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Article

Native Grass Enhances Bird, Dragonfly, Butterfly and Plant Biodiversity Relative to Conventional Crops in Midwest, USA

by
Steven I. Apfelbaum
1,*,
Susan M. Lehnhardt
2,
Michael Boston
2,
Lea Daly
2,
Gavin Pinnow
2,
Kris Gillespie
2 and
Donald M. Waller
3
1
Applied Ecological Institute, Inc., N673 Mill Rd., Juda, WI 53550, USA
2
Lower Sugar Watershed Association, N3941 Golf Course Rd., Brodhead, WI 53520, USA
3
Department of Botany, University of Wisconsin-Madison, Madison, WI 53706, USA
*
Author to whom correspondence should be addressed.
Agriculture 2025, 15(15), 1666; https://doi.org/10.3390/agriculture15151666
Submission received: 11 June 2025 / Revised: 28 July 2025 / Accepted: 31 July 2025 / Published: 1 August 2025
(This article belongs to the Section Agricultural Systems and Management)
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Abstract

Conspicuous declines in native grassland habitats have triggered sharp reductions in grassland birds, dragonflies, butterflies, and native plant populations and diversity. We compared these biotic groups among three crop type treatments: corn, alfalfa, and a perennial native grass, Virginia wild rye, (Elymus virginicus L.) or VWR. This crop type had 2-3X higher bird, dragonfly, butterfly and plant species richness, diversity, and faunal abundance relative to alfalfa and corn types. VWR crop fields also support more obligate grassland bird species and higher populations of dragonfly and butterfly species associated with grasslands and wet meadows. In contrast, the corn and alfalfa types support few or no obligatory grassland birds and mostly non-native insects such as the white cabbage looper (Artogeia rapae L.), the common yellow sulfur butterfly (Colias philodice Godart.), and the mobile and migratory common green darner dragonfly (Anax junius Drury.). In sum, the VWR perennial native grass crop type offers a special opportunity to improve the diversity and abundance of grassland bird species, beneficial insect species, and many native plant species within agricultural landscapes.

1. Introduction

Perennial native grass species dominated grasslands and savanna’s that historically occupied 5–6 billion hectares of the Earth’s surface [1]. In addition to sustaining biodiversity and wildlife habitats, native grasslands support many ecosystem services. These include enhancing soil fertility; building soil structure; fixing and storing large quantities of organic carbon that, in turn, helps moderate the Earth’s climate; and slowing runoff, erosion, and nutrient losses that sustain the integrity of these habitats [2,3,4,5]. The ability of historic native grasslands to sustain these services has declined in proportion to their conversion to farms to support food production.
Heavy grazing and intensive grassland haying have also reduced ecosystem services and biodiversity in ways that threaten many cultural, economic, and biological values [6]. In contrast, using perennial native grass crops in grazed or hayed rangelands to produce meat can work to maintain grassland health [7,8,9,10]. Native perennial grass crops like VWR can be planted once and then harvested repeatedly for a decade before yields decline [11]. This raises the possibility that using native long-lived, deep-rooted grassland species, adapted to stressful environments, might help agricultural lands resist degradation. Such efforts gain importance given that climate change has increased the frequency and intensity of droughts, windstorms, and flooding. Increasing plantings of native perennial crops might therefore better maintain the economic productivity of agricultural land at risk from global climate change.
VWR is a grass native to North America [12] that occurs commonly in floodplains, forests, savannas, and wetland margins. It grows from the East Coast of the United States (US) through the Midwest corn belt and along river systems west to the Rocky Mountains. Across the northern tier of states, VWR grows in moist, seasonally wet margins of prairie potholes, high margins of floodplains along rivers and around lakes, under semi-open to open riparian gallery forests, and in oak savannas. VWR was once widespread, occupying the deep fertile soils that became the corn belt. Its occurrence today is restricted to floodplains and forests where sufficient light reaches the understory. Because VWR is a C3 grass adapted to cooler climates and seasons, it rarely grows in southern states like Florida, Texas, or Arizona. It was often used as a food crop by Indigenous Americans [13] and shows promise as both human food and livestock forage derived from native perennial grasslands [11]. VWR is currently used in roadside plantings, in land reclamation after mining, and to restore prairie, wetland, oak forest, and savanna habitats in the Midwestern US.
Populations of grassland birds have declined significantly across the US [14,15,16,17,18]. These decreases reflect declines in the extent and health of perennial native grasslands. In the Midwest and Western US, these losses stem from both conversions to row crops and continuous grazing or overgrazing. Where corn, wheat, or planted non-native pastures replaced native grasslands, biodiversity and ecosystem functions have declined [19,20,21,22,23]. Declines in obligate grassland birds have been accompanied by similar reductions in plant diversity as well as declines in diversity and populations of native dragonflies and butterflies as documented in the US, United Kingdom, and elsewhere [24,25,26,27,28].
Native grasslands currently cover less than 1% of the land that they did historically. The prairie ecosystem was expansive: in the US there were once ~142 Mln acres (57.5 Mln ha) and in Wisconsin 2.1 Mln acres (0.850 Mln ha) of prairie [29]. Extensive conversions occurred in the northern Great Plains following European settlement, with a second surge occurring recently in response to US crop subsidies for grain ethanol [30]. Remaining native grassland cover continues to decline in response to wildfire suppression, landscape clearing, ex-urban land development, tree plantations, and drainage by ditching and tiling for crop production. Native grassland remnants left fallow and unmanaged are often quickly invaded by non-native plant species or succeed to brush or forest cover [31,32,33]. Remaining native grasslands in the Southeastern and Midwestern US are small and fragmented [34]. The Prairie Strips initiative (Iowa State University—[35]) and efforts by the Southeast Grassland Initiative [36] and others (e.g., [37]) seek to restore some of these losses. Nevertheless, most lands once occupied by native grasslands remain dominated by non-native grasslands, annually planted crop fields, and young to middle-aged forests.
We hypothesized that VWR fields would have higher native plants, birds, butterflies, and dragonflies richness and abundance compared to corn and alfalfa crop fields. We compared grassland bird, butterfly, dragonfly, and plant species between fields of perennial native VWR, a short-statured native perennial, and neighboring prevalent fields growing either corn, an annual crop, and alfalfa, a short-lived perennial legume. This study utilized the only large commercial production of VWR in southern Wisconsin that is being evaluated as a new perennial grain crop and as a soil conditioner in the Midwestern US [11]. We use the results of this study to suggest improvements in farmland management to enhance ecosystem health, improve ecosystem services, and better regenerate soil health and biodiversity.

