Evolutionary implications of hierarchical impacts of nonlethal injury on reproduction, including maternal effects
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Nonlethal injury is pervasive in metazoans, but surprisingly little is known about its impact upon reproductive allocation. The impact of injury on reproduction has been explored in some detail in lizards and salamanders, which have tails that are adapted for fat storage but are also injured or lost during predatory and social encounters. We synthesize diverse insights from these studies and propose new hypotheses using graphical models which highlight three distinct, hierarchical effects of injury on reproduction: reproductive inhibition, reduction in propagule number and diminished per-propagule investment (PPI), a maternal effect. Previous studies, which involved experimentally amputating the whole tail, have provided evidence of the first two effects, although there is little evidence of reduced PPI. We assayed these effects in free-ranging Desmognathus salamanders exhibiting naturally occurring injury. Whereas earlier studies found that tail injury prohibits reproduction (precipitating functional conflict), we found that females missing 80% of their tails, including the smallest mature individuals, still reproduce. We also detected a negative correlation between magnitude of injury and PPI, a continuous maternal effect. Continuous (graded) effects of injury on PPI have not been reported previously; neither discrete nor continuous maternal effects due to injury have been previously demonstrated in free-ranging vertebrates. The dearth of evidence for such effects may be due to the design of experimental manipulations that use all-or-nothing treatments. Future studies employing quantitative field data, or more realistic experimental treatments that mimic the continuous distribution of injury are likely to detect maternal effects arising from nonlethal injury. Although our comparative and empirical findings derive from studies of lizards and salamanders, we discuss how they apply in principle to all metazoans. Studies of these effects in nonvertebrate model systems are sorely needed. 2005 The Linnean Society of London.
