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Article

First-Year Mortality of Four Early-Successional Species on Severely Degraded Sites in Eastern Canada as Influenced by a Factorial of Site Preparation Treatments

by
Dominic Galea
and
John E. Major
*
Natural Resources Canada, Canadian Forest Service—Atlantic Forestry Centre, 1350 Regent St., Fredericton, NB E3B 5P7, Canada
*
Author to whom correspondence should be addressed.
Forests 2024, 15(1), 143; https://doi.org/10.3390/f15010143
Submission received: 9 November 2023 / Revised: 23 December 2023 / Accepted: 5 January 2024 / Published: 10 January 2024
(This article belongs to the Section Forest Ecology and Management)
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Abstract

:
Barren, severely disturbed sites lacking soil, such as mine sites and waste deposit sites, present severe challenges to ecological service restoration because of high temperatures, solar radiation, and wind speeds; extreme temperature changes; and low soil moisture and nutrient availability. An ecological restoration experiment using three site preparation treatments was conducted. Straw (S), Meri-Crusher (MC), and coarse woody debris (CWD) were assessed in a site, no site preparation 2 × 2 × 2 factorial, including a control treatment, on sites barren for 25 years. In addition, four early-successional species: white birch (WB, Betula papyrifera Marsh), gray birch (GB, Betula populifolia Marsh), green alder (GA, Alnus viridis Vill. ssp. crispa Ait) and speckled alder (SA, Alnus incana L. ssp. rugosa Du Roi), were examined for mortality. Mortality was measured after three time periods, summer-related 2021, winter-related, and frost heave mortality (spring 2022). Summer-related mortality was predominantly influenced by S treatments (reduced mortality) and their interactions. Straw’s ability to retain moisture strongly suggests it mitigated summer-related drought mortality. S interactions were not rank changes but magnitude effects. The species × straw interaction showed that SA had the greatest magnitude difference, with 25% and 3.6% summer-related mortality for NS and S treatments, respectively. SA, a hydrophilic species, accounted for nearly half the total summer-related mortality, and there were strong species effects and species interactions. The full combination of site preparation treatments had the lowest summer-related mortality, at 1%. Winter-related mortality only affected 1.9% of the total sample size, and there were no species effects or interactions, but contrary to other results, S was the leading cause of mortality due to fungal presence found on expired seedlings. For frost heave mortality, it was clear that the S treatment was effective, with 1.2% and 20.7% overall mortality for S and NS, respectively. MC alone had the greatest negative effect, with 46.9% frost heave mortality; however, when interacting with S or CWD, the mortality decreased substantially. Frost heave had no species interactions and only a species effect, with SA having the greatest mortality. Over the first full year, MC alone and control had the greatest mortality, with 60% and 38%, respectively, after one year. Overall, one-year mortality showed S reduced mortality by 27% and CWD by 19%, while MC increased mortality by approximately 4%. When treatments were combined in any way, mortality dropped significantly, showing an additive effect, with the three-combination treatment resulting in the lowest one-year mortality, of only 3.1%. Straw provided the strongest effect, both as an effective barrier to moisture evaporation, providing up to 10% more soil moisture under dry conditions and provided an effective thermal barrier that substantially reduced the frost heave mortality. Even early-successional species such as WB, GB, GA, and SA need site preparation treatments to establish and survive the first year on long-term barren lands.

1. Introduction

Barren, severely disturbed sites lacking soil, such as mine sites and waste deposit sites, present severe challenges to ecological service restoration because of high temperatures, solar radiation, and wind speeds; extreme temperature changes; and low soil moisture and nutrient availability. Rapid erosion from these sites creates more problems by removing soil and increasing sedimentation in local waterways, negatively affecting aquatic environments [1,2]. Severely disturbed soils are often compacted and lack water holding capacity and soil nutrients [3,4,5]. Thus, successional restoration should start with some development of soils and early-successional native species, which should allow sites to eventually return to natural successional processes [6]. Returning nutrients to the soil and eventually ameliorating the coarse substrate into a forest supporting substrate could be accelerated by using early-successional hardwood species, as it has been shown that nutrient content and organics within hardwood forests are often greater than in conifer forests [7,8]. Reintroduction of original or new topsoil to barren sites can appear to be a promising method to ameliorate poor sites; however, there is a very high likelihood of more erosion [9].
Seedlings in barren landscapes are often exposed to severe climatic conditions. Excessive heat and drought in the summer, lack of nutrients, frost heave in shallow soils, and cold damage in the winter are unfavorable conditions for plant survival. Shallow soils lack water holding capacity and dry rapidly, resulting in seedling mortality. In addition, shallow soil depth to bedrock and consequently a high water table increase susceptibility to frost heave from greater water concentrations near the soil surface [10,11,12]. Frost heave has two main effects. First, young plants may be damaged or killed from root exposure. Second, soils that have been frost heaved are susceptible to erosion [13,14]. According to Graber [15], four types of damage can occur to seedlings: (1) heaved out, with lethal exposure of root system, (2) partial heave, with exposed root damage commonly found, (3) girdling of plant stems via mechanical damage caused by substrate abrasion on the epidermis and cambium, and (4) seedling decapitation: separation of plant parts such as the roots from the aboveground plant parts.
Early-successional species are known for their ability to grow in post-disturbance landscapes [16,17]. With their need for full sunlight, they tend to grow rapidly, which can help capture sites quickly [16,18,19]. We examined four such species: white birch (WB, Betula papyrifera Marsh), gray birch (GB, Betula populifolia Marsh), green alder (GA, Alnus viridis Vill. ssp. crispa Ait) and speckled alder (SA, Alnus incana L. ssp. rugosa Du Roi). All four species are classified as early-successional, relatively fast-growing species with relatively short lifespans [20]. All four species have some histories in restoration efforts due to their ability to withstand and reclaim harsh sites [16,18,19,21,22,23]. White birch prefers deep, well-drained, rich soil, but can still compete in a wide range of soil types ranging from gravels to silts and organic bog to peat soils [24]. Gray birch grows best on a moist, well-drained site, but is quite adaptable and can grow well on dry sandy or even gravelly soil [25,26]. Green alders prefer moist sites but can tolerate dry sites with soil types from gravelly to sandy and will regularly be found on upland glacial till sites [27]. Speckled alders compete best on wet sites, such as beside waterways, bogs, and fens, but will also establish in mires with peaty soils [28,29]. In Minnesota, it was noted that speckled alder will grow in mesic to moist sites but was found more in moist sites [30].
In this study, we examined a factorial of three site preparations: (1) site crusher to create a loose aggregate using a Meri-Crusher (MC) (MJ hydraulic 1.4 ST Meri-Crusher, Voukatti, Finland), (2) straw (S) coverage, and (3) coarse woody debris (CWD) coverage. The site crusher treatment involved mechanically ripping the top layer of aggregate to create a substrate that can be used to restore terrestrial ecosystems with little to no existing substrate available [31,32]. Breaking in situ soil beds is better than replacing with soil from elsewhere [33], and benefits of ripping the soil include improved aeration, water infiltration, and root penetration [34]. Straw coverage can affect the processes occurring in the top layer of the soil, such as reducing surface evaporation [35], improving water infiltration [36], and increasing moisture retention [37,38,39], and can act as an insulator and regulate extreme temperatures in the winter to help prevent frost heave of planted seedlings [40]. Coarse woody debris can create protected microsites and enhance biodiversity on a reclaimed site [41]. The addition of CWD has been shown to decrease water loss and seedling desiccation by providing shelter from the elements [42]. There is a preponderance towards larger CWD, as smaller pieces like mulch can draw what little nitrogen is available from the soil via different fungi to fuel decomposition [43].
Our goal was to quantify seedling mortality from the effects of three site preparation treatments in a site, no site preparation 2 × 2 × 2 factorial design resulting in eight separate treatments including control, testing four early-successional species. We hypothesized that the presence of a site preparation treatment will have a positive effect on the survival and establishment of the seedings. We also hypothesized that when site preparation treatments were used together, their effects might be additive. In addition, we hypothesized that there will be species differences in survival. Our objective was to quantify mortality (1) over the first summer, (2) over the following winter, and (3) mortality due to frost heave. In addition, we examined the combined full first-year mortality to assess the best practices and the best species to begin the restoration and forest succession process on highly disturbed sites.