2. Methods

2.1. Study Field Selection and Agricultural Matrix Assessment

In this study, we used farm fields located within a homogenous agricultural landscape in S-central Wisconsin (Figure 1), that included fields planted to corn, alfalfa, and to VWR (Figure 2). All sites and the larger landscape historically were dominated by native prairie and oak savanna vegetation until the late 1800s when the land was plowed and converted for wheat production. In the early 1920s, a rotation of corn and oats was planted, which changed in the 1950s with increased dairy production, as alfalfa (for forage) and soybean, became part of a rotation, now focused on corn and alfalfa. VWR production was introduced in 2023 only at the VWR sites. All crop type sites are bordered by fence rows with shrubs and tree cover and are traversed by drainageways that can flood into the crop field after major storm events. All sites have large rural residential lots within 0.5 to 1 km with lawns, a home and garage.
Each conventional agricultural site was at least 20 ha (50 ac) and abutted thousands of hectares of the same crop types; VWR abutted this same matrix of conventional crops and was planted in five sites for a total of 73 ha (191 ac) in this study area. This study site scale supported randomly placing up to five independent sample points within each crop type, for sampling birds, butterflies, and dragonflies. Vegetation sampling occurred randomly at two selected of the five sampled locations within each crop type. The VWR study site (Figure 2) is located ~2 km north of Brodhead, Wisconsin on Green County Highway E, was planted to VWR after decades of conventional corn, soybean, and alfalfa rotation. All sites have similar hydrology to agricultural ditching to convey stormwater although parts of each study site still flood, often annually. The 100-acre (40.4 ha) alfalfa and 100-acre (40.4 ha) corn sites are located approximately 4 km north of the VWR field (Figure 1 and Figure 2). Randomly selected corn and alfalfa sites are typical for regional conventional production operations, and crop yields found over the region’s large acreage of alfalfa and corn (Figure 1 and Figure 2) in the study region (USDA, NRCS).
Each site includes multiple fields and farming operators; before VWR planting, this same cropping history occurred over the VWR fields in this study site. We selected the conventional crop study sites as typical, local, and with representative cropping procedures and histories on land with similar topography, acreages, soils, and land with similar historic land management, and nearby vegetation and hydrology settings (Table 1). The similarity was confirmed by comparing physical characteristics (slope, aspect, topography, landforms, soil, geology, hydrology, and land cover) and land cover/use between 1937 and 2024 and their cropping and land management history over the last 15 years using USDA, NRCS Soil Web Survey, and StratifyX™ (Table 1) [38]. Stratifyx is a software application (Stratifyx corporation) used during site screening, farmer discussions, and mapping to confirm land use history and biophysical setting (Figure 1 and Figure 2).
In spring 2023, VWR was planted into previous years corn crop stubble. The alfalfa was planted in 2021 after five years of successive annual corn crops. All sites were planted using no-till seeding. The alfalfa and corn sites used herbicide, fertilizer, and fungicide during establishment or production. This involved applications of a pre-emergent herbicide (over Roundup Ready corn Glyphosphate is used) and nitrogen-phosphorus-potassium fertilizer with 120+ lbs. (54 kg) of nitrogen (N) as anhydrous ammonium per acre. The Alfalfa site is mowed typically three times annually for hay crops (Table 1).

