In total, we collected 8532 insects across three years of sampling, comprising 7 orders and 71 families. Of this total number, 3762 (44.1%) were in Dolichopodidae, 2202 (25.8%) were in Sarcophagidae, and 555 (6.5%) were in Halictidae. We collected 34 unique taxa in 2018, 55 in 2019, and 41 in 2020. A sampling abundance accumulation curve of family richness indicates that species richness in 2019 and 2020 was similar. The overall family diversity in 2018 does not approach an asymptote, suggesting that sampling may have inadequately captured family diversity during that year (Supp. Figure 1). Abundance of insects per sampling date varied widely, with a range of 1-289 individuals captured on a given sampling date pooled across all sites (Supp. Figure 2.).
Family abundance
Of the 18 insect families included in our analyses, nine exhibited a significant interaction effect between year and Julian day on abundance per sampling event (model results summarized in Table 1). This included Aphididae (F2,92 = 3.35, p = 0.04, Fig. 1a), Apidae (F2,92 = 8.86, p < 0.001, Fig. 1b), Dolichopodidae (F2,91 = 5.80, p < 0.01, Fig. 1c), Formicidae (F2,93 = 7.60, p < 0.001, Fig. 1d), Hesperiidae (F2,93 = 4.46, p = 0.01, Fig. 1e), Miridae (F2,93 = 4.53, p = 0.01, Fig. 1f), Scoliidae (F2,91 = 3.45, p < 0.05, Fig. 1g), Sphecidae (F2,91 = 3.45, p = 0.04, Fig. 1h), and Vespidae (F2,92 = 4.73, p = 0.01, Fig. 1i). In general, there were more apids, scoliids, sphecids, and vespids in late 2020 compared to late 2018 and late 2019 (Figs. 1b, 1g, 1h, 1i, respectively). In both late 2019 and late 2020, there were more dolichopodids than in late 2018 (Fig. 1c). Several groups showed an apparent drop in abundance between equivalent periods in subsequent years, with more aphids and formicids in late 2019 compared to late 2020 (Figs. 1a and 1d, respectively), as well as more hesperiids in late 2018 and late 2019 compared to late 2020 (Fig. 1e).
We observed a significant year effect for five families; Calliphoridae (F2,94 = 4.87, p = 0.01, Fig. 1j), Cicadellidae (F2,94 = 6.77, p < 0.01, Fig. 1k), Crabronidae (F2,93 = 4.95, p = 0.01, Fig. 1l), Halictidae (F2,92 = 10.16, p < 0.001, Fig. 1m), Muscidae (F2,94 = 7.02, p < 0.01, Fig. 1n) and Sarcophagidae (F2,92 = 10.62, p < 0.001, Fig. 1o). There were significantly more calliphorids (t95.7 = 2.86, p = 0.04), cicadellids (t95.7 = 3.64, p < 0.01), halictids (t92.7 = 4.38, p < 0.001), and muscids (t95.7 = 2.87, p = 0.01) in 2019 than in 2020. There were significantly more crabronids (t91.2 = 2.78, p = 0.01) and sarcophagids (t91.1 = 2.33, p < 0.001) in 2020 than 2018. Finally, there were significantly more muscids (t93.8 = 3.11, p = 0.01) and sarcophagids (t92.0 = 4.31, p < 0.001) 2019 compared to 2018.
There was a slight but significant effect of Julian day on abundance per sampling bout for Muscidae (F1,93 = 11.47, p < 0.001) and Syrphidae (F1,93 = 40.55, p < 0.001, Fig. 1p), with a small increase in abundance over a given year for both families (0.00004 and 0.0003 individuals for each successive Julian day, respectively).
There was no significant effect of either year, Julian day, or their interaction on the abundance of Chloropidae or Ichneumonidae (0.05 < p < 0.81, Fig. 1q and 1r, respectively). However, there was a marginally non-significant interaction effect between year and Julian day on ichneumonid abundance, such that ichneumonids were more abundant in late 2019 and late 2020 compared to late 2018 (F2,91 = 3.08, p = 0.05, Fig. 1r).
Table 1
Results of linear mixed-effects models examining abundance of each family per sampling event (ANOVA (Type II or III Wald F-tests with Kenward-Roger degrees of freedom).