2. Materials and Methods

2.1. Study Area

The study area is located within the Department of National Defence, CFB Gagetown, NB, Canada (45.68108, −66.50179). The study area had three sites located at an average altitude of 87 m above sea level and lies on a small western aspect. Average annual precipitation is 1376 mm year−1, with most of it falling in summer to winter months. Rainfall ranges from 94 mm in March to 133 mm in June. Snowfall during the winter months ranges from 41 cm in March to 72 cm in February. Average temperature for the hottest months (July and August) is around 20 °C and for the coldest months (January and February) −9 °C. Soil is predominantly parent material consisting of felsic pebble conglomerates and mixed igneous rocks. Surrounding forest is dominated by gray birch (Betula populifolia), red oak (Quercus rubra), and trembling aspen (Populus tremuloides).

2.2. Site Preparation Treatments

Three separate sites were selected at CFB Gagetown, NB, Canada (45.68108, −66.50179). A randomized site preparation design with site, no site preparation factorial of: S, NS; MC, NMC; and CWD, NCWD in a 2 × 2 × 2 factorial resulted in eight treatments, including a control. Note, a prefix of N on the site preparation treatment indicates no such treatment. The eight-treatment factorial was (1) NS × NMC × NCWD, the control, (2) S × NMC × NCWD, (3) NS × MC × NCWD, (4) NS × NMC × CWD, (5) S × MC × NCWD, (6) S × NMC × CWD, (7) NS × MC × CWD, and (8) S × MC × CWD. The eight treatments were replicated (blocked) four times per site; this equated to 32 plots per site, on three sites, resulting in 96 plots in the experiment. The MC treatment was created using a John Deere 200D excavator equipped with an MJ hydraulic 1.4 ST Meri-Crusher (Voukatti, Finland). The MC plots produced a 3.2 m by 3.2 m square of broken aggregate from loose soil, loose rock and solid rock substrate. Straw bales were used to distribute straw over approximately 18 m2 of area or roughly 2 experimental plots to a depth of 5 cm. The CWD was sourced from Department of National Defence (DND) near Gagetown, NB, using primarily GB. All CWD was processed into ~3 m length to match the MC plots. An average volume of 0.203 m3 ± 0.020 m3 of CWD was applied to 48 plots. The placement of CWD was performed last to hold the straw in place, and the CWD was distributed over each plot in a north to south orientation to provide shade at different times of the day.

2.3. Plant Materials

Seedlings from the four tree species were grown from seedlots bulked from seed collected from sources listed in Table 1. WB, GB and GA were grown during 2020 at the Atlantic Forestry Center (AFC), Fredericton, NB (45.93501, −66.65745); SA were grown during 2020 at Scot and Stewart nurseries in Antigonish, Nova Scotia (45.53292, −61.86888). The seedlings grew in trays with root container volume of 110 mL. In January 2021, 1-year-old seedlings were cut to a height of roughly 10 cm to reduce the root-to-shoot ratio for these harsh sites and had a root-to-shoot ratio of 1:1 by dry weight. Seedlings were packed into boxes for frozen storage at −4 °C in AFC freezers. The seedlings were removed from frozen storage 3–4 days prior to planting to thaw out. On 7–8 May 2021, four individual replicates per species were randomly planted per plot, 32 plots per site, thus 128 individual species per site, and 384 individual species across the three sites. The seedlings were planted with narrow shovels at 0.5 × 0.5 m spacing. The experiment contained 1536 total planted seedlings across the three sites.

2.4. Soil Properties

For each plot, three depths to bedrock measurements were made using an iron rod hammered into the soil until it contacted the solid rock substrate. Soil samples were collected in October 2020 from 3 sites × 2 treatments MC, no MC (NMC) × 4 block replicates for a total of 24 total soil samples. Soil analysis included texture sand, silt, clay, and rock (%), and soil macronutrient analysis including organic matter, carbon (C), and nitrogen (N) using phosphorus (P) available potassium (K), calcium (Ca), magnesium (K), concentration, and pH by the Laboratory for Forest Soils and Environmental Quality at the University of New Brunswick using techniques as outlined in Major et al. 2022 [44]. Soil bulk density assessment was performed from 16 plots per site, 8 meri-crushed plots and 8 non-meri-crushed plots, for a total of 48 samples in the fall of 2022. Table 2 provides a summary of soil properties among sites. Soil moisture (%) was measured periodically using an ML3 Theta Probe soil moisture sensor (Cambridge, UK), by recording three readings per plot in two randomly selected blocks of each site, varying site order each time they were taken. Soil moisture readings were taken on 13 collection dates ranging from 3 March 2021 to 9 September 2021.

