Rocky Mountain Lodgepole Pine Research Articles

Vegetation response to wildfire and climate forcing in a Rocky Mountain lodgepole pine forest over the past 2500 years

Wildfire is a ubiquitous disturbance agent in subalpine forests in western North America. Lodgepole pine ( Pinus contorta var. latifolia), a dominant tree species in these forests, is largely resilient to high-severity fires, but this resilience may be compromised under future scenarios of altered climate and fire activity. We investigated fire occurrence and post-fire vegetation change in a lodgepole pine forest over the past 2500 years to understand ecosystem responses to variability in wildfire and climate. We reconstructed vegetation composition from pollen preserved in a sediment core from Chickaree Lake, Colorado, USA (1.5-ha lake), in Rocky Mountain National Park, and compared vegetation change to an existing fire history record. Pollen samples ( n = 52) were analyzed to characterize millennial-scale and short-term (decadal-scale) changes in vegetation associated with multiple high-severity fire events. Pollen assemblages were dominated by Pinus throughout the record, reflecting the persistence of lodgepole pine. Wildfires resulted in significant declines in Pinus pollen percentages, but pollen assemblages returned to pre-fire conditions after 18 fire events, within c.75 years. The primary broad-scale change was an increase in Picea, Artemisia, Rosaceae, and Arceuthobium pollen types, around 1155 calibrated years before present. The timing of this change is coincident with changes in regional pollen records, and a shift toward wetter winter conditions identified from regional paleoclimate records. Our results indicate the overall stability of vegetation in Rocky Mountain lodgepole pine forests during climate changes and repeated high-severity fires. Contemporary deviations from this pattern of resilience could indicate future recovery challenges in these ecosystems.

Enhanced seed defenses potentially relax selection by seed predators against serotiny in lodgepole pine.

Serotiny, the retention of seeds in a canopy seed bank until high temperatures cause seeds to be released, is an important life history trait for many woody plants in fire‐prone habitats. Serotiny provides a competitive advantage after fire but increases vulnerability to predispersal seed predation, due to the seeds being retained in clusters in predictable locations for extended periods. This creates opposing selection pressures. Serotiny is favored in areas of high fire frequency, but is selected against by predispersal seed predators. However, predation also selects for cone traits associated with seed defense that could reduce predation on serotinous cones and thereby relax selection against serotiny. This helps explain the elevated defenses in highly serotinous species. However, whether such interactions drive variation in seed defenses within variably serotinous populations has been studied rarely. We investigated the effects of phenotypic selection exerted by red squirrel (Tamiasciurus hudsonicus) predation on Rocky Mountain lodgepole pine (Pinus contorta latifolia) seeds. Squirrels preferentially harvested cones with more and larger seeds, indicating a preference for a higher food reward. We found evidence for stronger selection on trees with serotinous cones, which presumably accounts for the elevated defenses of and lower predation on serotinous compared to non‐serotinous cones. Lower levels of predation on serotinous cones in turn lessen selection against serotiny by squirrels. This has important implications because the frequency of serotiny in lodgepole pine has profound consequences for post‐fire communities and ecosystems widespread in the Rocky Mountains.

Habitat associations and abundance of a range-restricted specialist, the Cassia Crossbill (Loxia sinesciuris)

ABSTRACT The recently discovered Cassia Crossbill (Loxia sinesciuris) occurs only in 2 small, isolated mountain ranges in southern Idaho, USA: the South Hills and the Albion Mountains. The species faces 2 major threats from climate change related to its reliance on seeds of Rocky Mountain lodgepole pine (Pinus contorta latifolia). First, increased numbers of hot summer days (≥32°C) since 2003 apparently caused premature cone opening and seed shedding, leading to reduced seed availability and an 80% decline in Cassia Crossbill densities between 2003 and 2011. Second, climate change is predicted to prevent recruitment and could potentially cause the extirpation of lodgepole pine from the South Hills and Albion Mountains by 2080. This extirpation would cause the extinction of Cassia Crossbills, because they are unable to compete for lodgepole pine seeds elsewhere. Although recent summers with fewer hot days have allowed Cassia Crossbills to recover, determining their status will require using density across ...

