Intraspecific Morphological Root Trait Variation Is Less Affected by Soil Fertility Than Biomass Allocation Or Root Nutrient Concentrations
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Root traits of New Zealand trees: Community-level responses to a soil fertility gradient, and plastic responses to nutrient availability A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science in Biological Sciences at The University of Waikato by Kris Rushton Kramer-Walter 2015 Abstract The study of plant traits has great application for understanding plant distribution patterns and community assembly at a variety of scales. Roots are a vital component of plant water and nutrient uptake strategy, and yet root traits are not as well understood as leaf or stem traits. In this thesis I aimed to determine the relationships between seedling fine root, leaf, and stem traits of New Zealand tree species, and investigate whether these traits related to species‟ abundance along a soil fertility gradient. I also investigated how seedling traits compare with adult traits, and whether nutrient availability or method of fertiliser application affected seedling trait expression. To achieve these goals, I measured root, leaf, and stem traits on seedlings of 66 native tree species, and combined this with relative abundance data along a strong soil fertility gradient at Puketī forest. I used principle component analysis and ordination to assess the dimensionality of trait variation across species; and linear regressions to compare community-weighted mean (CWM) traits vs. a summarised axis of soil fertility. I also performed regression analysis between seedling traits and adult traits from the literature. To assess the effects of nutrient availability and application I grew four native tree species under three nutrient treatments: low, pulsed, and slow release. I used two- way ANOVA and co-efficient of variation analyses to determine the strength of responses to nutrient treatments. Dry matter content was positively associated across leaves, roots, and stems, and negatively associated with root nitrogen concentration and relative growth rate, suggesting that at least as seedlings, traits associated with a fast or slow growth strategy are co-ordinated across organs. Root diameter and SRL were independent from this axis of fast-slow growth, suggesting that SRL does not have a direct effect on seedling growth rates and nutrient foraging. A third axis of variation was also identified, strongly influenced by root phosphorus and nitrogen concentrations, but was difficult to interpret. Regression of CWM traits vs. soil fertility showed that traits which comprised the fast-slow PCA axis co-varied strongly with soil fertility. Neither root diameter nor SRL were significantly related to soil fertility, supporting the conclusion that SRL is not adaptive to nutrient foraging ability. Seedling morphology traits are generally well correlated with adults, but tend to be oriented towards a more “acquisitive” growth strategy, suggesting that species may down-regulate their growth over ontogenetic i development. The absence of correlation between SRL and soil fertility suggests that alternative root traits may be more applicable for understanding species foraging strategy. Species‟ responses to the nutrient treatments differed for most traits. Root-to- shoot ratio, growth rate, and root nutrient concentrations responded strongly to nutrient availability, and morphological traits did not respond as strongly. Slow release and pulse treatments were typically similar, and both significantly different to low nutrient treatment. These results suggest that it is practical to compare morphological trait data between studies, provided plants are raised in environments conducive to growth, but that nutrient concentrations and biomass allocation traits can be strongly influenced by soil fertility. ii Acknowledgements I would like to thank all the academic staff, family, and friends who have helped me to achieve this goal. Firstly, thank you to Daniel Laughlin for suggesting this project, and for all the guidance and advice along the way. I would like to thank Lynne Baxter for her help and advice in setting up and maintaining all the plants in the glasshouses. Thank you also to Colin Monk for his help in the laboratory. A huge thank you Toni Cornes and Adam Purcell for their assistance in processing and grinding roots when I had both of my arms in casts. I can‟t even begin to express my gratitude to my partner Catherine Walden, and my dad, Phil Kramer-Walter, for coming to my aid and getting stuck in with the messy job of root washing. I would also like to thank Melissa Jager, who laid the groundwork for this research, and gave me great advice during my study. Thank you to the nurseries across the country who supplied the seedlings for my experiments: Forest Flora, Oratia Nursery, Taupo Native Plant Nursery, and Pukerau Nursery. A special thank you to Wayne and Julia from Forest Flora, who gave me great plants, great advice, and great experience doing restoration planting. I would also like to thank the scholarship and funding bodies which have aided me in this project: The University of Waikato, Waikato Botanical Society and the Royal Society of New Zealand Marsden Fund (12-UOW-041). Thank you to my Mum and sisters for being such greenies, and inspiring me to study ecology. Thank you to my flatmates and labmates for providing great company throughout my University journey. Finally, than you again to my partner Cat, for being there whenever I needed you, and taking such good care of me. iii Table of Contents Abstract ............................................................................................................... i Acknowledgements ........................................................................................... iii Table of Contents .............................................................................................. iv List of Figures ................................................................................................... vi List of Tables ................................................................................................... viii Chapter 1: Literature review ............................................................................ 1 1.1 Chapter overviews ................................................................................ 11 1.2 References............................................................................................ 13 Chapter Two: A multi-trait test of the whole-plant economics spectrum using fine root, leaf, and stem traits and community-level trait responses to soil fertility ................................................................................................. 23 2.1 Abstract ................................................................................................ 23 2.2 Introduction .......................................................................................... 25 2.3 Methods ............................................................................................... 30 2.4 Results ................................................................................................. 46 2.4.1 Correlations between seedling and adult traits................................... 46 2.4.2 Plant trait dimensionality and the whole-plant economic spectrum.... 49 2.4.3 Community-level root traits along a soil fertility gradient ................. 52 2.5 Discussion ............................................................................................ 55 2.5.1 Relationships between traits of seedling and adult plants .................. 55 2.5.2 The whole-plant economics spectrum (PC1) ..................................... 57 2.5.3 Root diameter and SRL (PC2) .......................................................... 59 2.5.4 Root phosphorus (PC3) ..................................................................... 62 2.5.5 Root traits selection along a soil fertility gradient ............................. 64 2.6 Recommendations ................................................................................ 66 2.7 References: .......................................................................................... 68 Chapter Three: Intraspecific morphological root trait variation is less affected by soil fertility than biomass allocation or root nutrient concentrations ................................................................................................ 79 3.1 Abstract ................................................................................................ 79 3.2 Introduction .......................................................................................... 81 3.3 Methods ............................................................................................... 85 3.4 Results ................................................................................................