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CALIFORNIA STATE UNIVERSITY, NORTHRIDGE Climate-Growth Sensitivity of a California Endemic Tree, Pseudotsuga Macrocarpa (Bigcone

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CALIFORNIA STATE UNIVERSITY, NORTHRIDGE Climate-Growth Sensitivity of a California Endemic Tree, Pseudotsuga Macrocarpa (Bigcone CALIFORNIA STATE UNIVERSITY, NORTHRIDGE Climate-growth sensitivity of a California endemic tree, Pseudotsuga macrocarpa (Bigcone douglas fir) A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Biology By Cody Alexander Russell May 2018 The thesis of Cody A. Russell is approved: _______________________________________ _______________ Dr. Paul Wilson, Ph.D. Date _______________________________________ _______________ Dr. Fritz Hertel, Ph.D. Date _______________________________________ _______________ Dr. Paula Schiffman, Ph.D. Chair Date California State University, Northridge ii TABLE OF CONTENTS Page SIGNATURE PAGE……………………………………………………….. ii LIST OF FIGURES………………………………………………………… iv ABSTRACT…………………………………………..….………………… v INTRODUCTION………………………………………………………….. 1 HYPOTHESIS ……………………………………………………………... 3 MATERIALS AND METHODS…………………………………………… 5 RESULTS…………………………………………………….….….……… 12 DISCUSSION………………………………………………….…………… 19 LITERATURE CITED……………………………………………….…….. 25 APPENDIX: Angeles National Forest low elevation site ………………….. 28 iii LIST OF TABLES AND FIGURES Page 1. FIGURE 1 A-C: …………………………………………………... 11 2. TABLE 1: ……………………………………………………….... 12 3. FIGURE 2: …………………………………………………........... 13 4. FIGURE 3: …………………………………………………........... 14 5. FIGURE 4 A&B: …………………………………………………. 15 6. FIGURE 5: ………………………………………………………... 16 7. FIGURE 6 A&B: …………………………………………………. 17 8. FIGURE 7: ………………………………………………………... 18 9. FIGURE 8: ………………………………………………………... 18 iv ABSTRACT Climate-growth sensitivity of a California endemic tree, Pseudotsuga macrocarpa (Bigcone douglas fir) By Cody A. Russell Master of Science in Biology Anthropogenic climate change is predicted to increase temperatures and reduce precipitation throughout most Mediterranean-climate regions. Projections indicate that species with narrow, limited distributions are at the greatest risk of climate-induced extinction. One such potentially vulnerable species is Pseudotsuga macrocarpa (Bigcone douglas fir), a large coniferous tree that is endemic to the Transverse Mountain Ranges of southern California. Although the species’ geographic extent is limited, it does occur at a wide range of elevations. Trees growing at lower elevations (300-1400 m) are relegated to cooler, mesic micro-habitats composed of steep, north facing canyons. The most common surrounding vegetation in these semi-arid environments is chaparral and coastal sage scrub. Conversely, trees growing at higher elevations (1400-2500 m) are found in continuous habitat and are more spatially extensive. Stands of P. macrocarpa at these elevations are not relegated to north facing exposure or steep slopes. v In semi-arid ecosystems, such as where P. macrocarpa is found, precipitation and to a lesser extent, temperature, are the most common factors that limit growth. The dynamic relationship between limiting climatic factors and a tree’s annual radial growth is commonly referred to as climate-growth sensitivity. Research has shown that tree species growing at their dry distribution limit are more sensitive to changes in climate. I hypothesized that the growth of P. macrocarpa individuals had the highest degree of climate sensitivity at low elevations. To address this hypothesis, I used dendrochronological techniques to extract tree core samples from individuals growing at sites of different elevations. Growth chronologies were created from the annual rings of core samples and the growth patterns were then correlated with local climate data. The analysis showed that lower elevation stands had smaller trunk diameters, greater inter-annual growth variability, increased sensitivity to drought, and a higher percentage of missing annual growth rings. If projected climate scenarios hold true, P. macrocarpa populations are at risk of extirpation at these more stressful lower elevations. vi Introduction Anthropogenic climate change is predicted to increase temperatures and reduce precipitation throughout most Mediterranean climate regions (Giorgi & Piero 2008). Many of these regions are already experiencing extreme weather conditions including drought and record-setting temperatures (AghaKouchak et al. 2014, Griffin & Anchukaitis 2014). Mediterranean-climate biodiversity hotspots such as the California Floristic Province, are habitat to thousands of species with limited ranges and specific ecological requirements (Dallman 1998m, Calsbeek et al. 2003). It is predicted that up to 66% of California’s native plants will experience climate change induced geographic range reductions of as much as 80% (Loarie et al. 2008). In addition, projections indicate that