Cone and seed traits of two Juniperus species influence roles of frugivores and scatter-hoarding rodents as seed dispersal agents

- Utah juniper (UJ) cones are dry with large seeds and do not attract frugivores.
- Western juniper (WJ) cones are succulent with smaller seeds and attract frugivores.
- Rodents, predominantly piñon mice, scatter-hoard WJ and UJ seeds.
- Seed-caching rodents are the primary, and often sole, dispersers of UJ seeds.
- Rodents are complimentary and secondary dispersers of WJ.

Seed dispersal in Juniperus is generally attributed to frugivores that consume the berry-like female cones. Some juniper cones are fleshy and resinous such as those of western juniper (Juniperus occidentalis), while others are dry and leathery such as those of Utah juniper (J. osteosperma). Rodents have been recorded harvesting Juniperus seeds and cones but are mostly considered seed predators. Our study sought to determine if rodents play a role in dispersal of western and Utah juniper seeds. We documented rodent harvest of cones and seeds of the locally-occurring juniper species and the alternate (non-local) juniper species in removal experiments at a western juniper site in northeastern California and a Utah juniper site in western Nevada. Characteristics of western and Utah juniper cones appeared to influence removal, as cones from the local juniper species were preferred at both sites. Conversely, removal of local and non-local seeds was similar. Piñon mice (Peromyscus truei) were responsible for most removal of cones and seeds at both sites. We used radioactively labeled seeds to follow seed fate and found many of these seeds in scattered caches (western juniper: 415 seeds in 82 caches, 63.0% of seeds found; Utah juniper: 458 seeds in 127 caches, 39.5% of seeds found) most of which were attributed to piñon mice. We found little evidence of frugivores dispersing Utah juniper seeds, thus scatter-hoarding rodents appear to be the main dispersal agents. Western juniper cones were eaten by frugivores, and scatter-hoarding is a complimentary or secondary form of seed dispersal. Our results support the notion that Utah juniper has adapted to xeric environments by conserving water through the loss of fleshy fruits that attract frugivores and instead relies on scatter-hoarding rodents as effective dispersal agents.