2.5. Mortality Assessment

Percent mortality per species per plot was determined from the four replicates per plot. Mortality assessments were performed at the end of August 2021 for summer-related mortality, and April 2022 for frost heave mortality. If a root plug lifted 25% or more and had died, it was considered frost heave mortality. A large majority were completely heaved from the ground. If the root plug lifted less than 25% and died, it was considered part of the winter mortality. Winter-related mortality was largely due to fungal presence found on expired seedlings. The difference between summer-related mortality 2021 and what was not due to spring frost heave mortality 2022 was considered winter-related mortality.

2.6. Statistical Analysis

The experiment was established as a randomized block design. Initial soil properties were subjected to analyses of variance (ANOVA); site and MC were fixed effects, and block was a random effect. The following ANOVA model was used:
Yijk = µ + Bi(Sj) + Sj + Mk+ SMjk + eijk
where Yijk is the dependent soil property trait of the ith block, jth site, and the kth MC treatment, and µ is the overall mean. Bi(Sj) is the random effect of block nested within site (i = 1, 2, 3, 4), Sj is the effect of the jth site (j = 1, 2, 3), Mk is the effect of the kth MC treatment (k = 1, 2), SMjk is the interaction effect between jth site effect and kth MC treatment effect, and eijk is the random error component.
Available soil moisture in 2021 was subjected to analyses of variance (ANOVA); site, S, MC and CWD were fixed effects, and block was a random effect. The following ANOVA model was used:
Yijklm = µ +Bi(Tj) + Tj + Sk +Ml + Cm + TSjk + TMjl + TCjm + SMkl + SCkm + MClm + TSMjkl + TSCjkm + TMCjlm + SMCklm + eijklm
where Yijklm is the dependent soil moisture availability of the ith block, the jth site, kth S treatment, lth MC treatment, and mth CWD treatment, and µ is the overall mean. Bi(Tj) is the random effect of the ith block nested in the jth site (i = 1, 2, 3, 4), Tj is the effect of the jth site (j = 1, 2, 3), Sk is the effect of kth S treatment (k = 1, 2), Ml is the effect of lth MC treatment (l = 1, 2), Cm is the effect of the mth CWD treatment (m = 1, 2), and eijklm is the random error component. Interactions of the single effects are included up to three-way interactions; four- and five-way interactions are folded into the error term.
Summer-related, winter-related, frost heave, and total first year mortality were analyzed using analysis of variance (ANOVA). MC, S, CWD, species, and site were all considered as fixed effects. Block was a random effect. The following ANOVA model was used:
Yijklno = µ + Bi(Tj) + Tj + Pk + Sl + Cn + Mo + TPjk + TSjl + TCjn + TMjo + PSkl + PCkn + PMko + SCln + SMlo + CMno + TPSjkl + TPCjkn + TPMjko + TSCjln + TSMjlo + TCMjno + PSCkln + PSMklo + PCMkno + SCMlno + eijklno
where Yijklun is the dependent mortality factor of the ith block, within the jth site, of the kth species, in the lth S treatment, nth CWD treatment, oth MC treatment, and µ is the overall mean. Bi(Tj) is the random effect of block nested in site (i = 1, 2, 3, 4), Tj is the effect of the jth site (j = 1, 2, 3), Pk is the effect of the kth species (k = 1, 2, 3, 4), Sl is the effect of the lth S treatment (l = 1, 2), Cn is the effect of the nth CWD treatment (n = 1, 2), Mo is the effect of the oth MC treatment (o = 1, 2), and eijklno is the random error component. Interactions of the single effects are included up to three-way interactions; four- and five-way interactions are folded into the error term.
Percent mortality was arcsine, square root transformed to fulfill normality. All effects were considered significant at the p = 0.05 level, though all p-values are presented for individual consideration. Assumptions of independence of data, normality of distribution, and normality of variance were met prior to assessing each statistical analysis (ANOVA). The general linear model from Systat Software Inc. (Chicago, IL, USA) was used for analysis. If significant, site and species were post hoc tested using Tukey mean separation test (p = 0.05). Significant interactions are referred to as either rank change interaction, or magnitude effect interaction. A rank change interaction occurs when one effect has a greater mean under one scenario and lower mean under another scenario. A magnitude effect would refer to when the effect mean is always greater than under both scenarios, but of different magnitudes.