Stand dynamics 11 years after retention harvest in a lodgepole pine forest

Structurally diverse forests provide resilience to an array of disturbances and are a mainstay of multiple-resource management. Silviculture based on natural disturbance can increase structural heterogeneity while providing other ecological and economic benefits. One useful silvicultural tool for promoting structural heterogeneity is retention harvesting, whereby a portion of forest stands are left unlogged, transitioning even-aged stands to multi-aged. We report stand and tree dynamics 11 years after retention harvest in a central Montana Rocky Mountain lodgepole pine forest with evidence for a mixed-severity fire regime. Treatments were implemented on 16 experimental units with prescriptions for two 50% overstory basal area retention patterns (Aggregated and Dispersed) crossed with two levels of prescribed fire use (Burned and Unburned). The aim of this study was to identify (1) how retention harvest spatial pattern affects stand dynamics, (2) if stand dynamics after retention treatments are more simply attributable to tree size and competition, or if retention pattern affects dynamics beyond those measures, and (3) how stem and basal area heterogeneity varied over the 11-year measurement period. Retention pattern affected overstory density, growth, mortality, and regeneration density and stocking. After controlling for the fine-scale factors of tree size and competition, overstory mortality, regeneration stocking, and regeneration height growth did not vary by treatment-scale factors. Fine-scaled factors explained significant variation in overstory basal area growth, but at the scale of experimental units, growth was also greater in Dispersed treatments. Prescribed burning interacted with retention pattern to influence overstory tree growth, increased overstory mortality, and increased regeneration height growth. Overstory heterogeneity (e.g., in basal area) degraded more rapidly in treatments with the Dispersed spatial pattern than Aggregated. This study evaluates novel silvicultural treatments in a lodgepole pine forest and highlights the tradeoffs between retention patterns combined with broadcast burning on forest change. Our results are useful for planning silvicultural treatments in multiple-use forests designed to promote structural complexity and resilience to disturbances.

Limited evidence for CO2 -related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients.

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO2 may enhance photosynthesis and/or decrease stomatal conductance (gs ) thereby enhancing intrinsic water-use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO2 and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ13 C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in gs . Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing gs may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.

The Natural History of the South Hills Crossbill in Relation to Its Impending Extinction.

Increasingly, the species that we discover will be uncommon, area restricted, and vulnerable to extinction. I describe the natural history of a newly discovered seed-eating finch from the Rocky Mountain region, the South Hills crossbill (Loxia curvirostra complex). It relies on seeds in the closed cones of the fire-adapted Rocky Mountain lodgepole pine (Pinus contorta latifolia) and is found only in the higher elevations of two small mountain ranges in southern Idaho. Here crossbills and pine are engaged in a coevolutionary arms race. Although most of the seeds remain secured within the cones for decades until the heat of a stand-replacing fire causes the cone scales to separate, seeds become accessible to crossbills slowly as cones weather and gaps form between some of the scales. However, hot days (≥32°C), especially four or more hot days, seem to mimic the effect of fire, apparently causing the immediate release of a fraction of the seeds. Such events caused a 20% annual decline in crossbills that lasted up to 4 years and an 80% decline in the population between 2003 and 2011. This is an example of a novel trophic mismatch between a consumer and its resource caused by a shift in the phenology of the resource arising from climate change. Not only do these phenological shifts have the potential to cause seed consumers to decline, these shifts are also likely to cause reduced recruitment of the plants. The South Hills crossbill is especially vulnerable and will likely go extinct this century before lodgepole pine is extirpated from the South Hills.

From the ground up: biotic and abiotic features that set the course from genes to ecosystems.

Spatial variation in cone serotiny in Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) across Yellowstone National Park influences initial pine recruitment after stand‐replacing fire with tremendous population, community, and ecosystem consequences. A previous study showed that much of the spatial variation in serotiny results from the balance of selection arising from high frequencies of fire favoring serotiny countered by opposing selection exerted by American red squirrels (Tamiasciurus hudsonicus) as seed predators. This earlier study, however, assumed stable local red squirrel densities over multiple generations of pines. Here, we examine environmental properties that might contribute to long‐term stability in the densities of red squirrels among sites. We found that the amount of clay in the soil, an indicator of plant and fungal growth—the latter an important food resource for red squirrels—and the coefficient of variation (CV) in diameter at breast height (DBH) of forest trees together account for a substantial amount of variation in red squirrel density. Soil development occurs over very long time scales, and thus, intersite variation in the amount of clay is unlikely to shift across pine generations. However, CV of DBH and squirrel density increase with stand age, which acts to amplify selection against serotiny with increasing interfire interval. Regardless, much of the variation in the CV of DBH is accounted for by soil bulk density, mean annual temperature, and surface curvature, which are unlikely to vary in their relative differences among sites over time. Consequently, these soil and abiotic attributes could contribute to consistent spatial patterns of red squirrel densities from one pine generation to the next, resulting in consistent local and spatial variation in selection exerted by red squirrels against serotiny.

