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Western Spruce Budworm Effects on Throughfall N, P, C Fluxes and Soil Nutrient Status in the Pacific Northwest

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Western Spruce Budworm Effects on Throughfall N, P, C Fluxes and Soil Nutrient Status in the Pacific Northwest Canadian Journal of Forest Research Western spruce budworm effects on throughfall N, P, C fluxes and soil nutrient status in the Pacific Northwest Journal: Canadian Journal of Forest Research Manuscript ID cjfr-2018-0523.R2 Manuscript Type: Article Date Submitted by the 02-Jun-2019 Author: Complete List of Authors: Arango, Clay; Central Washington University, Biological Sciences Ponette-González, Alexandra; University of North Texas System, Department of Geography and the Environment Neziri, Izak; Central Washington University, Biological Sciences Bailey, Jen;Draft University of North Texas System, Department of Geography and the Environment Keyword: herbivory, coniferous forest, outbreak insect, climate change, lepidoptera Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : https://mc06.manuscriptcentral.com/cjfr-pubs Page 1 of 45 Canadian Journal of Forest Research 1 Western spruce budworm effects on throughfall N, P, C fluxes and soil nutrient status in 2 the Pacific Northwest 3 4 Clay Arango1*, Alexandra Ponette-González2, Izak Neziri1, Jennifer Bailey2 5 6 1Department of Biological Sciences, Central Washington University, 400 E University Ave, 7 Ellensburg, Washington 98926-7537, USA 8 9 2Department of Geography and the Environment, University of North Texas, 1155 Union Circle 10 #305279, Denton, Texas 76203, USA 11 Draft 12 *Corresponding author: [email protected], (509) 963-3163, fax (509) 963-2730 13 1 https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 2 of 45 14 Abstract 15 Western spruce budworm (Choristoneura freemani) is the most widely distributed insect 16 herbivore in western North American coniferous forests. By partially or completely defoliating 17 tree crowns, budworms influence fluxes of water, nutrients, and organic carbon from forest 18 canopies to soils and, in turn, soil chemistry. To quantify these effects, throughfall water, 19 inorganic nitrogen (N), phosphorus (P), and dissolved organic carbon (DOC) concentrations and 20 fluxes, and soil N and P concentrations were measured in coniferous forest sites with high and 21 background levels of budworm herbivory. Throughfall N and P concentrations and fluxes 22 increased at high budworm sites during and/or immediately after larval stage budworm feeding, 23 indicating reduced uptake and/or greater leaching from canopies as a result of budworm activity. 24 Annual throughfall N fluxes (<67-71 g NDraft ha-1 yr-1) and soil N concentrations were low regardless 25 of herbivory level. In contrast, throughfall P was considerably greater at sites with high (2174 g 26 P ha-1 yr-1) compared to background (1357 g P ha-1 yr-1) herbivory, and this was reflected in 27 nearly 3-fold higher soil P concentrations at high budworm sites. Our findings suggest that by 28 altering throughfall chemistry and soil N:P, budworms could influence elemental export from 29 watersheds. 30 31 Keywords: herbivory, coniferous forest, outbreak insect 2 https://mc06.manuscriptcentral.com/cjfr-pubs Page 3 of 45 Canadian Journal of Forest Research 32 Introduction 33 Western spruce budworm (Christoneura freemani, hereafter WSB) is the most widely 34 distributed and destructive defoliator in western North American coniferous forests and a major 35 agent of forest disturbance (Fellin and Dewey 1982). In Washington State, insect forest damage 36 affected ~222,500 hectares in 2014, killing approximately 2.4 million trees by 2016, with nearly 37 all defoliation (~40,500 hectares) caused by WSB (WADNR 2016). The area affected by insects 38 is often similar to that burned by wildfires (156,000 hectares in 2014), underscoring the role of 39 insects in forest change. 40 Leaf-feeding canopy herbivores, such as WSB, influence the quantity and chemical 41 composition of water delivered to the soil in throughfall (water that falls through the canopy to 42 the soil; Stadler et al. 2001). First, throughDraft partial or complete defoliation, herbivores often 43 increase throughfall water flux (Michalzik 2011). Second, by fragmenting and damaging 44 foliage, herbivores stimulate leaching of organic and inorganic solutes into throughfall 45 (Michalzik 2011). Third, canopy herbivores deposit frass to canopies and soils during feeding 46 (Hunter 2001). Carbon-(C-) and nitrogen-(N-) rich frass in canopies and the litter layer can be 47 readily leached during the first seasonal rains (Hollinger 1986), increasing dissolved organic C 48 (DOC) and N fluxes into soils (Michalzik 2011). Increased DOC fluxes have, in turn, been 49 shown to fuel leaching of DOC and dissolved organic nitrogen (DON) from the forest floor 50 (Michalzik et al. 2001) and to promote microbial immobilization of N and phosphorus (P) 51 (Michalzik and Stadler 2005). DOC and frass-derived nutrients have also been found to 52 accelerate soil N transformation rates (Huber 2005). For example, in coniferous forests with 53 pine