3. Results

There was little or no difference in soil nutrients or texture between MC and NMC plots, so they were folded into the site-only analysis. Additionally, soil bulk density showed no significant difference between sites or MC and NMC plots. Soil analysis in fall 2020 showed greater total N in site 1 compared to sites 2 and 3 (Table 2). Site 1 had a significantly lower Ca content and pH than sites 2 and 3. Two soil texture properties were significantly different: silt, which was greater on site 1 (43.8%) than on sites 2 and 3, and sand, which was greater on site 3 (58.2%) than on sites 1 and 2.
Soil moisture content corresponded with the precipitation graph (Figure 1A). When the greatest rainfall occurred (1 July onwards), the average soil moisture content increased. Dry conditions from 3 June to 30 June resulted in significantly lower soil moisture content for NS than for S treatments. As the soil accumulated more rain, the effect of straw was not significant.
All summer-related mortality main effects excluding CWD were significant (Table 3). Straw was highly significant and accounted for the greatest amount of variance (9.2%) in summer-related mortality and appeared in most other significant interactions that were magnitude effects.
The significant MC × S × site (p = 0.003) interaction (Table 3) for summer-related mortality was a result of a single site rank change on only NS plots. Site 3 had the greatest summer-related mortality on NS, NMC treatments (25.8%), whereas it had low summer-related mortality on NS, MC treatment plots (7.0%) compared to site 2 (21.0%) (Figure 2A). The S × site × species interaction in summer-related mortality was largely due to a magnitude effect under NS. Site 2 NS had the greatest mortality particularly for SA (45.3%), which is also seen in Figure 2B, whereas sites 2 and 3 were somewhat equal in summer-related mortality for the other species.
The significant MC × S × CWD interaction in summer-related mortality was a result of a single rank change again under NS (Figure 3). Under the NS, CWD treatment, NMC plots had greater summer-related mortality than MC plots, with 20.7% and 7.8%, respectively. Summer-related mortality was reversed to a lesser degree under the NS, NCWD plots, with 11 and 14%, respectively.
Overall, S and NS had 2.7% and 13.2% summer-related mortality, respectively. Site 1 mortality of 3.3% was significantly lower than that of sites 2 and 3 with 11.1% and 9.4%, respectively.
Winter-related mortality had the smallest mortality impact, resulting in the loss of only 36 seedlings, affecting only 1.9% of the total seedlings. Winter-related mortality had only one three-way interaction, site × S × CWD interaction (Table 4), and it was a small rank and largely a magnitude effect driven by S on site 2 (Figure 4).
S, NCWD on site 2 had the greatest mortality, with 12% winter-related mortality (Figure 4). This was also driving the two significant two-way interactions: site × CWD and S × CWD (Figure 5A,B). Overall, S and NS resulted in winter-related mortality of 1.7% and 4%, respectively. Overall, CWD, NCWD had 1.6% and 3.6% winter-related mortality, respectively. MC, NMC had a winter-related mortality of 3.5% and 1.7%, respectively.
There was only one frost heaved mortality three-way interaction (Table 5); it was largely a magnitude effect among the three site treatments and was driven by the large frost heaved mortality under NS, NCWD with NMC, MC having 25% and 47% mortality, respectively (Figure 6).
Frost heaved mortality with all other treatment combinations was 5% or less, with S, CWD having zero frost heaved mortality. The three two-way interactions S × MC, CWD × MC and CWD × S were largely magnitude effects found in the three-way interaction figure (Figure 6). The one other two-way interaction site × S was also a magnitude effect, with sites 2 and 3 having the largest magnitude with site 1 lesser (Figure 7).
ANOVA for frost heaved mortality showed that all main effects were significant (Table 3). Straw accounted for the most variance in the model (19%), and CWD had the second largest variance (12.9%). Overall, S vs. NS had 1.1% and 20.7% frost heaved mortality, CWD, NCWD had 2.5% and 19.4%, and MC, NMC had 13.4% and 8.5%, respectively. The species that was affected the most was SA, with 14.8% mortality, then green alder at 11.7%, and both birches at 8.6%. Site was significant because of the reduced mortality on site 1 (6.8%) while sites 2 and 3 had increased mortalities of 12.9% and 13.1%, respectively.
There was also only one total first-year mortality three-way interaction (Table 6), and it was largely a magnitude effect, with some rank change, and it was again among the three site treatments and driven by the large frost heave mortality findings.
Again, under NS, NCWD with NMC, MC had 39 and 60.9% mortality, respectively (Figure 6). The three-way interaction MC × S × CWD (p < 0.001) (Figure 8) showed trends very similar to that discussed in the frost heaved model, with MC alone having the greatest mortality (60.9%). Since spring 2022 mortality was the sum of the past three mortalities, it followed similar trends. MC presence was largely negative by itself without other treatments; however, once CWD or S was added, MC had a positive effect.
The species × straw interaction (p < 0.001) (Figure 9A) was a magnitude effect. When S was present, all species mortality stayed under 10% (Figure 9A), but when S was not present, there was an average 21.3% increase in mortality for WB, GB, and GA, and the greatest increase of 45.4% mortality in SA, which resulted in the greatest mortality through the year of around 56.3% (Figure 9A). Straw × site interaction (p < 0.001) was also due to a magnitude effect. The lack of S on site 1 (16.8%) had a more muted effect when compared to site 2 and 3 at 46.9% and 41%, respectively (Figure 9B).
After the first full year, the overall survival of the seedlings was significantly affected by all main effects, excluding MC (Table 3). However, MC was significantly present in two-way interactions with CWD (p < 0.001) and site (p = 0.007) and also a three-way interaction with S and CWD (p = 0.001). Like the other models, S had the highest variance component (19.1%), and it was present in four of the six significant interactions. Overall, S and NS had 7.7% and 34.9% total mortality, respectively. CWD and NCWD had 11.8% and 30.7% total mortality. Sites 1, 2 and 3 overall had 11.3%, 27.9%, and 24.6% total mortality, respectively, by spring 2022.
Total mortality was greatest in SA, with 33.6% (Figure 10A). WB, GB, and GA all had under 25% mortality, with GA being at exactly 20% mortality through the first year. Frost heave was the largest contributor to mortality regardless of species, with WB, GB, GA, and SA experiencing 8.6%, 8.6%, 11.7%, and 14.8% mortality, respectively, across all sites. The effect of winter-related mortality was relatively minor, with only 36 cases. Of all species, 1.9% were affected, with most mortality occurring in SA, with 4.4% of all SA dying (Figure 10A). When broken down by treatments in Figure 10B, the addition of S greatly limited the high level of frost heaved mortality, as did the addition of CWD but to a lesser extent. The treatment of MC alone has the greatest mortality, with 60.9% of all MC plots succumbing to 13.5% summer-related mortality and 46.9% frost heave mortality. An additive effect could be seen where the more treatments being added reduced the mortality. The treatment of MCSCWD has the lowest mortality rate, with only 3.1% of the species dying within that plot type (Figure 10B).

4. Discussion

4.1. Summer-Related Mortality

Straw alone had the largest summer-related mortality effect, accounting for 9.2% of the total variance and resulting in an overall 10.5% reduction in summer-related mortality when S was present. Straw can reduce summer moisture evaporation [45,46,47], increase soil water retention [37], reduce erosion [46,48], regulate surficial soil temperatures [35], increase soil enzyme activity [49], and reduce weeds that may compete for above and below ground resources [38]. Straw also reduces solar radiation reaching the soil surface, lowering soil temperatures during early growth stages [35,40]. All species were significantly better on site 1 than on either site 2 or 3. Site 1 had deeper soil and hence had better soil moisture conditions compared to sites 2 and 3; these conditions allowed all species to survive better regardless of the treatments. Straw effects differed depending on the site. Site 2 had a shallower soil, more visible bedrock, and was more open to the elements, and that is where S had the greatest impact.
The three-way site preparation treatment interaction that included NS, NMC and CWD had ~10% greater summer-related mortality than control plots. This result was somewhat unexpected; however, other combinations with CWD decreased summer-related mortality (see below). CWD can often create microsites and enhance future biodiversity on reclaimed sites [41]. The addition of CWD decreases water loss and seedling desiccation by providing shelter from the elements [42,50]. However, the presence of CWD is not always homogenous since its physical properties and rate of decay differ both in time and among different forest communities [51,52,53]. While the immediate effects of CWD alone seem to have hindered the summer-related survival of the planted species, the evidence points towards a propensity for a more favorable environment once the CWD begins decomposing and amassing organics within the plots [54]. A theory as to why it may have heightened summer-related mortality in CWD plots is that total area of good planting spots is reduced with CWD, as it takes up space on plots of limited size and available planting area. This issue was more important on site 2, which was already somewhat limited in available planting spots due to its shallow depth to bedrock.
As demonstrated above, MC interactions were dependent on other site preparation treatments. Treated with S or CWD, there was very little summer-related mortality. MC by itself had 3% greater summer-related mortality than control plots. The benefits of ripping the soil with the MC treatment include improved aeration, water infiltration, and root penetration [34]. Loose aggregate also enhances moisture infiltration and retention but instead of controlling erosion, the churned earth was raised above the surrounding ground layer and probably suffered from increased rain and wind erosion. The MC plots without the implementation of S suffered too much water loss since the breaking of the aggregate created a substrate that should lower the soil bulk density compared to NMC plots and increase the water infiltration rate. Bulk density showed no significant difference, but this may be due to measuring in the fall of 2022, 2 years after MC treatment.
Species interactions with sites for summer-related mortality were driven by the greater SA mortality on site 2. Overall, GB had the least amount of summer-related mortality (4.7%), with GA and WB at 5.7% and 7%, respectively, and SA mortality was the greatest, at 14.3%. The difference in species quickly became evident, as SA seedlings began to die off because they could not overcome the transplant shock on the harsh dry sites; SA are hydrophilic and are better adapted to mesic sites [28,29]. In addition, SA showed the greatest benefit from the S treatment and particularly on site 2, which had shallow soil and experienced the greatest stress. GA prefers soil types ranging from sand to gravel and can regularly be found on upland glacial till [27]. Although GB prefers moist, well-drained sites like other species, it can withstand harsh environments and tolerate arid gravelly soils, and has performed well in reclamation efforts on Pennsylvanian mine sites [22,25,26]. WB has been used in the past to stabilize soil on former mine sites [55,56].