| Year | Julian day | Year*Julian day |
Family | F statistic | df | df residual | p-value | F statistic | df | df residual | p-value | F statistic | df | df residual | p-value |
Aphididae | 1.98 | 2 | 91.54 | 0.14 | 0.09 | 1 | 92.39 | 0.77 | 3.35 | 2 | 91.54 | 0.04* |
Apidae | 6.84 | 2 | 91.50 | < 0.01* | 1.41 | 1 | 92.31 | 0.24 | 8.86 | 2 | 91.50 | < 0.001* |
Calliphoridae | 4.87 | 2 | 94.46 | < 0.01* | 0.68 | 1 | 93.41 | 0.41 | 1.49 | 2 | 93.47 | 0.23 |
Chloropidae | 1.49 | 2 | 92.12 | 0.23 | 0.10 | 1 | 93.83 | 0.76 | 2.45 | 2 | 92.15 | 0.09 |
Cicadellidae | 6.77 | 2 | 94.29 | < 0.01* | 0.09 | 1 | 92.99 | 0.76 | 1.13 | 2 | 93.11 | 0.33 |
Crabronidae | 4.95 | 2 | 92.90 | 0.01* | 1.19 | 1 | 91.56 | 0.28 | 0.97 | 2 | 91.68 | 0.38 |
Dolichopodidae | 2.90 | 2 | 91.30 | 0.06 | 2.06 | 1 | 91.78 | 0.15 | 5.80 | 2 | 91.30 | < 0.01* |
Formicidae | 6.36 | 2 | 92.54 | < 0.01* | 6.34 | 1 | 94.73 | 0.01* | 7.60 | 2 | 92.61 | < 0.001* |
Halictidae | 10.16 | 2 | 91.95 | < 0.001* | 3.25 | 1 | 91.22 | 0.07 | 0.55 | 2 | 91.28 | 0.58 |
Hesperiidae | 13.03 | 2 | 94.25 | < 0.001* | 17.27 | 1 | 92.90 | < 0.001* | 4.46 | 2 | 93.03 | 0.01* |
Ichneumonidae | 1.91 | 2 | 91.33 | 0.15 | 0.06 | 1 | 91.86 | 0.81 | 3.08 | 2 | 91.33 | 0.05 |
Miridae | 4.24 | 2 | 92.68 | 0.02* | 4.07 | 1 | 94.99 | < 0.05* | 4.53 | 2 | 92.75 | 0.01* |
Muscidae | 7.02 | 2 | 94.37 | < 0.01* | 11.47 | 1 | 93.17 | < 0.01* | 0.26 | 2 | 93.26 | 0.78 |
Sarcophagidae | 10.62 | 2 | 92.39 | < 0.001* | 1.76 | 1 | 91.36 | 0.19 | 0.02 | 2 | 91.44 | 0.98 |
Scoliidae | 12.44 | 2 | 92.71 | < 0.001* | 2.44 | 1 | 91.48 | 0.12 | 3.10 | 2 | 91.58 | < 0.05* |
Sphecidae | 2.28 | 2 | 91.48 | 0.11 | 0.24 | 1 | 92.24 | 0.62 | 3.45 | 2 | 91.48 | 0.04* |
Syrphidae | 1.19 | 2 | 94.07 | 0.31 | 40.55 | 1 | 92.57 | < 0.001* | 2.22 | 2 | 92.72 | 0.11 |
Vespidae | 2.23 | 2 | 92.37 | 0.11 | 0.00 | 1 | 94.37 | 0.97 | 4.73 | 2 | 92.42 | 0.01* |
Significance levels: *p < 0.05, **p < 0.01, ***p < 0.001 |
Guild abundance
Model results for guilds are summarized in Table 2. There was no significant effect of either year (F2,91 = 2.53, p = 0.09), Julian day (F1,91 = 4.44, p = 0.04), or their interaction (F1,91 = 2.34, p = 0.10, Fig. 2a) on overall bee abundance.
There was a significant effect of year on butterfly (F2,94 = 13.01, p < 0.001, Fig. 2b) abundance. There were significantly more butterflies in 2019 than both 2018 (t93.3 = 2.53, p = 0.03) and 2020 (t95.4 = 5.28, p < 0.001). In addition to the main effect of year, there was also a significant effect of Julian day on abundance per bout in butterflies. Butterflies demonstrated a slight but significant positive correlation between abundance per bout and Julian day (F1,92 = 19.99, p < 0.001), with an average increase of 0.0002 individuals for each subsequent Julian day.
Predatory wasps exhibited a significant interaction effect between year and Julian day on abundance per sampling event (F2,91 = 5.70, p = 0.01, Fig. 2c), with a general higher abundance of wasps in late 2019 and 2020 compared to late 2018. There was also a significant interaction day between year and Julian day on predatory fly abundance (F2,91 = 3.97, p = 0.02, Fig. 2d), again with a general higher abundance of predatory wasps in late 2019 and 2020 compared to late 2018.
Table 2
Results of linear mixed-effects models examining abundance of each guild per sampling event (ANOVA (Type II or III Wald F-tests with Kenward-Roger degrees of freedom). Starred values indicate significance (α = 0.05).
| Year | Julian day | Year*Julian day |
Family | F statistic | df | df residuals | p-value | F statistic | df | df residuals | p-value | F statistic | df | df residuals | p-value |
bees | 2.53 | 2 | 91.36 | 0.09 | 4.44 | 1 | 91.93 | 0.04* | 2.34 | 2 | 91.35 | 0.10 |
butterflies | 13.01 | 2 | 93.79 | < 0.001* | 19.99 | 1 | 92.19 | < 0.001* | 3.06 | 2 | 92.35 | 0.05 |
predatory flies | 1.74 | 2 | 91.30 | 0.18 | 0.22 | 1 | 91.78 | 0.64 | 3.97 | 2 | 91.29 | 0.02 |
predatory wasps | 2.80 | 2 | 91.25 | 0.07 | 0.94 | 1 | 91.66 | 0.33 | 5.25 | 2 | 91.25 | 0.01* |
Significance levels: *p < 0.05, **p < 0.01, ***p < 0.001 |