2.2. Sampling Design

We randomly located vegetation sampling transects and faunal sampling plots using the biophysical maps at each site, including proximity to wetland and floodplain habitats, and habitat changes using StratifyX™ [38]. All sampling locations were placed 100+ m from any habitat changes such as crop field edges, and 500 m from one another. We used GPS to locate transact ends for vegetation sampling and random magnetic compass headings to randomly locate two 50 m linear transects, in a single randomly selected of the five study location sites in each of the five independent study fields sampled in each cropping system. To sample vegetation, we placed ten 1 m2 circular quadrats along each transect at 10 m interval (total = 20 vegetation sampling plots representative field). In each of five replicated crop type fields we placed two randomized points where we sampled birds, butterflies, and dragonflies. These points were separated by at least 500 m to ensure independent sampling. For purposes of statistics and to eliminate concern of pseudo-replication, all fauna data are summarized and evaluated with reference to the two vegetation sampling locations within each crop type.
Because of the differing phenology of each crop and crop field, we recorded faunal group uses during the rapid plant structural changes during growth and pre-maturation that occurred during the survey period. Planted in early May, corn height was 2 m during the first survey and >3–4 m in height during the last survey 11 days later; alfalfa height was 0.25 m during the first survey, then was mowed for hay to 2–3 cm of the ground shortly after the last survey; VWR was 0.25 m and ~1.25 m in height at the start and end of the survey, respectively.

2.3. Bird, Dragonfly, Butterfly Survey Protocols

Fauna surveys documented the use of each crop type by each faunal group. Direct use (e.g., observed foraging, breeding behaviors, roosting, etc.) were primary indicators of habitat use of each crop type. The key metrics for all faunal groups are species richness, and numbers of individuals, as a measure of relative abundance of use in each crop type.
Surveys included visual observations and detected audible song detection [39] per crop type. Observed and heard birds were recorded for 10 minutes, then binned at intervals of 0–3, 3–5, and 5–10 min. During the bird surveys, the same bird sample points were visually surveyed for dragonfly and butterfly species. To maintain independence between the five sampled points in each crop type, we truncated bird observations to 200 m of each observation point, and dragonfly and butterfly observations to 50 m.
Between 15 and 26 July 2024, bird, butterfly, and dragonfly surveys at five faunal sample points in each crop, representing the five sites surveyed for each crop type were each surveyed four times (5 points × 4 surveys = 20 surveys in each crop type); two surveys at each point were in the early morning, and the remaining two at each point occurred in the afternoons, to capture increased insect activity, and avian foraging activity. For statistical summary and analysis, we used n = 20 for vegetation and faunal samples within each crop type. This was performed to conservatively reduce pseudo-replication concerns, integrate the morning and afternoon surveys for all fauna and to take into account the condensed eleven days of sampling. In total, sixty independent surveys were conducted for fauna in the eleven-day sampling period. This sampling period attempted to normalize plant phenology and to avoid the initial harvest of alfalfa hay which commenced the day after the study ended.
All surveys were completed only during clement weather to avoid high winds and heavy rain. Morning surveys occurred during the early morning (5 to 8:30 a.m. CDT) for all three groups. They were again surveyed in the afternoons when warmer temperatures prevailed (13:00–15:00 CDT), when temperatures were >26.7 °C (80 °F), ensured heightened insect activity. We recorded temperature, wind speed (mph), wind direction (cardinal direction), cloud cover (percent), and any precipitation at the start of each survey. Nomenclature followed the American Ornithological Society [15] for birds; [40] for dragonflies; and [41] for butterflies; these references provide full polynomials plus taxonomic authority reference by readers and as such this information is not included in this paper (See Supplemental Data, Table S1).
We observed both species’ occurrences and behavior from the faunal sample points to infer how birds, dragonflies, and butterflies used the study area. Observations were conducted using 10 × 50 mm binoculars and an 80 mm spotting scope. For each individual or group of individuals observed, we estimated the horizontal distance from the observer, flight direction relative to magnetic north, the numbers and behavior observed. Behaviors have been recorded but not analyzed in this paper: for flying birds, dragonflies, and butterflies, we also estimated height above the ground. Data not analyzed in this paper, such as for birds, included standard categories: perching, flying, foraging, territorial singing, displaying aggression or a distraction display, or carrying fecal sacs and nesting material [42]. This study occurred after the main songbird breeding season, and is thus not a breeding bird survey, but we did pay attention to the few individual male birds still singing and defending territories, that were only recorded once to avoid over-detecting individuals [43]. The plotted distance and direction of each individual aided in minimizing double-counting individual birds, and the insects, especially butterflies. No adjustments were necessary for dragonflies and butterflies: relative abundance was based on the number of individual observations by species during the surveys at each sample point location. Bird, dragonfly, and butterfly species observed during the surveys are listed in Table S1.

