Abstract

Natural enemies of plants such as insect herbivores can contribute to structuring and maintaining plant diversity in tropical forests. Most research in this area has focused on the role of specialized enemies and the extent to which herbivory on individual plant species is density‐dependent. Relatively few insect herbivores specialize on a single host plant species. Insect herbivores that feed on more than one plant species may link the regeneration dynamics of their host species through “apparent competition” or “apparent mutualism.” We investigated herbivory and survival of seedlings of two tropical tree species (Cordia alliodora and Cordia bicolor) in the forests of Barro Colorado Island (Panama). We used experiments and observations to assess seedling fate in relation to the presence of conspecifics and heterospecifics across a range of spatial scales. Herbivory significantly increased seedling mortality and was highest at high local densities of C. alliodora seedlings. There was also evidence that high local densities of C. alliodora increased herbivory on co‐occurring C. bicolor seedlings. Synthesis. The elevated rates of seedling herbivory at high densities of conspecifics documented in our study are consistent with the predictions of the Janzen–Connell hypothesis, which explains how so many plant species can coexist in tropical forests. Our data also highlight the possibility that herbivore‐mediated density‐dependence, facilitated by herbivores that feed on multiple plant species, can also occur across plant species. Enemy‐mediated indirect effects of this sort have the potential to structure plant communities.

Highlights

  • Plants and their insect herbivores account for around half of the macroscopic diversity on earth

  • The elevated rates of seedling herbivory at high densities of conspecifics documented in our study are consistent with the predictions of the Janzen–Connell hypothesis, which explains how so many plant species can coexist in tropical forests

  • Host species sharing one or more enemies could experience enemy‐mediated interspecific density‐dependence, in‐ creasing or decreasing each other’s mortality rates via mechanisms such as “apparent competition” and “apparent mutualism” in loca‐ tions where they co‐occur (Abrams, Holt, & Roth, 1998; Holt, 1977). This could lead to the regeneration dynamics of plant species being linked to heterospecific as well as conspecific plant densities, and to non‐specialist enemies shaping distributions of plant species at the community level

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Summary

| INTRODUCTION

Plants and their insect herbivores account for around half of the macroscopic diversity on earth. The Janzen–Connell hy‐ pothesis predicts that host‐specific pests such as insect herbivores will cause seed and seedling mortality to be highest in areas where the density of conspecifics is high, and/or in the vicinity of conspe‐ cific adults This mechanism can facilitate high local plant diversity in tropical forests by limiting population growth of species that are common and favoring locally rare species (Connell, 1971; Janzen, 1970). In terms of their diet breadths, insect herbivores are known to range from specialists to generalists that attack a wide range of species (Novotny et al, 2010) In these instances, host species sharing one or more enemies could experience enemy‐mediated interspecific density‐dependence, in‐ creasing or decreasing each other’s mortality rates via mechanisms such as “apparent competition” and “apparent mutualism” in loca‐ tions where they co‐occur (Abrams, Holt, & Roth, 1998; Holt, 1977). Measuring the effects of congeneric distance‐ and density‐dependent herbivory allows us to assess whether enemy‐mediated indirect interactions between Cordia spe‐ cies occur at our study site

| MATERIALS AND METHODS
| DISCUSSION
Findings
CONFLICT OF INTEREST
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