Forest composition change after a mountain pine beetle outbreak, Rocky Mountain National Park, CO, USA

Recent severe and extensive mountain pine beetle (Dendroctonus ponderosae; MPB) outbreaks have created novel conditions in Southern Rocky Mountain lodgepole pine forests which historically had disturbance regimes dominated by extensive, stand-replacing fires. The goal of this study is to investigate patterns of and potential mechanisms in post-outbreak forest change in order to better understand the ecological legacy of the recent outbreak in the context of its implications for resilience to future disturbances and adaptation to climate change. To this end, we collected field data on forest structure and species composition in 2012 in lodgepole pine dominant forests in Rocky Mountain National Park. We then used a combination of modeling and statistical methods to identify possible mechanisms in post-outbreak forest conditions and evaluate the effect of the MPB outbreak on forest heterogeneity. We found that the outbreak initiated a shift in forest structure from single-cohort lodgepole pine stands to stands with greater diversity in age classes and species composition. This increase in landscape asynchrony may increase resiliency to future disturbances. However, this heterogeneity is a result of more spruce and fir on the landscape, species which are less adapted to projected future climate conditions. Our results indicate that disturbances do not necessarily increase the rate at which vegetation adapts to a changing climate, and that it is essential to consider disturbance type and available seed sources when predicting future forest conditions.

Conflicting selection from fire and seed predation drives fine-scaled phenotypic variation in a widespread North American conifer.

Recent work has demonstrated that evolutionary processes shape ecological dynamics on relatively short timescales (eco-evolutionary dynamics), but demonstrating these effects at large spatial scales in natural landscapes has proven difficult. We used empirical studies and modeling to investigate how selective pressures from fire and predispersal seed predation affect the evolution of serotiny, an ecologically important trait. Serotiny is a highly heritable key reproductive trait in Rocky Mountain lodgepole pine (Pinus contorta subsp. latifolia), a conifer that dominates millions of hectares in western North America. In these forests, the frequency of serotiny determines postfire seedling density with corresponding community- and ecosystem-level effects. We found that serotinous individuals have a selective advantage at high fire frequencies and low predation pressure; however, very high seed predation shifted the selective advantage to nonserotinous individuals even at high fire frequencies. Simulation modeling suggests that spatial variation in the frequency of serotiny results from heterogeneity in these two selective agents. These results, combined with previous findings showing a negative association between the density of seed predators and the frequency of serotiny at both landscape and continental scales, demonstrate that contemporary patterns in serotiny reflect an evolutionary response to conflicting selection pressures from fire and seed predation. Thus, we show that variation in the frequency of a heritable polygenic trait depends on spatial variation in two dominant selective agents, and, importantly, the effects of the local trait variation propagate with profound consequences to the structure and function of communities and ecosystems across a large landscape.

Variable resource availability when resource replenishment is constant: The coupling of predators and prey

ABSTRACT. Although food resources are thought to limit many populations, the extent to which the population dynamics of predators and prey are coupled is rarely known. We examined a sedentary population of Red Crossbills (Loxia curvirostra L. complex) that relies on seeds in cones that accumulate in the canopy of Rocky Mountain Lodgepole Pine (Pinus contorta latifolia Engelm.). Nearly constant annual seed production and gradual weathering over many years of initially impenetrable cones in the tree canopy results in a continuous and perhaps roughly constant replenishment of accessible seeds. However, seed availability varies seasonally. We estimated the seasonal variation in the energy demands of the study population. Our results demonstrate that seed predation by these sedentary Red Crossbills potentially drives the seasonal variation in seed availability and likely causes the Red Crossbill population to be regulated. The results are also consistent with a nearly constant replenishment of accessible seeds...

