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On Santa Rosa Island, California Environmental 2019-20 Baseline Demographic Survey of the Bishop Pine (Pinus muricata) on Santa Rosa Island, California Environmental Science and Resource Management Capstone Project Andrew Tegley, Eric Schwarz, Victoria Joyce, and Grant Hassinger Submitted in partial fulfillment of the requirements for a Bachelors of Science degree in Environmental Science and Resource Management from California State University Channel Islands 1 Table of Contents: I. A b stract 3 II. Study A rea 6 III. H y p o th eses 7 IV. Materials and Methods 8 V. R esu lts 18 VI. D iscu ssio n 37 VII. C onclusion 43 VIII. Acknowledgements 45 IX. Literature Cited 45 X. A p p en d ix 51 2 Abstract Environmental factors, including drought, ungulate grazers, bark beetle infestation, and competition for resources have affected the abundance, distribution, and health of the Bishop Pine population on Santa Rosa Island, California. Santa Rosa Island, one of the northern Channel Islands, is the second-largest island in California, with an area of 217 km2. Since the 1980s, the island has been managed by the National Park Service, which brought an end to the era of ranching operations that had caused significant damage to the island’s native vegetation via ungulate grazing. The complete removal of ungulate grazers by 2011 provided a novel opportunity to monitor the regeneration of the island’s native vegetation. In 2016, a preliminary survey of the Bishop Pine population was conducted, which resulted in the establishment of thirty-five circular (6.3 m diameter) plots in the vicinity of Black Mountain. In 2019, we completed a thorough demography survey of the Bishop Pine population, thereby providing an assessment of the Bishop Pines’ status following the extraordinary drought that California experienced from 2012-2016 while simultaneously establishing a baseline set of data that future researchers can reference and build-upon. In our study, we discovered that there were five times more seedlings in 2019 as compared to 2016, and that most seedlings were found on slopes that ranged from 40-65% with lesser amounts of crust/moss. We also discovered that seedling and sapling growth did not occur in plots with levels of vegetation species richness higher than 10. 48% of Bishop Pine plots showed evidence of bark beetle infestation in 2019. Trees that had larger diameters, as well as trees that were unhealthy or dead prior to infestation in 2016, were associated with higher levels of infestation in 2019. Beetle samples were collected from underneath the bark of Bishop Pines throughout the study area. The samples were identified as weevils of the genusStenoscelis , i.e. inquilines, which take advantage of the damage wrought by the initial invasion of another species, and are often associated with a more-aggressive, primary invader. As such, the bark beetle species causing the initial damage and, potentially, contributing to the mortality of Bishop Pines on Santa Rosa Island is yet to be identified. Keywords: Santa Rosa Island, California, Bishop Pine, Pinus muricata, Channel Islands National Park, Bark Beetle, Stenoscelis, Demography Survey, DBH, Drought, Recruitment, Species Richness 3 Introduction Drought is a significant driver of forest mortality across the world. Anthropogenic activities have resulted in a trend of increasing global temperatures, which is causing droughts to increase in frequency and severity (Allen et al. 2010). Given that drought has such a pronounced impact on forest health and structure, it is important to understand the impacts of past and current droughts in order to better-predict, and mitigate, the effects of future droughts. The California drought of 2012-2016, a product of low precipitation and record-high temperatures, was the worst drought that California has experienced in the past 1,200 years (Griffin and Anchukaitis 2014). Since 2010, an estimated 147 million trees have died in California, primarily due to drought and drought-driven impacts, e.g. bark beetle epidemics (USDA 2019). These extreme events have raised concerns among conservationists and land managers regarding the future viability of native trees, particularly those with a limited distribution and small population size, such as the Bishop Pine (Pinus muricata D. Don) (Fischer 2009). The Bishop Pine is a drought-sensitive, sparsely distributed species of closed-cone pine that is endemic to the coast of California, including the northern Channel Islands, and Baja California, Mexico (Williams et al. 2008). Approximately 12,000 years ago, a warming climate reduced the once-extensive Bishop Pine forests of the Channel Islands to a small number of isolated stands that are spottily distributed across Santa Cruz and Santa Rosa Islands (Rick et al. 2014). Several efforts have been made to study the characteristics and structure of Bishop Pine communities on Santa Cruz Island, but little knowledge exists with regards to the structure and population dynamics of the Santa Rosa population, particularly following the