Insect species interactions and resource effects in treeholes: are helodid beetles bottom-up facilitators of midge populations?

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Abstract
The insect community living in central Pennsylvania treeholes in autumn consists primarily of larvae of two species of helodid beetles, Prionocyphon discoideus and Helodes pulchella, and larvae of one species of ceratopogonid midge,Culicoides guttipennis. We manipulated treehole volume and the densities of these insects in laboratory microcosms. We hypothesized that: (1) helodid beetle larvae, which are shredders, would enhance growth and survival of ceratopogonid midge larvae (deposit feeders) in a processing chain commensalism, and (2) the quantity of resources expressed as water volume plus leaf litter would affect helodids and protozoans directly. Intraspecific competition was not found in midges, nor was interspecific competition between the two helodid species. Protozoan population densities decreased or grew slower in the presence of insects and in smaller microcosms. Development time and adult wing length of the midge (C. guttipennis) were affected by both total microcosm volume and insect species combination. Under resource limitation, midges grew larger in the presence of helodids, and in general, midges were larger in treatments with higher ratios of helodids to midges. Water chemistry in the microcosms was affected both temporally and by insect presence. Hydrogen ion levels decreased over time, and microcosms with no insects had lower hydrogen ion levels. Specific conductance increased in all treatments over time, and microcosms with no insects had lower conductivity than most treatments. Helodid larvae have a positive effect on midges, possibly due to a processing chain facilitation. If helodids are keystone decomposers in this system, their presence could affect resource availability and affect other organisms in the community. Similar processing chain commensalisms occur in other phytotelmata. These types of interactions may therefore be important in the structure and function of detritus-based communities.

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