Comparison of vegetation between S. altissima-dominant areas and uninvaded areas
In the area dominated by S. altissima, we detected 44 and 37 plant species in 2018 and 2019, respectively (Table S3). Plants belonging to Asteraceae were most frequent, followed by Gramineae and Rosaceae. In the area dominated by native species, we found 53 plant species in 2018 (Table S4). Asteraceae was most frequent, followed by Gramineae and Fabaceae. All species detected in the areas dominated by S. altissima or native species in 2018 and 2019 are listed in Table S2.
The average relative coverage of S. altissima was 83.2% in S. altissima-dominant sites, compared with 5.6% at sites dominated by native species (Fig. 1a). The average number of total species per quadrat differed significantly between S. altissima-dominated and or native species-dominated areas (i.e., 4.8 and 7.8, respectively) (Fig. 1b). The average number of native species also differed significantly between S. altissima-dominated and or native species-dominated areas (i.e., 3.1 and 4.9, respectively).
In the area dominated by S. altissima, the importance value was highest for S. altissima (0.409), followed by Pueraria lobata (0.141) and Artemisia princeps (0.087) in July 2018 (Table S3). In the area dominated by native species, Glycine soja (0.166), P. lobata (0.158), and Erigeron canadensis (0.154) had the high importance values. The importance value for S. altissima was 0.104, ranking 6th in July 2018 (Table S4).
Management of S. altissima-dominated community by repeated mowing and eco-friendly herbicide application
The coverages of S. altissima did not differ significantly between 2018 and 2019, regardless of the treatment applied to quadrats (Fig. S2). The quadrats that were mowed three times in 2018 had a lower S. altissima coverage in 2019 than in 2018; however, the variance was high and the difference was not statistically significant. Therefore, mowing treatment for 1 year did not affect the coverage of S. altissima, regardless of mowing frequency.
The importance value for S. altissima changed significantly after mowing treatment for 1 year. The importance value for S. altissima was 0.409 in 2018 and decreased to 0.321 in 2019. Consistent results were obtained at all sites subjected to mowing (Table S5).
In July 2018, 2 months after the first treatment (mowing or Eco200), the height of S. altissima in the mowing group was significantly different from that of the control group, with no difference in the height of S. altissima between the Eco 200 treatment and control group (Table S6). After 1 year of mowing or Eco 200 treatment, in April 2019, the height of the dead shoots differed significantly between the control group and the Mowing 1, Mowing 2, and Mowing 3 groups (Table 1). In particular, the average heights in the Mowing 2 and Mowing 3 groups were one-tenth that of the control group (Table 1). However, the height of the dead shoots in the Eco 200 group was not significantly different from that in the control group, and was larger than those in the mowing treatment groups.
Table 1
Heights of dead shoots of S. altissima after 1 year of treatment in April 2019
Site | Control | Mowing 1 | Mowing 2 | Mowing 3 | Eco 200 |
Site 1 | 162.8 ± 6.62a | 85.7 ± 5.52b | 29.5 ± 2.74c | 21.3 ± 2.09c | 77.9 ± 8.94b |
Site 2 | 213.2 ± 12.08a | 87.1 ± 7.33c | 38.4 ± 2.17d | 29.0 ± 2.01d | 144.8 ± 9.57b |
Site 3 | 163.4 ± 7.31a | 66.3 ± 4.55c | 40.7 ± 2.57d | 33.4 ± 2.11d | 143.4 ± 7.13b |
Site 4 | 191.6 ± 3.84a | 160.8 ± 7.90b | 59.2 ± 1.66d | 81.9 ± 6.50c | 166.9 ± 12.33b |
Site 5 | 181.7 ± 4.76a | 114.7 ± 2.90b | 89.5 ± 10.23c | 46.6 ± 5.12d | 183.8 ± 9.00a |
Site 6 | 174.8 ± 4.34a | 82.9 ± 4.49c | 39.7 ± 2.51d | 31.6 ± 2.47d | 153.3 ± 3.12b |
Site 7 | 165.5 ± 4.79a | 91.2 ± 6.02b | 64.7 ± 3.24c | 44.0 ± 3.67d | 153.8 ± 7.25a |
Mowing 1: mowed once in July; Mowing 2: mowed twice in May and September; Mowing 3: mowed three times in May, July, and September; Eco 200: 30% of shoots in the quadrats were cut near the ground, and the cut surfaces were covered with the Eco 200 block. Data are presented as means ± standard errors of 20 replicates. Means within a column followed by different letters are significantly different at p < 0.05 (ANOVA with post hoc Tukey’s test). Units: cm. |
We continued the experiment to determine the effects of mowing on the growth of S. altissima in 2019. We measured the height of shoots of S. altissima in May, July, and September, before each mowing treatment. The height of the shoot already differed between the control and Mowing 2 or Mowing 3 groups in May 2019, before mowing (Table 2). Height did not differ between the Mowing 1 and control groups. In July, heights in the Mowing 2 and Mowing 3 groups were significantly different from that in the control group, with no difference between the Mowing 1 and control groups (Table S7). In September, heights in all treatment groups were significantly different from that in the control group (Table 3).
