title: Tree-ring δ13C and δ18O, leaf δ13C and wood and leaf N status demonstrate tree growth strategies and predict susceptibility to disturbance.
abstract: Understanding how tree growth strategies may influence tree susceptibility to disturbance is an important goal, especially given projected increases in diverse ecological disturbances this century. We use growth responses of tree rings to climate, relationships between tree-ring stable isotopic signatures of carbon (δ(13)C) and oxygen (δ(18)O), wood nitrogen concentration [N], and contemporary leaf [N] and δ(13)C values to assess potential historic drivers of tree photosynthesis in dying and apparently healthy co-occurring northern red oak (Quercus rubra L. (Fagaceae)) during a region-wide oak decline event in Arkansas, USA. Bole growth of both healthy and dying trees responded negatively to drought severity (Palmer Drought Severity Index) and temperature; healthy trees exhibited a positive, but small, response to growing season precipitation. Contrary to expectations, tree-ring δ(13)C did not increase with drought severity. A significantly positive relationship between tree-ring δ(13)C and δ(18)O was evident in dying trees (P < 0.05) but not in healthy trees. Healthy trees' wood exhibited lower [N] than that of dying trees throughout most of their lives (P < 0.05), and we observed a significant, positive relationship (P < 0.05) in healthy trees between contemporary leaf δ(13)C and leaf N (by mass), but not in dying trees. Our work provides evidence that for plants in which strong relationships between δ(13)C and δ(18)O are not evident, δ(13)C may be governed by plant N status. The data further imply that historic photosynthesis in healthy trees was linked to N status and, perhaps, C sink strength to a greater extent than in dying trees, in which tree-ring stable isotopes suggest that historic photosynthesis was governed primarily by stomatal regulation. This, in turn, suggests that assessing the relative dominance of photosynthetic capacity vs stomatal regulation as drivers of trees' C accrual may be a feasible means of predicting tree responses to some disturbance events. Our work demonstrates that a dual isotope, tree-ring approach can be integrated with tree N status to begin to unravel a fundamental question in forest ecology: why do some trees die during a disturbance, while other conspecifics with apparently similar access to resources remain healthy?
authors.affiliation: Department of Ecology and Evolutionary Biology, Kansas Biological Survey, University of Kansas, Lawrence, KS 66047, USA sharon.billings@ku.edu.
authors.forename: S A
authors.initials: SA
authors.lastname: Billings
authors.affiliation: Department of Ecology and Evolutionary Biology, Kansas Biological Survey, University of Kansas, Lawrence, KS 66047, USA.
authors.forename: A S
authors.initials: AS
authors.lastname: Boone
authors.affiliation: Department of Entomology, University of Arkansas, Fayetteville, AR 72701, USA.
authors.forename: F M
authors.initials: FM
authors.lastname: Stephen
mesh_terms: D001131:Arkansas; D017554:Carbon Compounds, Inorganic / Q000378:metabolism*; D002247:Carbon Isotopes / Q000032:analysis; D002482:Cellulose / Q000378:metabolism; D002980:Climate; D065928:Forests; D009584:Nitrogen / Q000378:metabolism*; D010103:Oxygen Isotopes / Q000032:analysis; D018515:Plant Leaves / Q000254:growth & development / Q000378:metabolism; D018547:Plant Stems / Q000254:growth & development / Q000378:metabolism; D029963:Quercus / Q000254:growth & development* / Q000378:metabolism*; D013440:Sulfides / Q000378:metabolism*; D014934:Wood / Q000254:growth & development / Q000378:metabolism
chemical_list: D017554:Carbon Compounds, Inorganic; D002247:Carbon Isotopes; D010103:Oxygen Isotopes; D013440:Sulfides; C092035:carbon sulfide; D002482:Cellulose; D009584:Nitrogen
keywords: Quercus rubra; forest disturbance; insect outbreak; leaf N; δ13C; δ18O