INTERACTION OF A BIOTIC FACTOR (PREDATOR PRESENCE) AND AN ABIOTIC FACTOR (LOW-OXYGEN) AS AN INFLUENCE ON BENTHIC INVERTEBRATE COMMUNITIES

We examined the response of benthic invertebrates to hypoxia and predation risk in bioassay and behavioral experiments. In the bioassay, four invertebrate species differed widely in their tolerance of hypoxia. The mayfly, Callibaetis montanus, and the beetle larva, Hydaticus modestus, exhibited a low tolerance of hypoxia, the amphipod, Gammarus lacustris, was intermediate in its response and the caddisfly, Hesperophylax occidentalis, showed high tolerance of hypoxia. In the behavioral experiments, we observed the response of these benthic invertebrates, which differ in locomotor abilities, to vertical oxygen and temperature gradients similar to those in an ice-covered pond. With adequate oxygen, invertebrates typically remained on the bottom substrate. As benthic oxygen declined in the absence of fish, all taxa moved above the benthic refuge to areas with higher oxygen concentrations. In the presence of fish mayflies increased activity whereas all other taxa decreased activity in response to hypoxia. Mayflies and amphipods remained in the benthic refuge longer and endured lower oxygen concentrations whereas the vertical distribution of caddisflies and beetle larvae was not influenced by the presence of fish. As benthic oxygen declined in the presence of fish, all but the beetle larva reduced activity over all oxygen concentrations compared to when fish were absent. As benthic oxygen continued to decline, mayflies and amphipods moved above the benthic refuge and were preyed upon by fish. Thus, highly mobile taxa unable to tolerate hypoxia (mayflies and amphipods) responded behaviorally to declining oxygen concentrations by migrating upward in the water column. Taxa that were less mobile (beetle larvae) or hypoxia-tolerant (caddisflies) showed less of a response. Taxa most vulnerable to fish predation (mayflies and amphipods) showed a stronger behavioral response to predator presence than those less vulnerable (caddisflies and beetle larvae). Because invertebrates differ in their ability to withstand hypoxia, episodes of winter hypoxia could have long-lasting effects on benthic invertebrate communities either by direct mortality or selective predation on less tolerant taxa.