extreme drought of 2012-2016 (Baguskas et al. 2016). These stands are primarily distributed on precipitous, north-facing slopes that are inundated by the coastal fog belt, which provides a critical source of moisture during the dry summer months (Rastogi et al. 2016, Buckley 2016, Pritchard 2016). On the Channel Islands, the Bishop Pine fulfills an ecological role as a keystone species by making water available to the surrounding flora and fauna during the dry summer months. The pine’s needles sequester and condense the moisture provided by the fog belt, thereby providing water to the surrounding environment via the process of “fog drip.” This mechanism is critical for the survival of other endemic species that have evolved in isolation on the islands, reliant on the availability of water during the summer (Williams et al. 2008, Fischer et al. 2009, Baguskas et al. 2016). Drought is a primary influence on the mortality of Bishop Pines, evidenced by large-scale diebacks on Santa Cruz Island during the drought years of 1976-1977, 1988-1989, and most-recently in 2012-2016 (Fischer et al. 2009, Taylor et al. 2019). Severe drought influences the population dynamics and structure of pine communities in several ways: carbon starvation; 4 hydraulic failure; competition over resources; bark beetle infestations; and soil desiccation (Allen et al. 2010, Grogan et al. 2000). In response to drought, trees close their stomata in order to prevent the excess loss of water, thereby halting the process of photosynthesis and, subsequently, energy production. If the drought outlasts the tree’s stored provisions then the tree will die as a result of “carbon starvation” (Allen et al. 2010). In addition, the rapid desiccation of a tree may result in the formation of emboli within the xylem, the tissues that transport water throughout the tree, thereby blocking the flow of water (Choat et al. 2012). Amongst these internal changes to the Bishop Pine, drought also leads to competition between the pine and its neighboring vegetation. During times of drought, pines compete with neighboring species for underground resources, especially water. Studies have shown that this competition for water, particularly between pines and oaks, has led to a decline in pine survival and growth rate (Jucker et al. 2014). Areas with high species richness are more likely to encounter such competition due to limited space and finite resources available. The physiological stressors that forest communities experience during periods of severe drought, e.g. carbon starvation and hydraulic failure, also make trees vulnerable to exploitation by insects, particularly bark beetles (Kolb et al. 2013). As temperatures and, subsequently, drought severity and frequency have increased on a global level, the range and prevalence of bark beetle populations has greatly increased (Raffa et al. 2008). These environmental conditions result in the impairment of metabolic processes that provide a conifer with the materials necessary to defend against foreign invaders. In order to complete their life cycle, bark beetles tunnel their way through the bark of a conifer, into the phloem or vascular tissues. Within the phloem, the insects carve galleries, which are laden with eggs. After hatching, the larvae feed on the tree’s tissues until they develop into adults and emerge from the bark in search of a host. Drought-stressed conifers have fewer nutrients and water resources to allocate to the production of resin, a viscous substance that is utilized as a primary defense mechanism, serving as both a physical barrier, and as a chemical repellent that expels invasive insects (Raffa et al. 2008). Drought can also threaten the health and recruitment of Bishop Pines via the desiccation of local soils. Pine growth is dependent upon the ability to take up water for cell expansion and growth (Hsiao 2000). This trait can be threatened in the case of extreme drought, like the one that occured in 2012-2016. Another stressor for Bishop Pine recruitment on the Channel Islands is ungulate grazers which would consume seedlings and saplings before they were big enough to not be a food source for these animals. Now post-drought and post-removal of grazers (McEachern et al. 2009), Bishop Pines have an opportunity for recruitment without these barriers that they have faced for more than a century of ranching land use. In this study, we provide insight into the overall status of the Bishop Pine community on Santa Rosa Island via an analysis of several biotic and abiotic factors, including current regeneration rates, bark beetle infestation, species richness levels, and resource competition measured in 36 plots that were previously established and sampled in 2016 (Buckley 2016, Pritchard 2016). 5 Study Area Santa Rosa Island is one of the northern Channel Islands, located 43 km offshore of mainland California within Channel Islands National Park. Managed by the National Park Service, Santa Rosa is the second-largest island in California with an area of 217 km2 (McEachern et al.
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