4.2. Winter-Related Mortality

Despite an extensive review of the literature, we could not find species or site preparation effects on winter-related mortality. Interestingly, there were no species or species interactions for winter-related mortality. Though only 36 trees (1.9%) were affected, the treatments that experienced the greatest winter-related mortality occurred in plots with S alone or S treatment interactions. The increased mortality with the presence of S appeared to be largely due to a mold fungus that was observed on seedlings that died overwinter. There were significant but minor interactions with S, but it appeared that CWD with S reduced winter-related mortality and that the site × CWD interaction was due to increased mortality on the shallow, more-stressed site 2 without CWD. This was driving the three-way interaction found on the shallowest site 2, where S with NCWD had the greatest winter-related mortality, at 12.5%. It is possible that CWD limited winter-related mortality since its sheltering properties [41] limited the elements that may have cause nonfungal related mortality.

4.3. Frost Heave Mortality

Frost heave occurs when ice forms within the soil surface layers and within wet soil, causing a volumetric expansion and lifting the seedlings from their original planting positions [57]. The three-way site preparation (S × MC × CWD) interaction was driven by the NS NCWD combined treatment, which had the greatest frost heave mortality regardless of MC treatment. Overall, it was clear that there was a strong positive S treatment effect on frost heave since S and NS had 1.2% and 20.7% frost heave mortality, respectively. Frost heave can be mitigated by adding an insulating cover to the surface of the soil [58,59,60]. It has been reported that straw has an ability to insulate and maintain a higher temperature below ground in winter [59,60]. Insulation covers such as straw, forest litter, or wooden laths are effective in reducing frost heave mortality [58,61,62,63]. In 2019, [58] found that straw mulch was useful in controlling the freeze–thaw action by slowing water transport and heat transfer.
Site × S interaction showed that despite site differences in frost heave, S always had a positive effect on mortality. In fact, the more severe the site, the greater the benefit S had in preventing frost heave. It has been reported that frost heave is dependent on soil texture and depth to the water table [11,12]. These site factors along with the temperature and precipitation rates over the winter months severely effect the extent to which a seedling can be lifted from its original planting position [64]. An initial study by Beskow (1935) [10], the findings of which were again demonstrated by Kinosita (1979) [11], showed that the higher the ground water table, the greater the rate of frost heave, and the opposite occurs with a lower ground water table. A full saturation of soil leads to a greater rate of frost heave [65,66], and combined with a high ground water table, a maximum amount of frost heave likely occurred in some areas of the sites, especially sites 2 and 3, which had some of the shallowest soils in the experiment. Site 1 experienced the lowest frost heave mortality (6.8%), likely due to the average depth to bedrock of 31.93 cm ± 12.18 cm, whereas sites 2 and 3 had a 12.9% and 13.1% frost heave mortality, respectively.
In addition, water permeability is another factor that determines the frost heave susceptibility of a site. Permeability considers several factors, including soil texture. Sandy soils have a larger pore space and, therefore, higher permeability, but the moisture suction will be lower. This lower moisture suction means that water cannot easily move to the freezing front and thus minimizes frost heave. In contrast, clay and silt have a fine-grained soil texture resulting in higher moisture suction but also a lower water permeability, which can hinder water movement to the freezing front, mitigating frost heave [67].
CWD and its interactions had a positive effect on reducing frost heave mortality. Overall, CWD and NCWD had 2.5% and 19.4% frost heave mortality, respectively. The addition of CWD does insulate the ground by reducing direct environmental impacts; however, more importantly, it creates a heavier overburden layer that also lowers the rate of frost heave [68]. Over time, CWD will ameliorate the soil by trapping organic matter within the plots, increasing the insulation of the soil from frost penetration and the effects of frost heave [61,62]. In addition, with each passing year, the seedlings will have developed a more extensive root system that reduces the risk of frost heave mortality.
The MC treatment increased frost heave mortality from NMC at 8.5% to MC at 13.4%. In solely MC-treated plots, frost heave mortality was 47%, the greatest mortality of any treatment. The MC treatment created a slightly deeper soil profile than the NMC plots around them. The MC plots were easier to plant, providing seedling root systems a better growth substrate, but the new substrate had a higher susceptibility to heave, perhaps due to lower soil bulk density, and created intermediate soil pore sizes that may have promoted frost heave [67,69]. Studies on soil preparation for Norway spruce in tilled soils found that frost heave was more likely in disturbed or mixed soil [70].
The plant species effect was relatively small, accounting for 1.3% of the total variation, and there were no significant species interactions for frost heave mortality. Alders had slightly greater mortality than birches. A species that survives a period of frost heave has a much higher chance at surviving to adulthood [71,72]. The type of root system a plant puts out influences the survival of the species [73,74]. White birch and gray birch are both shallow rooting species depending on the soil depth [26,75], and GA and SA are also shallow rooting species [20,21]. Schramm 1958 looked at tree seedlings being heaved out of the soil and concluded that the top 2–3 inches had the largest effect, which was further confirmed by Heidmann in 1974 [74] with their use of “Ontario” plugs being heaved out when planted flush with the soil surface. The alders, particularly SA, suffered the greatest frost heave mortality, with 14.8%, and GA with 11.7% compared to birches at 8.6%.

