Evidence for an adult summer diapause in mountain pine beetle (Coleoptera: Curculionidae) that varies geographically and among haplogroups.

Identifying dormancy traits is important for predicting insect population success, particularly in a changing climate that could disrupt evolved traits. The mountain pine beetle (Dendroctonus ponderosae Hopkins) is native to North America, is responsible for millions of acres of tree mortality, and is expanding northward in Canada. Research has identified thermal traits important to epidemic-phase ecology that vary among populations. Genomic research identified 3 mountain pine beetle haplogroups representing Pleistocene glacial refugia. Significant variation in generation timing aligning with the haplogroups has been observed. The adult stage was previously identified as the likely cause of differences among populations, although the mechanism(s) remain unclear. We tested for an adult summer diapause that varies among populations from 2 haplogroups, southern Colorado (CO) (central haplogroup) and southern Idaho (ID) (eastern haplogroup) using respirometry and reproduction experiments. Warm temperatures (25 °C) resulted in reduced respiration rates of central haplogroup mountain pine beetle compared to a cool temperature treatment (15 °C), whereas respiration of the eastern haplogroup did not differ between the treatments. Mated pairs of central haplogroup mountain pine beetle reared/held at 15 °C were more likely to be classified with a higher reproductive success rating compared to pairs reared/held at 25 °C. These results support a facultative summer adult diapause in southern CO central haplogroup mountain pine beetle. Manifestation of this diapause was low/absent among adults from the northerly ID location. This diapause likely serves to maintain univoltinism shown to be important for mountain pine beetle epidemic-phase ecology. The variation occurring among haplogroups highlights the long-term, evolved processes driving local adaptations in mountain pine beetle.

Advances in Semiochemical Repellents to Mitigate Host Mortality From the Spruce Beetle (Coleoptera: Curculionidae).

We tested 3-methyl-2-cyclohexen-1-one (MCH) and novel semiochemicals as potential spruce beetle (Dendroctonus rufipennis Kirby) (Coleoptera: Curculionidae, Scolytinae) repellents over multiple years in Utah and Colorado trapping bioassays. MCH is a known spruce beetle repellent and our testing revealed Acer kairomone blend (AKB) and isophorone plus sulcatone as repellents. We subsequently tested these semiochemicals for area and single tree protection to prevent spruce beetle attacks at locations in Utah, Colorado, Wyoming, New Mexico, and Alaska. Individual tree protection trials found MCH-AKB provided significant protection against spruce beetle attacks in the southern Rocky Mountains but not in Alaska. Adding sulcatone or doubling MCH-AKB pouches did not further enhance protection. A degree of protection was extended to spruce at least 10 m distant from the repellents, including in Alaska. Tree diameter was not a significant covariate among treated trees but was positively correlated with the probability of infestation for surrounding spruce. In area protection trials, spruce in control plots were 2.4 times more likely to be in a higher severity attack class compared with spruce in plots treated with MCH-AKB pouches deployed at 30 sets per hectare. Tree diameter had a significant, positive relationship to the probability of infestation. We found MCH-AKB to offer a high degree of protection against beetle attack in Engelmann spruce (Picea engelmannii Parry ex Engelm.) (Pinales: Pinaceae) (Picea engelmannii Parry ex Engelm.) (Pinales: Pinaceae), especially for single tree protection (66% of control trees were strip- or mass-attacked compared with 6% of repellent-treated trees). AKB requires registration and labeling, however, before this economical and environmentally benign semiochemical can be used operationally.

Evidence for a Prepupal Diapause in the Mountain Pine Beetle (Dendroctonus ponderosae).

Dormancy strategies, including diapause and quiescence, enable insects to evade adverse conditions and ensure seasonally appropriate life stages. A mechanistic understanding of a species' dormancy is necessary to predict population response in a changing climate. Climate change is influencing distribution patterns and population success of many species, including Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae: Scolytinae), the most important mortality agent of pines in western North America. Diapause is considered absent in D. ponderosae, and quiescence in the final larval stage prior to pupation (i.e., prepupal) is considered the main dormancy strategy. We evaluated if a facultative diapause in the prepupal stage, rather than a pupation threshold ~15°C (i.e., quiescence), could describe pupation patterns in two latitudinally separated D. ponderosae populations in the western United States. We hypothesized that if pupation occurs at lower temperatures than previously described, and if significant prepupal developmental delays occur, diapause is a likely physiological mechanism. Although there was considerable variation within and between populations, pupation occurred below the previously established threshold suggesting a prepupal facultative diapause that is induced when late instars experience cool temperatures. Individuals that pupated at temperatures below 15°C also had developmental delays, relative to development at warmer temperatures, consistent with diapause development. Pupation patterns differed between populations wherein diapause was induced at cooler temperatures and diapause development was shorter in southern compared with northern D. ponderosae. Recognition of a facultative diapause that varies among and between populations is critical for making predictions about future population response and range expansion in a changing climate.