bark beetles, greater inorganic soil N levels after outbreak contributed to more rapid N 54 mineralization and nitrification (Griffin and Turner 2012). Reduced nutrient uptake by 3 https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 4 of 45 55 weakened trees (Frost and Hunter 2007) and tree death in stands experiencing herbivory can lead 56 to increased soil N and P as well via reduced assimilatory uptake and increased litter 57 decomposition (Mikkelson et al 2013). Both increased transformation rates and reduced nutrient 58 uptake in forests experiencing outbreak conditions have the potential to increase nutrient 59 leaching losses as much as 30 fold (Houle et al. 2009). While it is clear that herbivores can alter 60 throughfall chemistry and nutrient cycling in forest soils, ecosystem responses are highly 61 variable, and the magnitude and direction of herbivore impact is dependent on the system and the 62 environmental context (Hunter 2001). 63 Despite the persistent role of WSB as a disturbance agent in western North America, we 64 know of no research that has investigated WSB ecosystem effects in seasonally dry coniferous 65 forests in this region. WSB is a native, Draftoutbreak lepidopteran whose larvae feed mostly upon 66 freshly grown needles of Douglas fir and grand fir trees (Alfaro 2014). As an endemic 67 defoliator, WSB always exists at background levels (i.e., during non-outbreak years). However, 68 during outbreak years, these herbivores can reach densities high enough to defoliate tree crowns 69 within a season and to completely strip trees of their needles during a multi-season outbreak 70 (Zhao 2014). Dendrochronological analysis of the past three centuries shows that historic 71 outbreaks occur at the end of regional droughts, last up to ten years, and tend to be synchronized 72 across broad areas (Flower et al. 2014). 73 The WSB life cycle is tied closely to the seasonality of western coniferous forests (Nealis 74 2012). With warming spring temperatures in mid- to late-May, budworm larvae emerge from 75 hibernacula about two weeks prior to budburst and begin dispersing through the canopy. After 76 budburst in late May to mid-June, the larvae begin mining into buds and new needles mostly at 77 the top of the tree crown and on the fringes of branches. Needles begin to turn an orange color 4 https://mc06.manuscriptcentral.com/cjfr-pubs Page 5 of 45 Canadian Journal of Forest Research 78 as the larvae feed and grow. After 30-40 days of continuous feeding in June and July, sixth 79 instar larvae construct a pupal case on the underside of the branches from which adults emerge 80 after about 2 weeks in mid-July to early August. Adults disperse to mate and lay eggs, dying 81 shortly thereafter. Larvae emerge from eggs after about 10 days whereupon they immediately 82 seek shelter in the bark by building hibernacula to protect them over the winter. 83 Coupled with decades of fire suppression that have increased forest basal area and 84 density, projections of an ever-warming climate suggest that drought stress in overstocked 85 western coniferous forests (Dalton et al. 2013) will further amplify favorable conditions for 86 WSB. Future WSB outbreaks are predicted to increase in frequency, intensity, and spatial extent 87 (Bentz et al. 2010), with unknown ecosystem effects. Given these projected changes in WSB 88 disturbance, we quantified throughfall andDraft soil chemistry under canopies with high and 89 background levels of WSB herbivory. Specifically, we hypothesized that net throughfall N, P, 90 and DOC fluxes would be higher under canopies with high compared to background levels of 91 WSB herbivory due to increased throughfall water fluxes and accelerated canopy leaching. We 92 also hypothesized that soil inorganic N and P concentrations would mirror the patterns observed 93 in net throughfall. We anticipate that these findings will shed light on how changes in WSB 94 populations could affect nutrient cycling in western coniferous forests under future climate 95 change scenarios. 96 97 Study Area and Site Selection 98 We conducted this study in the Okanogan-Wenatchee National Forest and the Teanaway 99 Community Forest, both located in the rain shadow (east slope) of the Cascade Range in central 100 Washington State. This region is characterized by a continental climate with dry summers (May- 5 https://mc06.manuscriptcentral.com/cjfr-pubs Canadian Journal of Forest Research Page 6 of 45 101 September) and wet winters (October-April). Long term climate data from Blewett Pass, a 102 weather station about 4 km from our study sites, shows that mean annual precipitation is 894 mm 103 with ~86% falling between October and April, mostly as snow from November through March 104 (National Oceanic and Atmospheric Administration (NOAA) 2018). Although peak 105 precipitation is concentrated between October and April, total precipitation at any individual site 106 in this mountainous region can vary considerably by aspect and elevation. Mean monthly 107 temperature is -1.8°C during the winter (Dec, Jan, Feb) and 15.2°C during the summer (Jun, Jul, 108 Aug) (NOAA 2018).
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