Long-term deer exclosure alters soil properties, plant traits, understory plant community and insect herbivory, but not the functional relationships among them

Evidence of the indirect effects of increasing global deer populations on other trophic levels is increasing. However, it remains unknown if excluding deer alters ecosystem functional relationships. We investigated how sika deer exclosure after 18 years changed soil conditions, the understory plant community, the traits of a dominant understory plant (Sasa palmata), herbivory by three insect-feeding guilds, and the functional relationships between these properties. Deer absence decreased understory plant diversity, but increased soil organic matter and ammonium concentrations. When deer were absent, S. palmata plants grew taller, with more, larger, and tougher leaves with higher polyphenol concentrations. Deer absence led to higher leaf area consumed by all insect guilds, but lower insect herbivory per plant due to increased resource abundance (i.e., a dilution effect). This indicates that deer presence strengthened insect herbivory per plant, while in deer absence plants compensated losses with growth. Because plant defenses increased in the absence of deer, higher insect abundances in deer absence may have outweighed lower consumption rates. A path model revealed that the functional relationships between the measured properties were similar between deer absence versus presence. Taken together, deer altered the abiotic and biotic environment, thereby changing insect herbivory, which might impact upon nutrient cycling and primary productivity. These results provide evidence that deer can alter interactions between trophic levels, but that functional relationships between certain ecosystem components may remain constant. These findings highlight the need to consider how increasing global deer populations can have cascade effects that might alter ecosystem dynamics. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1007/s00442-017-3895-3) contains supplementary material, which is available to authorized users.