4.4. First Full-Year Mortality

Here, we have documented and discussed the three causes of mortality on severely degraded 25-year barren landscapes. Overall, frost heave was the greatest factor in mortality during the first full year, and the frost heave mortality interactions and main effects were similarly found significant in the first full-year mortality. From a species perspective, SA suffered the most mortality from all three causes of mortality, probably because its hydrophilic requirements [29] were not met in the barren shallow soil landscape. The survival rates of the remaining three species were comparable. Growth will also be a determinant of how each species is affected by site preparation treatments [76]. As for site preparation factorial treatments, there was a clear demarcation between NS and S on overall first-year mortality, with S having very low mortality as well interactions with other site preparation treatments on mortality. It is clear the combination of benefits that S provides, such as moisture retention in the summer, related to preventing drought mortality [77], and insulation in the winter, related to preventing frost heave mortality [60], makes it the best single treatment, as S-treated plots had 12% mortality. The worst overall treatment, worse than control, for first full-year mortality was MC alone, at 60.9%, which, as discussed earlier, was driven by frost heave, accounting for most of this mortality (47%). Second-worst overall mortality was control (no site preparation treatments), with 37.5% overall mortality, again driven mostly by frost heave mortality. The third-worst was CWD alone, accounting for 25% mortality; this, however, was largely driven by summer-related mortality, but CWD is expected to drastically reduce mortality and enhance the fertility of the plots in the future [54]. In the literature, there is a lack of information pertaining to combinations of the three treatments; however, data from this experiment and [76] regarding growth over two years provides insight on how these combinations affect the species. From these mortality causes and the overall full-year mortality, there were additive effects or benefits to site preparation treatments, such as S CWD and MC in that order, but also some negative site preparation effects such as MC alone. Next on the ranking list is MCCWD, with 16.2% mortality. With the addition of CWD on MC plots, initially the first summer results saw a 4% increase in mortality, but after the first full year there was a 27% reduction in mortality in plots containing both MC and CWD compared to plots with just MC. The accumulation of organic matter in CWD over time should increase the moisture levels of the microsites and accelerate the decomposition of the CWD and other organic matter [51,52,53]. SMC had 12.5% mortality, still 0.5% higher than S alone, but considering the 60.9% mortality of MC alone, the apparent downsides of MC allowing for greater water infiltration [34] and creating a substrate more susceptible to frost heave were balanced out by straw’s ability to retain moisture and insulate against frost heave [58,70,77,78]. MC had an additional benefit from a logistical view in that it made planting easier. The three preparation treatments combined and SCWD had the lowest first full-year mortality rates, both at 3.1 and 4.2%, respectively, due to the additive effects of the treatments.

4.5. Application

Many site restoration planting efforts fail in barren degraded areas due to the lack of soil, soil moisture, and nutrients and the presence of frost heave. Barren sites are exposed to some of the harshest conditions, which include high temperatures, solar radiation, wind, extreme temperature changes and low soil moisture. Different levels of site degradation will require different single or combined site preparation treatments to test and help achieve successful plant survival. The application of specific site preparation treatments to other sites can be informed from the results of this experiment but should strongly consider local conditions and the native early-successional species in their area. Site preparation treatments such as S, CWD and MC allow these sites to become more habitable for survival. First full-year results showed that the implementation of all three site preparation treatments in combination reduced mortality to 3.1%. While the use of MC provides benefits such as improved aeration, water infiltration, root penetration, and ease of planting, the synergies with straw reduce surface evaporation, improve erosion control, and increase soil moisture retention during critical times of the growing season, thus substantially reducing mortality. In addition, straw clearly acts as a temperature insulator to prevent frost heave and resulting mortality. Straw plots in combination with CWD provide protection from severe conditions but may be necessary on some sites to prevent straw from being blown or washed away by weather events, Early-successional species such as WB, GB, GA, and SA need site preparation treatments to establish themselves and survive the first year on long-term barren lands. SA, a hydrophilic species, had greater mortality from drought and thus appears to need more moist sites. After one year, GB proved itself to be the best at surviving the harsh site conditions; nevertheless, GA and WB had comparable survival rates under these harsh conditions.

Author Contributions

J.E.M. designed and co-analyzed the experiment and co-authored the manuscript. D.G. managed the experiment, co-analyzed the data, and co-authored the manuscript. All authors have read and agreed to the published version of the manuscript.

Funding

This research received Canadian Department of National Defense (DND) Environmental Services Branch (ESB) funding and was supported by Canadian Department of Natural Resources Canada.

Data Availability Statement

Please contact corresponding author for data.

Acknowledgments

We gratefully acknowledge the useful edits and comments received from John Kershaw, Alex Mosseler and Jasen Golding. We are also grateful for the support from Noah Pond, Deanna McCullum, and Meagan Betts from DND ESB. In addition, we are grateful for the technical help in the establishment and management of the experiment provided by Axel Brisebois, Shawn Palmer, John Malcom, Will Bradley, Megan Hall, and Josh Kilburn.

Conflicts of Interest

The authors declare no conflicts of interest.

