Theory

This section provides the background knowledge necessary to understand why and how the Mountain Pine Beetle has spread to epidemic proportions by outlining the biology and ecology of the beetle, describes the host trees and assessing how climate change impacts these factors.

Mountain Pine Beetle Biology and Ecology

The Mountain Pine Beetle is a bark beetle native to the the forests of Northwest North America. Its current range is from the Pacific Coast to Western South Dakota and from Northern British Columbia to Mexico. In its Northern habitats it can be found at elevations as low as sea-level, but in the Southern range it is restricted to mountain regions higher than 3000 meters (Amman et al., 1990).
The adult beetle typically infests its host in July. It makes pitch tubes under the bark and constructs long, straight egg galleries (10-120 cm). The eggs hatch after 10-14 days (Amman et al., 1990) and during the late summer the larvae feed on the phloem which is fatal for the tree (Bentz et al., 1991).
The larvae and the beetle survive the winter under the bark by going through three stages, as described by Regniere and Bentz (2007). In the first stage they feed on the phloem. In the intermediate stage the beetle voids its guts and eliminates ice-nucleating agents from the body as much as possible. Furthermore, it starts accumulating cryptoprotectants (alcohol) in the body.
In the final stage the beetle reaches its maximum accumulation of cryptoprotectants and is cold-hardened. With this technique it can survive temperatures down to -40ºC. The development continues in the spring with pupation and in July/August the adult beetle can jump to the next host.Mountain pine beetle
   
This phenologic cycle is completed under the bark normally in one year, but at low summer temperatures it might take two years. There are also examples from low altitude locations in California, where the beetle has completed two cycles within one year (Amman et al., 1990). This may be one of the future scenarios with a general heating of the climate. Generally though, each stage in the life cycle has a threshold temperature which helps to synchronize the phenology with the seasonal climate variation (Bentz et al., 1991).
Bentz et al. (2001) suggests that there is genetic variation within the MPB species associated with the latitudinal variation, so the life-pattern is not only a matter of climatic variables but also genetic variation.

The MPB has developed a symbiosis with the blue stain fungi (Grosmannia clavigera). The beetle hosts this fungus in its mouth and thus carries it from tree to tree. The fungus limits the production of resin in the host tree which is the pine's only defence against the beetle infestation. Furthermore, it markedly limits the water transportation to the crown, which gradually discolours (Amman et al., 1990).
The MPB does have some natural enemies including other insects as well as birds (e.g. competition from other beetle larvae; woodpeckers feeding on the larvae). In addition to this the tree can increase the pitch flow if the water supply is adequate. None of these factors are adequate to control the beetle at epidemic levels, but at an endemic level they do constrain the MPB. The most effective way of controlling it is silvicultural methods such as thinning of potential host tree stands and creation of patches with different age and species composition (Amman et al., 1990).

Susceptible Host Trees

Healthy lodgepole pineMost western pines (sugar pine (Pinus lambertiana), white pine (Pinus monticola), ponderosa pine (pinus ponderosa), and Lodgepole pine (Pinus contorta Douglas ex Louden and Pinus Contorata var. latifolia)) are susceptible hosts of the MPB (Amman et al., 1990), but Lodgepole pine (Pinus contorta Douglas) and Ponderosa pine (Pinus ponderosae Lawson) are the most important host species (Logan and Powell, 2001: 160).
The fire suppression that humans have put upon the forests for the last 100 years could be one of the reasons why the current lodgepole stand is more susceptible to MPB attacks. Forest fires are important for lodgepoles, since it prepares the seedbed, releases seeds from the serotinous cones (that are triggered to release seeds by heat of fire), and eliminates more shade-tolerant species that would eventually replace the lodgepole pine (Logan and Powell, 2001). Furthermore, research point to the fact that pine trees are only susceptible to the MPB when they are between 80-160 years old. Thus, in a natural BC environment, where wildfires burn down the forest stand on average every 60 years, the infested part of the susceptible pine stand will vary between 17-25 %. At present the infested part of the susceptible stand is as high as 55 % (Taylor & Carrol, 2003: 41).