Survival and population size of a resident bird species are declining as temperature increases

1. A large number of migratory bird species appear to be declining as the result of climate change, but whether resident bird species have or will be adversely affected by climate change is less clear. We focus on the South Hills crossbill (Loxia curvirostra complex), which is endemic to about 70 km(2) of Rocky Mountain lodgepole pine (Pinus contorta latifolia) forest in southern Idaho, USA. 2. Our results indicate that the South Hills crossbill has declined by over 60% between 2003 and 2008, and that decreasing adult survival drives this population decline. 3. We evaluated the relative support for multiple hypotheses linking crossbill survival to climate, an ectoparasitic mite (scaly-leg mites Knemidokoptes jamaicensis), and the recent emergence of West Nile virus. Changes in adult apparent survival rate were closely associated with average spring and annual temperatures, and with high temperatures (≥32 °C) during summer, which have increased during the last decade. In contrast, there was little evidence that scaly-leg mites or West Nile virus contributed to recent declines in adult survival. 4. The most probable mechanism causing the decline in adult survival and population size is a decrease in the availability of their primary food resource, seeds in serotinous pine cones. Cone production has declined with increasing annual temperatures, and these cones appear to be prematurely opening owing to increasingly hot summer conditions releasing their seeds and reducing the carrying capacity for crossbills later in the year. 5. In light of regional climate change forecasts, which include an increase in both annual temperature and hot days (>32 °C), and the likely disappearance of lodgepole pine from southern Idaho by the end of this century, additional research is needed to determine how to maintain lodgepole pine forests and their supply of seeds to conserve one of the few bird species endemic to the continental United States.

Seed release in serotinous lodgepole pine forests after mountain pine beetle outbreak

There are concerns that large-scale stand mortality due to mountain pine beetle (MPB) could greatly reduce natural regeneration of serotinous Rocky Mountain (RM) lodgepole pine (Pinus contorta var. latifolia) because the closed cones are held in place without the fire cue for cone opening. We selected 20 stands (five stands each of live [control], 3 years since MPB [3-yr-MPB], 6 years since MPB [6-yr-MPB], and 9 years since MPB [9-yr-MPB] mortality) in north central British Columbia, Canada. The goal was to determine partial loss of serotiny due to fall of crown-stored cones via breakage of branches and in situ opening of canopy cones throughout the 2008 and 2009 growing seasons. We also quantified seed release by the opening of forest-floor cones, loss of seed from rodent predation, and cone burial. Trees killed by MPB three years earlier dropped approximately 3.5 times more cones via branch breakage compared to live stands. After six years, MPB-killed stands had released 45% of their canopy seed bank through cone opening, cone fall due to breakage, and squirrel predation. Further losses of canopy seed banks are expected with time since we found 9-yr-MPB stands had 38% more open canopy cones. This was countered by the development of a modest forest-floor seed bank (6% of the original canopy seed bank) from burial of cones; this seed bank may be ecologically important if a fire or anthropogenic disturbance reexposes these cones. If adequate levels of regeneration are to occur, disturbances to create seedbeds must occur shortly after tree mortality, before the seed banks are lost. Our findings also suggest that the sustained seed rain (over at least nine years) after MPB outbreak may be beneficial for population growth of ground-foraging vertebrates. Our study adds insight to the seed ecology of serotinous pines under a potentially continental-wide insect outbreak, threatening vast forests adapted to regeneration after fire. Key words: biotic disturbance; cone burial; cone opening; Dendroctonus ponderosae; ground-foraging vertebrates; mountain pine beetle; natural regeneration; Pinus contorta var. latifolia; Rocky Mountain lodgepole pine; seed banks; serotiny (canopy seed storage); Tamiasciurus hudsonicus.

Fire severity and seed source influence lodgepole pine (Pinus contorta var. murrayana) regeneration in the southern cascades, Lassen volcanic National Park, California

Rocky Mountain lodgepole pine, (Pinus contorta var. latifolia) regenerates quickly after high severity fire because seeds from serotinous cones are released immediately post-fire. Sierra lodgepole pine (P. contorta var. murrayana) forests burn with variable intensity resulting in different levels of severity and because this variety of lodgepole pine does not have serotinous cones, little is known about what factors influence post-fire regeneration. This study quantifies tree regeneration in a low, moderate, and high severity burn patch in a Sierra lodgepole forest 24 years after fire. Regeneration was measured in ten plots in each severity type. In each plot, we quantified pre- and post-fire forest structure (basal area, density), counted and aged tree seedlings and saplings of all species, and measured distance to the nearest seed bearing tree. There was no difference in the density of seedlings and saplings among severity classes. Distance and direction to the nearest seed bearing lodgepole pine were the best predictors of lodgepole seedling and sapling density in high severity plots. In contrast to Rocky Mountain lodgepole pine, regeneration of Sierra lodgepole pine appears to rely on in-seeding from surviving trees in low or moderate severity burn patches or live trees next to high severity burn patches. Our data demonstrate that Sierra lodgepole pine follows stand development pathways hypothesized for non-serotinous stands of Rocky Mountain lodgepole pine.