Table 2
Heights of S. altissima before mowing treatment in May 2019
Site | Control | Mowing 1 | Mowing 2 | Mowing 3 |
Site 1 | 101.0 ± 3.2a | 101.0 ± 3.0a | 81.1 ± 2.6b | 64.9 ± 3.1c |
Site 2 | 116.5 ± 4.7a | 111.8 ± 4.7a | 82.9 ± 4.1b | 72.5 ± 6.6b |
Site 4 | 89.1 ± 2.2a | 74.8 ± 2.2b | 49.5 ± 3.1d | 59.8 ± 1.6c |
Site 5 | 93.5 ± 2.5a | 99.5 ± 2.3a | 78.0 ± 2.0b | 77.1 ± 2.0b |
Site 6 | 125.3 ± 5.2a | 117.0 ± 3.0a | 86.5 ± 3.0c | 105.0 ± 3.4b |
Site 7 | 123.3 ± 3.3a | 119.3 ± 2.9ab | 111.3 ± 2.8bc | 110.3 ± 3.1c |
Site 8 | 96.0 ± 4.0a | 94.5 ± 4.1ab | 84.5 ± 3.3bc | 81.0 ± 3.1c |
Mowing 1: mowed once in July; Mowing 2: mowed twice in May and September; Mowing 3: mowed three times in May, July, and September. Data are presented as means ± standard errors of 10 replicates. Means within a column followed by different letters are significantly different at p < 0.05 (ANOVA with post hoc Tukey’s test). Units: cm. |
Table 3
Heights of S. altissima before mowing in September 2019
Site | Control | Mowing 1 | Mowing 2 | Mowing 3 |
Site 1 | 168.0 ± 3.27a | 64.0 ± 3.32c | 100.5 ± 9.14b | 67.5 ± 3.1c |
Site 2 | 192.0 ± 11.3a | 79.0 ± 4.07c | 111.0 ± 6.9b | 73.0 ± 7.61c |
Site 3 | 159.0 ± 4.99a | 123.0 ± 7.12b | 84.5 ± 5.89c | 47.0 ± 2.71d |
Site 4 | 190.0 ± 2.98a | 102.0 ± 3.89b | 112.5 ± 8.07b | 62.0 ± 5.33c |
Site 5 | 217.0 ± 10.23a | 87.5 ± 4.9c | 137.0 ± 4.96b | 62.5 ± 3.1d |
Site 6 | 177.5 ± 6.76a | 97.5 ± 5.44c | 120.5 ± 4.97b | 60.0 ± 2.58d |
Site 7 | 143.5 ± 12.2a | 81.0 ± 2.77b | 138.5 ± 3.5a | 62.0 ± 4.67b |
Mowing 1: mowed once in July; Mowing 2: mowed twice in May and September; Mowing 3: mowed three times in May, July, and September. Data are presented as means ± standard errors of 10 replicates. Means within a column followed by different letters are significantly different at p < 0.05 (ANOVA with post hoc Tukey’s test). Units: cm. |
The number of shoots of S. altissima did not differ significantly between groups in March 2018 before the experiment started (Fig. 2). After 1 year of mowing or Eco 200 treatment, the number of shoots of S. altissima was significantly lower in the Mowing 3 group than in the control group in April 2019 (Fig. 2). Mowing 1, Mowing 2, and Eco 200 treatments did not reduce the number of shoots compared with the number observed in the control group.