Climate Change Impacts

According to the Intergovernmental Panel on Climate Change (IPCC) climate change is unequivocal, thus having a profound effect on several of the parameters controlling the area suitable for MPB: “At continental, regional and ocean basin scales, numerous long-term changes in climate have been observed. These include changes in arctic temperatures and ice, widespread changes in precipitation amounts, ocean salinity, wind patterns and aspects of extreme weather including droughts, heavy precipitation, heat waves and the intensity of tropical cyclones” (IPCC, 2007: 7).
The current outbreak is an order of magnitude greater in area than previous outbreaks, reaching epidemic proportions, spreading over 4.1x10^6 ha (41.000 km^2) (Hamann & Wang, 2006: 2773 after Ebata 2004).
This is mainly due to an increased area of susceptible hosts (caused by fire suppression, thus having forests reach older age) and change in climate allowing MPB to expand further north and east to areas previously uninhabitable (Kurz et al., 2008). The favorable climate is caused by changes in:
  • Minimum Winter Temperature
  • Average Summer Temperature
  • Spring Precipitation
  • Aridity
Projecting the spread of MBP and other insect pests in relation to climate change is often done using General Circulation Models (GCMs), of which several have been developed[1]. However, the application of GCMs has several limitations: the finest spatial scale used in these models is often far too coarse to be applied to ecological surveys[2] (Logan et al., 2003: 132). As such interpolation is often necessary, but local climate data is to be preferred to limit the amount of uncertainties and ease projection of future spread.

According to Hamann and Wang (2006) British Columbia (49°-60° latitude) has experienced a warming similar to that of the predictions from general circulation models, around 0.7 °C, why the use of these has been common in many studies of future MPB scenarios. Even such a small change has caused severe ecological and economic problems, as can be seen be the spread of MPB, a problem that will be further discussed later on.
As mentioned elsewhere the MPBs temperature-controlled development cycles and lack of diapauses (Logan and Powell, 2001) is highly dependent upon changes in temperature. Research suggests (Hamann and Wang, 2006; Logan et al., 2003) that this outbreak could be caused by a lack of low winterMPB infected lodgepole pine minimum temperatures, which would normally reduce the populations of this native-insect pest by killing the beetle. If temperatures drop to less than -40°C even the beetle natural defense mechanisms (as outlined above) can't keep it alive. Higher summer temperatures, especially in august where eggs hatch, are also a significant contributor, as well as higher summer temperatures allow the MPB to spread to higher altitudes and latitudes. Carroll et al. (2004) suggests that if minimum 5% of the days in August are at least 18, 3°C warm this increases the risk of epidemic spreading.
Furthermore research suggests (Carroll et al., 2004) that reduced amount of precipitation in the late spring / early summer, as well as a general warming of the climate (the MPB needs a minimum of 833 degree days a year) has further enhanced the spread of the MPB.  To be precise, if precipitation in late spring are below average, and if there's more than 833 degree days in a year, the risk of an epidemic spread increases.If precipitation are in late spring are above average the Lodgepole Pine's natural resistance are enhanced, thus reducing the risk of an epidemic outbreak.
The death of vast areas of forest further enhances the impact of climate change due to emissions of carbon dioxide. Kurz et al. (2008: 987) estimates that the impact of beetles on NBP [net biome production] was projected to reach -20 MtC yr^-1 […] in 2009. Cumulative beetle impact over the 21-year simulation period was projected to be -270 Mt C released into the atmosphere as a result of decaying trees. That is roughly one third of British Columbia’s anthropogenic carbon emissions[3].



Predicting future impacts

The future impacts is predicted by data predictions by ClimateBC. The predictions are based on the scenarios available in the latest IPCC report (IPCC, 2007).
We have chosen to use the B1 and the A1FI scenarios for our prediction of future impacts. The B1 predicts the least change in carbon dioxide levels and temperature, whereas the A1FI, FI meaning Fossil Intensive, predicts a much bigger change in the two parameters.
As such, it is hoped that the two scenarios will reflect the upper and lower boundary of the spread that can be expected in the future.
Forest infested by the MPB




[1] The CGCM1 model developed by the Canadian Centre for Climate Modeling and Analysis www.cccma.bc.ec.gc.ca), and another by the Hadley Centre for Climate Prediction and Research  (www.met-office.gov.uk/research/hadley-centre). (Logan et al., 2003: 132)
[2]. “This problem has been addressed through modeling and interpolation techniques that project GCM predictions to ecologically meaningful spatial and temporal scales”. (Logan et al., 2003: 132)