Diversifying Coevolution between Crossbills and Conifers

Coevolution between granivorous crossbills (Loxia spp.) and conifers has been a prominent process in the diversification of crossbills. A striking example occurs in western North America where coevolution between crossbills and Rocky Mountain lodgepole pine (Pinus contorta latifolia) is ongoing in isolated ranges without the crossbill’s dominant competitor for seeds, the red squirrel (Tamiasciurus hudsonicus). Preferential foraging by crossbills on lodgepole pine cones in the South Hills and Albion Mountains, two small mountain ranges in southern Idaho where red squirrels are absent, has led to the evolution of larger, thicker-scaled cones than in nearby ranges where red squirrels are present. This in turn has favored the evolution of larger-billed crossbills that have diverged from other crossbills in the region. However, such diversifying coevolution, resulting from geographic variation in the distribution of strongly interacting species, is vulnerable to species introductions. For example, the introduction of red squirrels caused the precipitous decline and perhaps extinction of the Newfoundland crossbill and perhaps a crossbill endemic to the Cypress Hills, Canada. In general, species introductions act to reduce the geographic variation in species interactions, which may be critical for the diversification of many taxa.

A Coevolutionary Arms Race Causes Ecological Speciation in Crossbills

We examined three ecological factors potentially causing premating reproductive isolation to determine whether divergent selection as a result of coevolution between South Hills crossbills (Loxia curvirostra complex) and Rocky Mountain lodgepole pine (Pinus contorta latifolia) promotes ecological speciation. One factor was habitat isolation arising because of enhanced seed defenses of lodgepole pine in the South Hills. This caused the crossbill call types (morphologically and vocally differentiated forms) adapted to alternative resources to be rare. Another occurred when crossbills of other call types moved into the South Hills late in the breeding season and feeding conditions were deteriorating so that relatively few non-South Hills crossbills bred ("immigrant infecundity"). Finally, among those crossbills that bred, pairing was strongly assortative by call type (behavioral isolation). Total reproductive isolation between South Hills crossbills and the two other crossbills most common in the South Hills (call types 2 and 5) summed to .9975 and .9998, respectively, on a scale of 0 (no reproductive isolation) to 1 (complete reproductive isolation). These extremely high levels of reproductive isolation indicate that the divergent selection resulting from the coevolutionary arms race between crossbills and lodgepole pine is causing the South Hills crossbill to speciate.

A Coevolutionary Arms Race Causes Ecological Speciation in Crossbills

We examined three ecological factors potentially causing premating reproductive isolation to determine whether divergent selection as a result of coevolution between South Hills crossbills (Loxia curvirostra complex) and Rocky Mountain lodgepole pine (Pinus contorta latifolia) promotes ecological speciation. One factor was habitat isolation arising because of enhanced seed defenses of lodgepole pine in the South Hills. This caused the crossbill call types (morphologically and vocally differentiated forms) adapted to alternative resources to be rare. Another occurred when crossbills of other call types moved into the South Hills late in the breeding season and feeding conditions were deteriorating so that relatively few non–South Hills crossbills bred (“immigrant infecundity”). Finally, among those crossbills that bred, pairing was strongly assortative by call type (behavioral isolation). Total reproductive isolation between South Hills crossbills and the two other crossbills most common in the South Hills (call types 2 and 5) summed to .9975 and .9998, respectively, on a scale of 0 (no reproductive isolation) to 1 (complete reproductive isolation). These extremely high levels of reproductive isolation indicate that the divergent selection resulting from the coevolutionary arms race between crossbills and lodgepole pine is causing the South Hills crossbill to speciate.

CAN SELECTION BY AN ECTOPARASITE DRIVE A POPULATION OF RED CROSSBILLS FROM ITS ADAPTIVE PEAK?

The bill structures of different call types of red crossbills (Loxia curvirostra complex) in western North America usually approximate the predicted optima for foraging on single species of conifers. One clear exception is the call type in the South Hills, Idaho, that is coevolving in an evolutionary arms race with Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia). Although South Hills crossbills forage only on the cones of these lodgepole pines, their average bill depth is smaller than that predicted to be optimal. Because preliminary data showed that large-billed males were more likely to exhibit symptoms of ectoparasitic mite (Knemidokoptes jamaicensis) infestation, the goal of our study was to further quantify the incidence of mite infestation and determine whether selection by mites may have favored smaller-billed crossbills and thus driven crossbills away from the foraging optimum. We estimated annual survival of both infected and uninfected South Hills crossbills using program MARK, which allows for auxiliary variables such as bill size and sex to be included in survival analyses. Mite infestation depressed crossbill survival and, especially for males, caused directional selection against larger-billed individuals. Such selection may explain why South Hills crossbills have smaller bills than the optimum and why average bill size for males has decreased from 1998 to 2003. This selection may also explain why the degree of sexual size dimorphism has decreased by nearly 50% since 1998.

