Resource Quantity and Quality Determine the Inter-Specific Associations between Ecosystem Engineers and Resource Users in a Cavity-Nest Web

While ecosystem engineering is a widespread structural force of ecological communities, the mechanisms underlying the inter-specific associations between ecosystem engineers and resource users are poorly understood. A proper knowledge of these mechanisms is, however, essential to understand how communities are structured. Previous studies suggest that increasing the quantity of resources provided by ecosystem engineers enhances populations of resource users. In a long-term study (1995-2011), we show that the quality of the resources (i.e. tree cavities) provided by ecosystem engineers is also a key feature that explains the inter-specific associations in a tree cavity-nest web. Red-naped sapsuckers ( Sphyrapicus nuchalis ) provided the most abundant cavities (52% of cavities, 0.49 cavities/ha). These cavities were less likely to be used than other cavity types by mountain bluebirds ( Sialia currucoides ), but provided numerous nest-sites (41% of nesting cavities) to tree swallows ( Tachycineta bicolour ). Swallows experienced low reproductive outputs in northern flicker ( Colaptes auratus ) cavities compared to those in sapsucker cavities (1.1 vs. 2.1 fledglings/nest), but the highly abundant flickers (33% of cavities, 0.25 cavities/ha) provided numerous suitable nest-sites for bluebirds (58%). The relative shortage of cavities supplied by hairy woodpeckers ( Picoides villosus ) and fungal/insect decay (<10% of cavities each, <0.09 cavities/ha) provided fewer breeding opportunities (<15% of nests), but represented high quality nest-sites for both bluebirds and swallows. Because both the quantity and quality of resources supplied by different ecosystem engineers may explain the amount of resources used by each resource user, conservation strategies may require different management actions to be implemented for the key ecosystem engineer of each resource user. We, therefore, urge the incorporation of both resource quantity and quality into models that assess community dynamics to improve conservation actions and our understanding of ecological communities based on ecosystem engineering.