that
occurred among partners. This approach can be used to test hypotheses on
the relative importance of the drivers underlying the development of
by-product, or other, mutualisms.