CAN SELECTION BY AN ECTOPARASITE DRIVE A POPULATION OF RED CROSSBILLS FROM ITS ADAPTIVE PEAK?

The bill structures of different call types of red crossbills (Loxia curvirostra complex) in western North America usually approximate the predicted optima for foraging on single species of conifers. One clear exception is the call type in the South Hills, Idaho, that is coevolving in an evolutionary arms race with Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia). Although South Hills crossbills forage only on the cones of these lodgepole pines, their average bill depth is smaller than that predicted to be optimal. Because preliminary data showed that large-billed males were more likely to exhibit symptoms of ectoparasitic mite (Knemidokoptes jamaicensis) infestation, the goal of our study was to further quantify the incidence of mite infestation and determine whether selection by mites may have favored smaller-billed crossbills and thus driven crossbills away from the foraging optimum. We estimated annual survival of both infected and uninfected South Hills crossbills using program MARK, which allows for auxiliary variables such as bill size and sex to be included in survival analyses. Mite infestation depressed crossbill survival and, especially for males, caused directional selection against larger-billed individuals. Such selection may explain why South Hills crossbills have smaller bills than the optimum and why average bill size for males has decreased from 1998 to 2003. This selection may also explain why the degree of sexual size dimorphism has decreased by nearly 50% since 1998.

A KEYSTONE SELECTIVE AGENT? PINE SQUIRRELS AND THE FREQUENCY OF SEROTINY IN LODGEPOLE PINE

Serotiny is a key life history trait in fire-prone habitats that is favored in plants that experience stand-replacing fires within their average life span. Although variation in fire frequency has been the focus of most studies attempting to understand variation in serotiny among populations and species of plants, other factors can select against serotiny. One agent in particular that can select against serotiny is an efficient pre-dispersal seed predator that eats a large fraction of the seeds held in the canopy. To test whether selection by such a predator selects against serotiny, we compared levels of serotiny in fire-adapted Rocky Mountain lodgepole pine (Pinus contorta ssp. latifolia) in five ranges lacking its dominant pre-dispersal seed predator, the pine squirrel (Tamiasciurus hudsonicus) to 344 stands with pine squirrels. Where pine squirrels were absent, the frequency of serotiny was consistently near 100%, whereas where squirrels were present, the frequency was variable, rarely approached the high levels found in areas without squirrels, and averaged much less than 50%. This indicates that squirrels select against serotiny and that, in the absence of squirrels, the frequency of serotiny would probably be uniformly higher throughout much of the Rocky Mountains. Because serotiny levels also influence the density of seedlings following a fire, squirrels, by selecting strongly against serotiny, have the potential to alter the early stages and perhaps the course of succession and various community attributes.

Reciprocal selection causes a coevolutionary arms race between crossbills and lodgepole pine.

Few studies have shown both reciprocal selection and reciprocal adaptations for a coevolving system in the wild. The goal of our study was to determine whether the patterns of selection on Rocky Mountain lodgepole pine (Pinus contorta spp. latifolia) and red crossbills (Loxia curvirostra complex) were concordant with earlier published evidence of reciprocal adaptations in lodgepole pine and crossbills on isolated mountain ranges in the absence of red squirrels (Tamiasciurus hudsonicus). We found that selection (directional) by crossbills on lodgepole pine where Tamiasciurus are absent was divergent from the selection (directional) exerted by Tamiasciurus on lodgepole pine. This resulted in divergent selection between areas with and without Tamiasciurus that was congruent with the geographic patterns of cone variation. In the South Hills, Idaho, where Tamiasciurus are absent and red crossbills are thought to be coevolving with lodgepole pine, crossbills experienced stabilizing selection on bill size, with cone structure as the agent of selection. These results show that crossbills and lodgepole pine exhibit reciprocal adaptations in response to reciprocal selection, and they provide insight into the traits mediating and responding to selection in a coevolutionary arms race.