1. Genetic Structure of Ips Pini (Say) Populations

1. Genetic Structure of Ips pini (Say) Populations Anthony I. Cognato1, Steven J. Seybold2, and Felix A.H. Sperling3 The pine engraver, Ips pini (Say) is an important British Columbia, Idaho, and Montana produce between pest of North American pine trees and has been the 91 percent and 95 percent (-) - ipsdienol while other subject of considerable biological research (Wood and western populations produce > 94 percent (-) - ipsdienol Bright 1992). Although these beetles feed mostly on (Seybold and others 1995). Based on pheromone and dead or dying trees, epidemic populations can attack and cuticular hydrocarbon data, Seybold and others (1995) kill living trees. Aggregation pheromones mediate the hypothesize that two phenotypes exist in western North attraction of I. pini and other bark beetles to suitable America south of the Canadian border. food sources (Wood 1982). A more extensive investigation of pheromone Pheromones and their role in aggregation behavior phenotypes in British Columbia showed the existence of have been investigated extensively in I. pini (for ex- “New York,” “California,” and intermediate pheromone ample, Miller and others 1996, 1997; Seybold and phenotypes in different individuals within and among others 1992, 1995; Teale and others 1991). The phero- populations (Miller and others 1996). British Columbia mone components lanierone and ipsdienol are known to may represent an area of intergradation of eastern and be active for I. pini; however beetle populations across western populations, given the variation of pheromone North America vary in the production of and response to production among individuals. These populations lanierone and to the enantiomeric blend of ipsdienol. presumably interbreed (Miller and others 1996), al- Generally, populations in eastern North America pro- though the rate of interbreeding will be affected by duce and respond to <60 percent (-) - ipsdienol (“New female pheromone phenotype preference. York” phenotype) whereas in western North America Eastern and western populations of I. pini were not they produce and respond to >90 percent (-) - ipsdienol always considered a single species. They were described (“California” phenotype) (Seybold and others 1995). as separate species: I. pini in the east and I. oregonis in Lanerione is produced in small quantities by male I. pini the west. Morphological differences, such as the density from New York (Teale and others 1991) and has recently of punctures on the pronotum and the smoothness of the been discovered in a California population as well elytra, were used to distinguish the eastern and western (Quilici and others, in press). The synergistic effect of species. However, these morphological characters lanierone with ipsdienol is weaker for California I. pini intergrade in western Canada and the species were when compared with New York I. pini (Seybold and synonymized by Hopping (1964). Lanier (1972) added others 1992). The attraction of California I. pini to >90 further evidence of conspecificity with laboratory percent (-) - ipsdienol is hindered by the presence of breeding experiments showing that the eastern and high quantities of (+) - ipsdienol (Birch and others western individuals produce viable offspring. 1980). Therefore, the preference of female I. pini for the However, the population variation found in phero- “conspecific” pheromone phenotype may preclude mone and morphological characters once again calls into mating among eastern and western I. pini. (Hager and question the species status of I. pini. Do two species Teale 1996; Piston and Lanier 1974; Teale and others exist and hybridize in British Columbia, or is I. pini one 1994) species whose populations were allopatrically separated Pheromone phenotypic variation also occurs within in the past? western North America. Populations from southeastern To address this hypothesis, a genetic marker inde- ____________________ pendent of pheromone production is needed to assess 1 Postdoctoral Fellow, Department of Entomology, University of the relationship among I. pini populations. DNA Minnesota, St. Paul, MN 55108. Current address: The Natural History Museum, Department of Entomology, Cromwell, Road, sequence from the mitochondrial cytochrome oxidase I London SW75BD. gene was used because it is presumably independent of 2 Assistant Professor, Department of Entomology, University of pheromone production, and provides enough nucleotide Minnesota, St. Paul, MN 55108. variation to resolve relationships among populations 3 Assistant Professor, Department of ESPM, Division of Insect (see Simon and others 1994, for review of mitochondrial Biology, University of California, Berkeley 94720. genome use in insect phylogenetics). 