Revealing cryptic interactions between large mammalian herbivores and plant-dwelling arthropods via DNA metabarcoding.

In the past decade, it has become clear that omnivory, feeding on more than one trophic level, is important in natural and agricultural systems. Large mammalian herbivores (LMH) frequently encounter plant-dwelling arthropods (PDA) on their food plants. Yet, ingestion of PDA by LMH is only rarely addressed and the extent of this direct trophic interaction, especially at the PDA community level, remains unknown. Using a DNA-metabarcoding analysis on feces of free-ranging cattle from a replicated field experiment of heavily and moderately grazed paddocks, we reveal that feeding cattle (incidentally) ingest an entire food chain of PDA including herbivores, predators and parasites. Overall, 25 families of insects and four families of arachnids were ingested, a pattern that varied over the season, but not with grazing intensity. We identified the functional groups of PDA vulnerable to ingestion, such as sessile species and immature life stages. Most of the fecal samples (76%) contained sequences belonging to PDA, indicating that direct interactions are frequent. This study highlights the complex trophic connections between LMH and PDA. It may even be appropriate to consider LMH as omnivorous enemies of PDA.

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