Invasive Pathogen Threatens Bird-Pine Mutualism: Implications for Sustaining a High-Elevation Ecosystem

Human-caused disruptions to seed-dispersal mutualisms increase the extinction risk for both plant and animal species. Large-seeded plants can be particularly vulnerable due to highly specialized dispersal systems and no compensatory regeneration mechanisms. Whitebark pine ( Pinus albicaulis), a keystone subalpine species, obligately depends upon the Clark\u27s Nutcracker (Nucifraga columbiana) for dispersal of its large, wingless seeds. Clark\u27s Nutcracker, a facultative mutualist with whitebark pine, is sensitive to rates of energy gain, and emigrates from subalpine forests during periods of cone shortages. The invasive fungal pathogen Cronartium ribicola, which causes white pine blister rust, reduces whitebark pine cone production by killing cone-bearing branches and trees. Mortality from blister rust reaches 90% or higher in some whitebark pine forests in the Northern Rocky Mountains, USA, and the rust now occurs nearly rangewide in whitebark pine. Our objectives were to identify the minimum level of cone production necessary to elicit seed dispersal by nutcrackers and to determine how cone production is influenced by forest structure and health. We quantified forest conditions and ecological interactions between nutcrackers and whitebark pine in three Rocky Mountain ecosystems that differ in levels of rust infection and mortality. Both the frequency of nutcracker occurrence and probability of seed dispersal were strongly related to annual whitebark pine cone production, which had a positive linear association with live whitebark pine basal area, and negative linear association with whitebark pine tree mortality and rust infection. From our data, we estimated that a threshold level of similar to 1000 cones/ha is needed for a high likelihood of seed dispersal by nutcrackers ( probability \u3e= 0.7), and that this level of cone production can be met by forests with live whitebark pine basal area \u3e5.0 m(2)/ha. The risk of mutualism disruption is greatest in northernmost Montana ( USA), where three-year mean cone production and live basal area fell below predicted threshold levels. There, nutcracker occurrence, seed dispersal, and whitebark pine regeneration were the lowest of the three ecosystems. Managers can use these threshold values to differentiate between restoration sites requiring planting of rust-resistant seedlings and sites where nutcracker seed dispersal can be expected

Whitebark Pine Stand Condition, Tree Abundance, and Cone Production as Predictors of Visitation by Clark's Nutcracker

Accurately quantifying key interactions between species is important for developing effective recovery strategies for threatened and endangered species. Whitebark pine (Pinus albicaulis), a candidate species for listing under the Endangered Species Act, depends on Clark's nutcracker (Nucifraga columbiana) for seed dispersal. As whitebark pine succumbs to exotic disease and mountain pine beetles (Dendroctonus ponderosae), cone production declines, and nutcrackers visit stands less frequently, reducing the probability of seed dispersal.We quantified whitebark pine forest structure, health metrics, and the frequency of nutcracker occurrence in national parks within the Northern and Central Rocky Mountains in 2008 and 2009. Forest health characteristics varied between the two regions, with the northern region in overall poorer health. Using these data, we show that a previously published model consistently under-predicts the proportion of survey hours resulting in nutcracker observations at all cone density levels. We present a new statistical model of the relationship between whitebark pine cone production and the probability of Clark's nutcracker occurrence based on combining data from this study and the previous study.Our model clarified earlier findings and suggested a lower cone production threshold value for predicting likely visitation by nutcrackers: Although nutcrackers do visit whitebark pine stands with few cones, the probability of visitation increases with increased cone production. We use information theoretics to show that beta regression is a more appropriate statistical framework for modeling the relationship between cone density and proportion of survey time resulting in nutcracker observations. We illustrate how resource managers may apply this model in the process of prioritizing areas for whitebark pine restoration

Record long-distance movement of a Deer Mouse, Peromyscus maniculatus, in a New England montane boreal forest

We report a single-season, non-homing movement of 4287 ± 10 m by an adult male Deer Mouse, Peromyscus maniculatus, in western Maine, United States, in summer 2014. The movement was achieved in two stages: 927 ± 3 m in four days and an additional 3360 ± 10 m in 44 days. This is approximately 40% greater than the previously documented maximum linear movement for an individual of this species

Whitebark and Foxtail Pine in Yosemite, Sequoia, and Kings Canyon National Parks: Initial Assessment of Stand Structure and Condition

The Inventory & Monitoring Division of the U.S. National Park Service conducts long-term monitoring to provide park managers information on the status and trends in biological and environmental attributes including white pines. White pines are foundational species in many subalpine ecosystems and are currently experiencing population declines. Here we present results on the status of whitebark and foxtail pine in the southern Sierra Nevada of California, an area understudied relative to other parts of their ranges. We selected random plot locations in Yosemite, Sequoia, and Kings Canyon national parks using an equal probability spatially-balanced approach. Tree- and plot-level data were collected on forest structure, composition, demography, cone production, crown mortality, and incidence of white pine blister rust and mountain pine beetle. We measured 7899 whitebark pine, 1112 foxtail pine, and 6085 other trees from 2012–2017. All factors for both species were spatially highly variable. Whitebark pine occurred in nearly-pure krummholz stands at or near treeline and as a minor component of mixed species forests. Ovulate cones were observed on 25% of whitebark pine and 69% of foxtail pine. Whitebark pine seedlings were recorded in 58% of plots, and foxtail pine seedlings in only 21% of plots. Crown mortality (8% in whitebark, 6% in foxtail) was low and significantly higher in 2017 compared to previous years. Less than 1% of whitebark and zero foxtail pine were infected with white pine blister rust and <1% of whitebark and foxtail pine displayed symptoms of mountain pine beetle attack. High elevation white pines in the southern Sierra Nevada are healthy compared to other portions of their range where population declines are significant and well documented. However, increasing white pine blister rust and mountain pine beetle occurrence, coupled with climate change projections, portend future declines for these species, underscoring the need for broad-scale collaborative monitoring

Detection Station Activity History

The detection station activity history used for Spatial Capture-Recapture modeling of American marten density. 238 rows correspond to distinct station, with 22 total checks corresponding to columns. Cell values of 1 indicate that a detection station was active during a specific check period, and cell values of 0 indicate it was not active

Individual Marten Captures

Individual marten encounter history used for Spatial Capture-recapture modeling of marten density in 'flat format': data must be coerced to a 3-dimensional array for model fitting using the BUGS language. Individual represents the identity of the marten encountered, trap represents the trap number at which the individual was encountered, check corresponds to the check # during which the individual marten was detected at the specific detection station, and detection type corresponds to the type of detection: 2 represents detection by camera only, 3 represents detection by non-invasive genetic sampling only, and 4 represents detection by both camera and non-invasive genetic sampling

Detection Station Locations

This file provides the x and y coordinates of 238 detection stations used to survey American marten in Maine, USA. Units are UTM/1000 (thus, km units); original projection of the data was UTM Zone 19N

Marten site covariates, occupancy models

Two site covariates used for occupancy modeling. each row corresponds to 1 of 238 detection stations: "Early" is a binary variable indicating whether a station was active during early winter (Jan. & Feb.), "MNEVC" corresponds to the z-standardized proportion of existing Tree vegetation cover within a 1km buffer surrounding the detection station location