Host selection behavior mediated by differential landing rates of the walnut twig beetle, Pityophthorus juglandis, and associated subcortical insect species, on two western North American walnut species, Juglans californica and J. major.
The walnut twig beetle, Pityophthorus juglandis Blackman (Coleoptera: Scolytidae), vectors a phytopathogenic fungus, Geosmithia morbida Kolařík et al. (Hypocreales), which causes thousand cankers disease (TCD) in walnut (Juglans sp.) and wingnut (Pterocarya sp., both Juglandaceae) trees. We investigated an early point in disease inception in two walnut species - Juglans californica S. Wats. and Juglans major (Torr. ex Sitsgr.) Heller - native to riparian forests of the western USA by comparing P. juglandis flight and landing responses to small-diameter branch sections. Twenty unbaited branch sections (10 each of J. californica and J. major) were presented in a completely randomized design to populations of P. juglandis at the USDA Agricultural Research Service, National Clonal Germplasm Repository (NCGR) Juglans collection located at Wolfskill Experimental Orchards (Winters, CA, USA) and at the California State University, Chico, Agricultural Teaching and Research Center (ATRC, Chico, CA). These assays were carried out within a 4- to 6-year period when weekly flight surveys with aggregation pheromone-baited multiple funnel traps revealed that P. juglandis flight activity-abundance was higher at the NCGR than at the ATRC. For the landing rate assays, adhesive-coated acetate sheets were wrapped around the branch sections and exchanged weekly. Three assays were completed at the NCGR (assays 1-3), whereas one assay was completed at the ATRC (assay 4). Landing rates on these traps were compared between J. californica and J. major. Two additional assays (5 and 6) were completed at the NCGR to compare responses to branch sections of J. californica and to similarly sized cardboard tubes (negative control). All six assays were completed over a 4-year span during the 4- to 6-year weekly flight survey period. Pooled landing rates of male and female P. juglandis (assays 1-4) demonstrated a preference by both sexes for J. californica over J. major. In assay 5 there was no preference by males or females for J. californica over the negative control, perhaps due to the low flight activity-abundance of P. juglandis during the assay. When repeated at a time of higher flight activity-abundance (assay 6), male and female landing rates on J. californica exceeded those on the negative control. Females of the invasive fruit-tree pinhole borer (an ambrosia beetle), Xyleborinus saxeseni (Ratzeburg), and an invasive bark beetle, Hypothenemus eruditus Westwood (both Coleoptera: Scolytidae), showed relatively higher flight responses than either sex of P. juglandis during most assays, suggesting higher population densities of these two other invasive species at the two orchards or a greater sensitivity to host volatiles. Xyleborinus saxeseni and H. eruditus preferred to land on J. major over J. californica and on J. californica over the negative control. Similarly, an invasive longhorned beetle, Nathrius brevipennis (Mulsant) (Coleoptera: Cerambycidae), showed a significant preference for J. major over J. californica, but not for J. californica over the negative control. More male N. brevipennis were trapped than females at both study sites [sex ratio ranged from 5:1 (assay 6) to 39:1 (assay 4)], and flight occurred only in the spring and early summer months. Another ambrosia beetle trapped at the NCGR and ATRC, Xyleborus affinis Eichhoff, represented the first records of this species from western North America. In summary, flight responses recorded in some of our assays for P. juglandis and several other subcortical insects on Juglans indicate that host preference by these insects may be determined by long-range olfactory cues that do not involve pheromones.