Diapause and overwintering of two spruce bark beetle species.

Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce-dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi- or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.

3-Methylcyclohex-2-en-1-one for area and individual tree protection against spruce beetle (Coleoptera: Curculionidae: Scolytinae) attack in the southern Rocky Mountains.

We tested 3-methylcyclohex-2-en-1-one (MCH) and an Acer kairomone blend (AKB) as repellent semiochemicals for area and single tree protection to prevent spruce beetle (Dendroctonus rufipennis Kirby) attacks at locations in Utah and New Mexico. In the area protection study, we compared host infestation rates of MCH applications at three densities (20, 40, and 80 g MCH ha-1) against a control treatment over 0.64 ha plots centered within ~1.25 ha treatment blocks. All treatments included two baited funnel traps within the plot to assure spruce beetle pressure. Following beetle attack, plots were surveyed for new spruce beetle attacks and to quantify stand characteristics. The probability of more severe spruce beetle attacks was significantly reduced, by ~50%, in each of the MCH area treatments compared with the control treatment but there was no significant treatment difference among the MCH deployment densities. For the single tree protection study, we compared attack rates of MCH, Acer kairomone blend (AKB), and MCH plus AKB on spruce beetle-baited trees against bait-only trees. Each treatment was applied over a range of host diameters to test for host size effects. Seventy-five percent of control trees were mass-attacked, about one-third of MCH- and AKB-alone spruce was mass-attacked, and no MCH plus AKB spruce were mass-attacked. These results suggest that MCH alone is a marginal area and single tree protectant against spruce beetle but that deployment with other repellents can significantly increase treatment efficacy.

Defense traits in the long-lived Great Basin bristlecone pine and resistance to the native herbivore mountain pine beetle.

Mountain pine beetle (MPB, Dendroctonus ponderosae) is a significant mortality agent of Pinus, and climate-driven range expansion is occurring. Pinus defenses in recently invaded areas, including high elevations, are predicted to be lower than in areas with longer term MPB presence. MPB was recently observed in high-elevation forests of the Great Basin (GB) region, North America. Defense and susceptibility in two long-lived species, GB bristlecone pine (Pinus longaeva) and foxtail pine (P. balfouriana), are unclear, although they are sympatric with a common MPB host, limber pine (P. flexilis). We surveyed stands with sympatric GB bristlecone-limber pine and foxtail-limber pine to determine relative MPB attack susceptibility and constitutive defenses. MPB-caused mortality was extensive in limber, low in foxtail and absent in GB bristlecone pine. Defense traits, including constitutive monoterpenes, resin ducts and wood density, were higher in GB bristlecone and foxtail than in limber pine. GB bristlecone and foxtail pines have relatively high levels of constitutive defenses which make them less vulnerable to climate-driven MPB range expansion relative to other high-elevation pines. Long-term selective herbivore pressure and exaptation of traits for tree longevity are potential explanations, highlighting the complexity of predicting plant-insect interactions under climate change.

Prepupal diapause and instar IV developmental rates of the spruce beetle, Dendroctonus rufipennis (Coleoptera: Curculionidae, Scolytinae).

The spruce beetle, Dendroctonus rufipennis (Kirby), is an important mortality agent of native spruces throughout North America. The life-cycle duration of this species varies from 1 to 3 years depending temperature. The univoltine cycle (one generation per year) is thought to maximize outbreak risk and accelerate host mortality in established outbreaks. Prepupal diapause is associated with the semivoltine cycle (one generation per 2 years) and we investigated thermal conditions that result in diapause induction. Preliminary experiments used respirometry in an attempt to distinguish the diapause state of experimental insects but the technique was apparently confounded by low respiration before and during pupation, regardless of diapause status. Therefore, diapause induction was deduced using developmental delays. The observed developmental response was not a "switch", with developmental delay either present or absent, but instead varied continuously. We found that temperatures <15°C from instar III through mid-instar IV were associated with developmental delays beyond that expected from cool temperatures. Moreover, the duration of exposure to cool temperatures was important in determining the degree of developmental delay. Small, if any, delays were observed if the cumulative exposure to <15°C was <20 d whereas >40 d cumulative exposure was associated with distinct developmental suppression. Intermediate exposure to cool temperatures resulted in minor developmental delays. We used our results to parameterize a maximum likelihood estimation model of temperature-dependent instar IV developmental rates, including the effect of diapause. This model can be included as part of a spruce beetle phenology model for predicting population dynamics.