The number of leaves was lower in mowing treatment groups than in the control group (Table S8). Mowing 3 (mowed twice before harvesting) had the fewest leaves between the control and treatment groups. The aboveground biomass of S. altissima was also significantly lower in the mowing treatment groups than in the control group. However, there were no significant differences between mowing treatments (Fig. 3).
The biomass of rhizomes was significantly affected by mowing treatment and by the frequency of mowing after 2 years (Fig. 4). Plants subjected to mowing once a year had 933.6 g/m2 rhizomes, similar to the rhizome content of control plants (987.7 g/m2). However, plants that were mowed two or three times had 732.0 g/m2 or 535.3 g/m2 rhizomes, respectively, which were significantly lower than the estimate for control plants.
Effects of mowing frequency on reproductive characteristics
We found that flowering rates of S. altissima were affected by mowing. All individuals in the Mowing 1 group had flowers, significantly higher than the flowering rate of the control group (93 ± 7%). However, the flowering rates in the Mowing 2 (0%) and Mowing 3 groups (20%) were significantly lower than that in the control group in Fall 2019 (Fig. 5).
Effects of mowing frequency and eco-friendly herbicide application on the early establishment of S. altissima invasion
Although there were no statistically significant differences between the control and treatment groups due to high variances, patches of S. altissima showed the greatest expansion in the control group, followed by the 20% Eco 200 and 100% Eco 200 groups (Table 4). In particular, 100% mowing and Eco 200 treatment decreased the patch area of S. altissima.
Table 4
Change in areas occupied by S. altissima in patch-type communities from March 2018 to March 2019
Control | Mowing 3 | Eco 20% | Eco 100% |
205.2 ± 107.3 | 24.7 ± 44.0 | 104.5 ± 85.2 | -31.7 ± 13.0 |
Mowing 3: mowed three times in May, July, and September; Eco 20%: 20% of shoots were mowed and treated with Eco 200; Eco 100%: 100% of shoots were mowed and treated with Eco 200. Data are presented as means ± standard errors of six replicates. |
The flowering rate decreased with the frequency of mowing in the patches (sites 8–10) of S. altissima (Fig. S3). All individuals flowered in the control quadrats. However, the flowering rate in the Mowing 3 group was 75%, which was significantly lower than that in the control group. The Eco 20% and Eco 100% patches had no flowering individuals (Fig. S4).
Allelopathic effect of S. altissima
S. altissima extract adversely affected the germination rates of Lactuca sativa L. and Oenothera biennis L. (Table S9). For L. sativa, the germination rate was 32.9% at 3 days and increased to 44.3% at 6 days in the control group. For O. biennis, the germination rate was 45.7% at 6 days and increased to 65.7% at 12 days in the control group. However, when treated with the extract of S. altissima, the germination of seeds of both plants was delayed or did not occur. L. sativa seeds did not germinate until 9 days after treatment with a 20% dilution of the S. altissima extract and did not germinate at all when treated with a 50% dilution of the S. altissima extract. The seeds of O. biennis showed similar responses to S. altissima extract, although small numbers of seeds germinated, even when treated with 50% S. altissima extract (Table S9). The growth of roots of both plants was also significantly reduced by treatment with the S. altissima extract (Table S10). L. sativa seedlings showed root elongation of only 0.8 mm 12 days after treatment with a 20% dilution of S. altissima extract. The roots of O. biennis seedlings did not grow when treated with S. altissima extract, regardless of the concentration.