species with narrow, limited distributions are at the greatest risk of climate-induced extinction (Midgley et al. 2002, Thomas et al. 2004,). One such potentially vulnerable species is Pseudotsuga macrocarpa (Bigcone douglas fir), a large coniferous tree that is endemic to the Transverse Ranges of southern California. Pseudotsuga macrocarpa has a narrow and patchy distribution from Santa Barbara County in the north to San Diego County in the south. Although the species’ geographic extent is limited, it does occur at a wide range of elevations (300–2700 m; McDonald et al. 1976). Depending on its topographic position, P. macrocarpa is found in a variety of plant communities such as chaparral, riparian zones, and mixed evergreen woodlands (Sawyer et al. 2009, Kauffmann et al. 2017). Low elevation populations (300-1400m) are most commonly found within semi- arid chaparral in relatively cool, mesic montane islands. North-facing canyons with slopes averaging 35° provide the most common microenvironment for these populations (McDonald et al. 1976). In these patchy communities, P. macrocarpa is the dominant 1 species with Quercus chrysolepis (“canyon live oak”) and Adenostoma fasticulatum (“chamise”) creating the sub-canopy layer (McDonald et al. 1976, Sawyer et al. 2009). Higher elevation populations (1400-2500 m) are found in more continuous habitat and are more spatially extensive. Stands of P. macrocarpa at these elevations are not relegated to north facing exposure or steep slopes (Kauffmann et al. 2017). With increasing elevation, other coniferous tree species such as Calocedrus decurrens (“incense cedar”), Pinus coulteri (“Coulter pine”), and Abies concolour (“white fir”) begin to replace the chaparral as associate vegetation (McDonald et al. 1976). Pseudotsuga macrocarpa is one of the most fire-adapted conifers in the world (Lombardo et al. 2009). Adaptations such as thick bark (15 ̶ 20cm) and the ability to re- sprout from epicormic buds allow this long-lived species to survive and recover from the intense heat of fires that can occur approximately every two or three decades (Mensing et al. 1999, Vander Wall et al. 2006). Several individuals of this species have been dated to over 600 years old which means that many of these trees can endure as many as 20-30 severe fires during their lifespan (McDonald et al. 1976). 2 Hypothesis In all ecosystems, there are factors that control the rate and extent of annual growth in plants. In tropical systems, the limiting factors are often macronutrients because temperature and precipitation remain constant throughout the year (Tanner et al. 1998). In semi-arid ecosystems, such as where P. macrocarpa is found, precipitation and to a lesser extent, temperature, are the most common factors that limit growth (Fritts 1966). The dynamic relationship between limiting climatic factors and a tree’s annual radial growth is commonly referred to as climate-growth sensitivity (Herrero et al. 2012). Recent research found that tree species in rear-edge populations (those at their dry distribution limit) at low elevations had the highest degree of radial growth sensitivity in response to changes in climate (Herrero et al. 2012, Dolanc et al. 2013). There has been very little research on the ecology of Pseudotsuga macrocarpa and, no past study has looked at the species’ response to climate change. In many of the low elevation habitats, P. macrocarpa is the only long-lived conifer (Minnich 2007). Therefore, it is an excellent candidate to examine this climate-growth sensitivity relationship in southern California. Because it is a narrow endemic, this relationship should be particularly important for the persistence P. macrocarpa. Increased environmental stress, coupled with the inability for populations to migrate out of insular canyons might lead to an extensive reduction in the range of P. macrocarpa. This could have serious implications for ecosystem functions because P. macrocarpa provides shade and valuable vertical structure in what are otherwise exposed shrub-dominated landscapes (Meretsky & Snyder 1992, Minnich 2007, Sawyer et al. 3 2009). It is a significant seasonal food source for birds and rodents as well (Vander Wall et al. 2006). For this study, I hypothesized that the growth of P. macrocarpa individuals had the highest degree of climatic sensitivity at low elevations. Specifically, I expected to find low elevation trees with increased inter-annual growth variability and smaller average trunk diameters as a result of reduced annual radial growth. If my hypothesis was correct, rear-edge P. macrocarpa populations would be at the highest risk of extirpation due to anthropogenic climate change. In addition, this information might provide insight about the futures of other endemic plant species with limited distributions in Mediterranean-climate regions. 4 Materials and Methods Field Sampling Pseudotsuga macrocarpa
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