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The Pennsylvania State University The Pennsylvania State University The Graduate School Department of Geography SPECIES COEXISTENCE IN AN ALPINE FOREST IN CENTRAL JAPAN A Dissertation in Geography by Amanda Beatrice Young 2016 Amanda Beatrice Young Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2016 The dissertation of Amanda B. Young was reviewed and approved* by the following: Alan Taylor Professor of Geography Dissertation Advisor Chair of Committee Andrew Carleton Professor of Geography Erica Smithwick Associate Professor of Geography Margot Kaye Associate Professor of Forest Ecology Cynthia Brewer Professor of Geography Head of the Department of Geography *Signatures are on file in the Graduate School iii ABSTRACT In this dissertation, I examine Bond’s “slow seedling” hypothesis in an alpine forest system in central Japan. Bond (1989) developed this idea to explain the spatial partitioning of angiosperms and gymnosperms, in which gymnosperms are less responsive to resources like light and soil nutrients. Angiosperms prefer nutrient rich, dry soils in areas with relatively warm climates, relegating gymnosperms to the cool nutrient poor environments. Thus, gymnosperms typically dominate at high elevations and latitudes in environments such as treeline, with low temperatures and poorly developed soils. The alpine treeline of central Japan is not composed of gymnosperms but an angiosperm, Betula ermanii Cham., and is bounded by a subalpine forest dominated by Abies mariesii Mast. below and a mat of scrub pine Pinus pumila (Pall.) Regel above. The B. ermanii treeline is abrupt and has had no apparent upward movement over the past 50 years despite significant warming (3ºC), indicating the ecosystem has long-term stability where gymnosperms and angiosperms coexist, though the dominance of each species is segregated by elevation. In this dissertation, I examine three drivers of the delineation of treelines - soil nutrients, the regeneration niche, and climate - from the subalpine forest through the B. ermanii treeline and into the P. pumila mat. In Chapter 2, I assessed Bond’s “slow seedling” hypothesis by examining relationships between soil nutrients and tree importance values across the three forest bands using linear mixed-effect models. I found that the predictions from Bond’s hypothesis were inconsistently correct. For example, soil nitrogen was highest in the B. ermanii treeline on volcanic mountains, as I expected, but was similar across forest belts on non-volcanic mountains. This mixed evidence suggests lithology plays a role in driving local composition. In Chapter 3, I explored differences in the regeneration niche across ontogeny (seedlings and trees) for B. ermanii and A. mariesii in the subalpine forest and treeline using a multivariate iv niche overlap approach. Evidence that B. ermanii, with its broad habitat niche but narrow regeneration niche, contrasted with A. mariesii, which has a broad regeneration niche but a narrow habitat niche, largely supports Bond’s hypothesis. The broad regeneration niche of A. mariesii likely stems from the large investment in seedlings through advanced regeneration; however, many of these individuals do not successfully reach maturity. B. ermanii has a narrow regeneration niche limited by suitable conditions in high light on raised surfaces with low litter, though over ontogeny the habitat niche broadens. In Chapter 4, I examined the influence of temperature, precipitation, PDSI on tree growth of B. ermanii and A. mariesii using a dendrochronological approach. B. ermanii responded positively to summer temperatures and negatively to summer precipitation, while A. mariesii had positive growth in the first year after warm and wet winters. 1.1% of the tree rings in B. ermanii were missing rings, possibly due to angiosperms’ responsiveness to changes in their environment. Missing rings may result from disturbance events. Though no known insect outbreaks or fires occurred in the Japanese Alps, missing rings may have been caused by volcanic activity. Understanding how and why the B. ermanii treeline exists is important for assessing responses to climate change as well as the cultural ecosystem services provided by alpine biodiversity. The climate-growth responses of B. ermanii and A. mariesii indicated that future warming, especially in winter, may reduce B. ermanii’s dominance at treeline. Warmer winter temperatures will likely increase A. mariesii’s growth and possibly reduce mechanical damage. Though some A. mariesii trees occur in the B. ermanii treeline and P. pumila mat, they frequently have mechanical damage (tops are sheared off at snow height, needles show browning and mortality). If A. mariesii establishes at treeline and above, then the B. ermanii treeline may be at risk of being replaced. The competition with P. pumila, mechanical damage of the A. mariesii, longevity of B. ermanii, and elevated nitrogen in the B. ermanii treeline are all factors that allow B. ermanii to persist in an environment where it is theoretically unexpected. v TABLE OF CONTENTS List of Figures .......................................................................................................................... v List of Tables ........................................................................................................................... vi Acknowledgements .................................................................................................................. vii Chapter 1 Introduction ............................................................................................................. 1 Theoretical Framework .................................................................................................... 2 Climatic Limitations ........................................................................................................ 4 Landscape Interactions ..................................................................................................... 5 Geomorphology ........................................................................................................ 5 Disturbance .............................................................................................................. 6 Alpine forests of central Japan ......................................................................................... 8 Climate ..................................................................................................................... 8 Species ...................................................................................................................... 9 Disturbance .............................................................................................................. 10 Summary .......................................................................................................................... 11 References ........................................................................................................................ 13 Chapter 2 Influence of topography and edaphic properties on broadleaf-deciduous versus evergreen conifers in a high elevation forest ................................................................... 27 Introduction ...................................................................................................................... 27 Methods ............................................................................................................................ 30 Study Area Description ............................................................................................ 30 Field Methods ........................................................................................................... 31 Statistical Analysis ................................................................................................... 33 Results .............................................................................................................................. 35 Topographic Properties ............................................................................................ 35 Shrubs ....................................................................................................................... 36 Light Environment ................................................................................................... 37 Soil Nutrients ........................................................................................................... 37 Canopy Structure ...................................................................................................... 38 Predictors of Importance Values .............................................................................. 39 Discussion ........................................................................................................................ 39 Canopy Structure ...................................................................................................... 40 Bedrock-Nutrients .................................................................................................... 41 Treeline Elevation .................................................................................................... 42 Nutrient Gradient...................................................................................................... 43 Microbial Activity .................................................................................................... 44 Nutrient Deposition .................................................................................................
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