1 A total of 97 individuals were PCR-amplified and this lineage. directly sequenced over 354 base pairs of the mitochon- More specimens from all regions, especially British drial cytochrome oxidase I gene. These sequences were Colombia and northwest US localities, will test the used to reconstruct a single most parsimonious phylog- robustness of these preliminary results. Mitochondrial eny (heuristic search with 10 random stepwise addi- markers only reflect maternal lineage, and the presence tions, Swofford 1993). Beetles were sampled from 10 of individuals with “eastern” lineage in Arizona, Mon- localities, including 3 from California, 2 from New York tana, and Washington may indicate introgression due to and 1 each from Alberta, Colorado, Washington, Mon- “eastern” lineage females breeding with “western” tana, and Arizona. Ten specimens were sampled from lineage males. The addition of a nuclear gene marker each locality except Montana, where only seven speci- will assess the phylogeny and gene flow for both sexes mens were obtained. and should help determine if “eastern” lineage males Two measures of gene flow (Lynch and Crease ever breed with “western” lineage females and con- 1990, Slatkin and Maddison 1989) indicate that the versely, determine if “western” lineage males breed with number of migrating individuals per generation (Nm ) is “eastern” lineage females. Concordance of the mito- 0.4 (by both measures) among all I. pini populations. chondrial and nuclear data sets would allow a better An Nm of approximately 1 is theoretically sufficient to assessment of the need for recognition of eastern and maintain genetic homogeneity among populations western I. pini as subspecies. (Avise 1994). Therefore, estimated gene flow for the sampled I. pini populations is moderate. Phylogenetic analysis revealed three monophyletic lineages: “eastern” (New York), “Rocky Mountain” (Alberta and Colorado), and “western” (all Californian populations and Arizona). One Arizona individual had a haplotype consistent with the “eastern” lineage. All three lineages were present in the Washington and Montana populations. This result shows genetic poly- morphism in regions of pheromone phenotype variation. The mitochondrial phylogeny is generally concor- dant with the geographic pattern of pheromone phenotypes (Seybold and others 1995). The “New York” and “California” pheromone phenotypes are associated, respectively, with the “eastern” and “western” lineages. The “Rocky Mountain” lineage may be associated with the second western pheromone phenotype (91 percent to 95 percent (-)- ipsdienol); however pheromone analysis from additional collection localities in this area are needed. The estimated moderate gene flow among populations supports the hypothesis that I. pini is one species. However, the mitochondrial phylogeny of populations indicates that there are three distinct lineages. Since female pheromone phenotype preferences exist between eastern and western I. pini, it may be reasonable to formalize the trivial names of “New York“ and “Califor- nia” I. pini with recognition of the “eastern” and “western” lineages as subspecies. The “Rocky Mountain” lineage may also represent a subspecies; however few populations have been sampled and diagnostic morphological characters have not yet been demonstrated for 2 2. Insecticide Resistance and Nucleotide Variability in the Coffee Berry Borer R.H. ffrench-Constant1 Abstract reduction in variability with the ecological success of The coffee berry borer beetle Hypothenemus hampei the insect and its ability to evolve insecticide resistance. (Ferrari)(Curculionidae: Scolytinae) is the major insect The coffee berry borer shows regular inbreeding. pest of coffee and has spread to most of the coffee Single mated female H. hampei burrow into coffee growing countries of the world. This beetle also dis- berries and produce large single families with highly plays an unusual life cycle, with regular sibling mating. distorted sex ratios (10 females to one male). The This regular inbreeding, and the population bottlenecks dwarfed males are flightless and mate with their sisters occurring on colonization of new regions, should lead to in the natal berry. The beetle is also ‘functionally low levels of genetic diversity. We were therefore haplodiploid,’ as paternal genes in males are neither interested in determining the level of nucleotide varia- expressed nor transmitted (Brun and others 1995a,b). tion in nuclear and mitochondrial genomes of this beetle Thus, both the nuclear and mitochondrial genome are worldwide. Here we show that two nuclear loci (Resis- strictly maternally inherited and should share an identi- tance to dieldrin and ITS2) are completely invariant, cal genealogy. Resistance to the cyclodiene type whilst some variability is maintained

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