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Forests 15 00143 g001
Figure 1. (A) Rainfall (mm) sourced from nearby Fredericton, New Brunswick (45.9636° N, 66.6431° W) from 1 April 2021–20 September 2021. (B) Soil moisture readings in 2021 showing differences between straw and no straw coverage on dates where field assessments were done. Dates marked with * are significantly different p < 0.05. Note x-axis dates range differ.
Figure 1. (A) Rainfall (mm) sourced from nearby Fredericton, New Brunswick (45.9636° N, 66.6431° W) from 1 April 2021–20 September 2021. (B) Soil moisture readings in 2021 showing differences between straw and no straw coverage on dates where field assessments were done. Dates marked with * are significantly different p < 0.05. Note x-axis dates range differ.
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Figure 2. Summer related 2021 mortality (mean ± SE) with (A) MC × S × Site interaction, (B) S × Site × Species interaction. NS: no straw, S: straw, NMC: no meri-crushing, MC: Meri-crushing. n = 384.
Figure 2. Summer related 2021 mortality (mean ± SE) with (A) MC × S × Site interaction, (B) S × Site × Species interaction. NS: no straw, S: straw, NMC: no meri-crushing, MC: Meri-crushing. n = 384.
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Figure 3. Summer related 2021 mortality (mean ± SE) with MC × S × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris, NMC: no meri-crushing, MC: meri-crushing. n = 384.
Figure 3. Summer related 2021 mortality (mean ± SE) with MC × S × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris, NMC: no meri-crushing, MC: meri-crushing. n = 384.
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Figure 4. Winter related mortality (mean ± SE) with Straw × CWD × Site interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris. n = 384.
Figure 4. Winter related mortality (mean ± SE) with Straw × CWD × Site interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris. n = 384.
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Forests 15 00143 g005
Figure 5. Winter related mortality (mean ± SE) with (A) Site × CWD interaction, (B) Straw × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris. n = 384.
Figure 5. Winter related mortality (mean ± SE) with (A) Site × CWD interaction, (B) Straw × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris. n = 384.
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Forests 15 00143 g006
Figure 6. Frost heave mortality (mean ± SE) showing MC × S × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris, NMC: no meri-crushing, MC: meri-crushing. n = 384.
Figure 6. Frost heave mortality (mean ± SE) showing MC × S × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris, NMC: no meri-crushing, MC: meri-crushing. n = 384.
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Forests 15 00143 g007
Figure 7. Frost heave mortality (mean ± SE) showing Site × S interaction. Different letters in two-way interactions represent significant differences according to Tukey’s mean separation test, p = 0.05. NS: no straw, S: straw. n = 384.
Figure 7. Frost heave mortality (mean ± SE) showing Site × S interaction. Different letters in two-way interactions represent significant differences according to Tukey’s mean separation test, p = 0.05. NS: no straw, S: straw. n = 384.
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Forests 15 00143 g008
Figure 8. Total first year mortality (mean ± SE) showing MC × S × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris, NMC: no meri-crushing, MC: meri-crushing. n = 384.
Figure 8. Total first year mortality (mean ± SE) showing MC × S × CWD interaction. NS: no straw, S: straw, NCWD: no coarse woody debris, CWD: coarse woody debris, NMC: no meri-crushing, MC: meri-crushing. n = 384.
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Forests 15 00143 g009
Figure 9. Total first year mortality (mean ± SE) showing (A) Species × Straw interaction, (B) Site × S interaction. Different letters in two-way interactions represent significant differences according to Tukey’s mean separation test, p = 0.05. NS: no straw, S: straw. n = 384.
Figure 9. Total first year mortality (mean ± SE) showing (A) Species × Straw interaction, (B) Site × S interaction. Different letters in two-way interactions represent significant differences according to Tukey’s mean separation test, p = 0.05. NS: no straw, S: straw. n = 384.
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Forests 15 00143 g010
Figure 10. Total first year mortality (mean ± SE) classified by period of death. (A) Species broken down by when they died. (B) Treatment types and their associated mortality. CRTL: no treatments, MC: meri-crushing, CWD: coarse woody debris, MCCWD: meri-crushing + coarse woody debris, S: straw, MCS: meri-crushing + straw, SCWD: straw + coarse woody debris, MCSCWD: meri-crushing + straw + coarse woody debris. n = 384.
Figure 10. Total first year mortality (mean ± SE) classified by period of death. (A) Species broken down by when they died. (B) Treatment types and their associated mortality. CRTL: no treatments, MC: meri-crushing, CWD: coarse woody debris, MCCWD: meri-crushing + coarse woody debris, S: straw, MCS: meri-crushing + straw, SCWD: straw + coarse woody debris, MCSCWD: meri-crushing + straw + coarse woody debris. n = 384.
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Table 1. Species and provenance of seedling material used in the experiment.
Table 1. Species and provenance of seedling material used in the experiment.
SpeciesSource Year Provenance Country Latitude (N)Longitude (W)Elevation (m)
Betula papyriferaWhite birch1998Wayeton, NBCAN47.21667−65.93333300
Betula papyriferaWhite birch1998Jewetts Creek, NBCAN45.83333−66.9833350
Betula populifoliaGray birch2008Newmarket, NBCAN45.80501−66.95634149
Betula populifoliaGray birch2008Newmarket, NBCAN45.83147−66.97115130
Betula populifoliaGray birch2008Newmarket, NBCAN45.8076−66.96825141
Betula populifoliaGray birch2008Newmarket, NBCAN45.83486−66.96272125
Betula populifoliaGray birch1999Bai-du-vin, NBCAN47.03333−65.166665
Alnus Viridis ssp. crispaGreen alder2002West Quaco, NBCAN45.33−65.5365
Alnus Viridis ssp. crispaGreen alder1999Lower Prince William, NBCAN45.87−6720
Alnus incana ssp. rugosaSpeckled alder1983Enmore, PEICAN46.58−64.0510
Alnus incana ssp. rugosaSpeckled alder1983Vallyfield, PEICAN46.13−62.7245
Alnus incana ssp. rugosaSpeckled alder1983Shediac, NBCAN46.23−64.615
Table 2. 2020 soil properties for three experimental sites on DND Base Gagetown.
Table 2. 2020 soil properties for three experimental sites on DND Base Gagetown.
IDOrganic Matter (%)Total Nitrogen (%)Carbon (%)C:N RatiopHPhopshorus (ppm)Pottasium (meq/100 g)
Site 11.744 ± 0.99 a0.219 ± 0.05 a1.013 ± 0.58 a4.350 ± 1.59 ab4.43 ± 0.16 b6.375 ± 3.02 a0.110 ± 0.03 a
Site 20.871 ± 0.37 a0.171 ± 0.03 b0.509 ± 0.21 a2.938 ± 0.93 b4.63 ± 0.15 a4.375 ± 1.19 a0.113 ± 0.03 a
Site 31.303 ± 0.54 a0.168 ± 0.03 b0.758 ± 0.31 a4.538 ± 0.72 a4.61 ± 0.08 a5.875 ± 2.10 a0.104 ± 0.02 a
IDCalcium (meq/100 g)Magnesium (ppm)Clay (%)Silt (%)Sand (%)Rocks (%) *Average Depth (cm)
Site 10.381 ± 0.15 b0.156 ± 0.08 a15.81 ± 3.41 a43.76 ± 7.45 a40.44 ± 10.69 b36.05 ± 17.36 a31.93 ± 12.18 a
Site 21.044 ± 0.32 a0.280 ± 0.09 a13.51 ± 3.01 a34.25 ± 6.20 b52.20 ± 9.04 b41.21 ± 12.46 a11.64 ± 7.27 b
Site 31.198 ± 0.72 a0.369 ± 0.24 a12.93 ± 2.69 a28.84 ± 5.13 b58.21 ± 7.66 a43.96 ± 15.46 a14.94 ± 9.78 b
Note: Sites with different letters are significantly different as per Tukey’s mean separation test (p < 0.05). * Rocks (%) was taken before the sane, silt, clay (%), which are equal to 100%.
Table 3. Site preparation treatments, species and site effects and interactions on summer-related 2021 mortality ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Table 3. Site preparation treatments, species and site effects and interactions on summer-related 2021 mortality ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Summer Related Mortality *
Source of VariationdfMSVC (%)p Value
MC10.2860.90.021
S13.0169.2<0.001
CWD10.0010.00.908
SITE20.5883.6<0.001
SPECIES30.4894.5<0.001
BLOCK(SITE)90.1654.50.001
S*MC10.1610.50.084
CWD*MC10.4461.40.004
SITE*MC20.3302.00.002
SPECIES*MC30.0010.00.997
CWD*S10.0460.10.356
SPECIES*S30.2982.70.001
SITE*S21.0766.6<0.001
SPECIES*CWD30.0500.50.421
SITE*CWD20.0820.50.219
SITE*SPECIES60.0841.50.154
CWD*S*MC10.2310.70.038
SPECIES*S*MC30.0310.30.626
SITE*S*MC20.3202.00.003
SPECIES*CWD*MC30.0920.80.162
SITE*CWD*MC20.0770.50.237
SITE*SPECIES*MC60.0340.60.701
SPECIES*CWD*S30.0090.10.923
SITE*CWD*S20.0720.40.261
SITE*SPECIES*S60.2444.5<0.001
SITE*SPECIES*CWD60.0751.40.209
Error3080.05350.2
R2 0.498
* arcsine square root transformed.
Table 4. Site preparation treatments, species and site effects and interactions on winter-related mortality. ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Table 4. Site preparation treatments, species and site effects and interactions on winter-related mortality. ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Winter Related Mortality *
Source of VariationdfMSVC (%)p Value
MC10.1031.00.043
S10.2582.40.001
CWD10.1211.10.029
SITE20.0541.00.118
SPECIES30.0601.70.066
BLOCK(SITE)90.0201.70.632
S*MC10.0060.10.612
CWD*MC10.0060.10.612
SITE*MC20.0020.00.918
SPECIES*MC30.0581.60.076
CWD*S10.3463.3<0.001
SPECIES*S30.0571.60.077
SITE*S20.0260.50.348
SPECIES*CWD30.0150.40.602
SITE*CWD20.0991.90.020
SITE*SPECIES60.0160.90.691
CWD*S*MC10.0260.20.311
SPECIES*S*MC30.0341.00.259
SITE*S*MC20.0090.20.709
SPECIES*CWD*MC30.0150.40.627
SITE*CWD*MC20.0000.01.000
SITE*SPECIES*MC60.0231.30.470
SPECIES*CWD*S30.0441.30.152
SITE*CWD*S20.1382.60.004
SITE*SPECIES*S60.0120.70.811
SITE*SPECIES*CWD60.0090.50.902
Error3080.02572.5
R2 0.275
* arcsine square root transformed.
Table 5. Site preparation treatments, species and site effects and interactions on frost heaved mortality. ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Table 5. Site preparation treatments, species and site effects and interactions on frost heaved mortality. ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Frost Heave Mortality *
Source of VariationdfMSVC(%)p Value
MC10.4830.90.010
S110.11019.0<0.001
CWD16.85712.9<0.001
SITE20.4511.70.002
SPECIES30.2261.30.026
BLOCK(SITE)90.1813.10.009
S*MC10.7771.50.001
CWD*MC10.3460.60.030
SITE*MC20.1550.60.119
SPECIES*MC30.0630.40.455
CWD*S14.9189.2<0.001
SPECIES*S30.1100.60.211
SITE*S20.2270.90.045
SPECIES*CWD30.1891.10.052
SITE*CWD20.0720.30.372
SITE*SPECIES60.0600.70.546
CWD*S*MC10.6001.10.004
SPECIES*S*MC30.0150.10.887
SITE*S*MC20.0170.10.795
SPECIES*CWD*MC30.0200.10.838
SITE*CWD*MC20.0080.00.899
SITE*SPECIES*MC60.0750.80.406
SPECIES*CWD*S30.1010.60.244
SITE*CWD*S20.0240.10.721
SITE*SPECIES*S60.0320.40.850
SITE*SPECIES*CWD60.0240.30.921
Error3080.07241.9
R2 0.581
* arcsine square root transformed.
Table 6. Site preparation treatments, species and site effects and interactions on total first-year mortality. ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Table 6. Site preparation treatments, species and site effects and interactions on total first-year mortality. ANOVA table including degrees of freedom (df), mean square values (MS), variance components (VC), and p values. p values < 0.05 are in bold print. Meri-crushing (MC), Straw (S), Coarse woody debris (CWD).
Total First Year Mortality *
Source of VariationdfMSVC(%)p Value
MC10.1930.20.202
S117.20419.1<0.001
CWD18.2219.1<0.001
SITE22.3175.1<0.001
SPECIES31.6275.4<0.001
BLOCK(SITE)90.1851.80.127
S*MC10.1940.20.201
CWD*MC11.5401.7<0.001
SITE*MC20.5951.30.007
SPECIES*MC30.0090.00.974
CWD*S11.8892.1<0.001
SPECIES*S30.8562.9<0.001
SITE*S21.6973.8<0.001
SPECIES*CWD30.2630.90.086
SITE*CWD20.0560.10.625
SITE*SPECIES60.1260.80.384
CWD*S*MC11.2841.40.001
SPECIES*S*MC30.0350.10.829
SITE*S*MC20.1310.30.332
SPECIES*CWD*MC30.0720.20.610
SITE*CWD*MC20.1320.30.329
SITE*SPECIES*MC60.0400.30.916
SPECIES*CWD*S30.1210.40.383
SITE*CWD*S20.0720.20.545
SITE*SPECIES*S60.1631.10.223
SITE*SPECIES*CWD60.0940.60.577
Error3070.11840.4
R2 0.597
* arcsine square root transformed.
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Galea, D.; Major, J.E. First-Year Mortality of Four Early-Successional Species on Severely Degraded Sites in Eastern Canada as Influenced by a Factorial of Site Preparation Treatments. Forests 2024, 15, 143. https://doi.org/10.3390/f15010143

AMA Style

Galea D, Major JE. First-Year Mortality of Four Early-Successional Species on Severely Degraded Sites in Eastern Canada as Influenced by a Factorial of Site Preparation Treatments. Forests. 2024; 15(1):143. https://doi.org/10.3390/f15010143

Chicago/Turabian Style

Galea, Dominic, and John E. Major. 2024. "First-Year Mortality of Four Early-Successional Species on Severely Degraded Sites in Eastern Canada as Influenced by a Factorial of Site Preparation Treatments" Forests 15, no. 1: 143. https://doi.org/10.3390/f15010143

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