2.4. Vegetation Sampling Protocols

Between 25 and 28 July 2024, we also surveyed vegetation and other % ground coverage in 1 m square circular quadrats; we recorded the presence of all species (vascular and avascular herbaceous plants; no woody species present) and estimated their % cover. Percentages of bare soil, fine litter (dead stem litter), coarse litter (>4 cm in any dimension), and rock were also estimated and recorded. Herbaceous plant species frequency was calculated as the percentage of all quadrats occupied by each species we averaged across all quadrats by species in each crop type. We estimated plant species richness as the total and mean number of species per quadrat, averaged across each crop type. We calculated species dominance using percent cover and frequency for each species across all quadrats by crop type. Absolute cover and frequency were relativized to express those as a percentage. To compare plant species, we summed relative frequency (RF) and relative cover (RC) to create an Importance Value (IV) (out of 200%) for each species in each crop type [31]. Plant taxonomy followed [44].

2.5. Statistical Analyses

We first estimated effective detection radii (EDR) for the bird observations using Rdistance [45,46]. This allowed us to determine: (a) the distance at which the likelihood of detection dropped by half for each species, and (b) any difference in detectability among VWR, corn, and alfalfa crop types. For dragonflies and butterflies, a 50 m cut-off distance optimized detectability. Bird species had similar EDRs across all three field types; significant differences only occurred beyond 200 m. We therefore set bird observation distances at 200 m and used unadjusted raw (detection) data using observation frequency to measure relative abundance. We did not compare bird species densities among treatments as estimating nesting density relies on singing territorial males and our observations occurred after the breeding season [47]. Instead, we estimated bird species relative abundance using the number of recorded observations of birds. We estimated relative abundance similarly in dragonflies and butterflies using raw, unadjusted observations.
We used a spreadsheet to enter and check the data and compute preliminary summaries. We compared observed frequencies of birds, dragonflies and butterflies using the described cut-off distances with the raw observation data to calculate unadjusted detections per point. Bird species richness was calculated as the total count of bird species at each point averaged over the twenty survey points in each crop type. We averaged unadjusted detections per point as total counts of birds, dragonflies, and butterflies across all surveys by crop type.
To compare species composition and diversity among the three crop types, we first tallied the recorded abundance for each species and overall species richness. We then checked how % cover (plants) and species count (in each faunal group) were distributed. These were usually Normal with uniform variance within crop types. We nevertheless applied a conservative (p < 0.01) minimum threshold (and report the actual significant of each test) for judging significance in all one-way analyses of variance.
To further characterize differences in community composition among the crop types, we applied multivariate methods including principal components analysis (PCA) and discriminant function analysis (DFA). PCA analyzes the principal axes of variation among the points representing each site in species space. We use bivariate diagrams to visualize differences among the crop types across the first three principal axes (components). The percent of variation accounted for by these three components reflects how efficiently the model accounts for variation among sites in species composition. Discriminant function analysis seeks to best predict crop type based on a linear combination of species’ abundances at each site. Results include the number of sites correctly classified, an entropy r2 value, and a canonical plot showing how species are distributed in prediction space (which species best discriminate among the crop types). Distances among the field type centroids in this space measure how distinct these communities are.
We first applied these analyses separately to the plant data and then merged all faunal groups to obtain a combined analysis. The combined faunal analysis focused on estimating overall responses among all faunal groups to the crop types. This analysis used the measured metrics—species richness, and relative abundance of each species combined across the three faunal groups and ordinated the species as response variables across the vegetation measurements. We performed all analyses and graphic visualizations using JMP Version 18 (SAS Institute, Cary, NC, USA). To summarize and compare plant and faunal group species richness, evenness, overlap, and diversity we also used PCORD v 7.0 [48] to summarize each ecological stratum. Descriptors include Species Richness and Shannon Diversity [49]:
H’ = S pi × ln (pi)
where pi = importance probability of element i (relativized by row total in the species data matrix). They also include Species Evenness calculated as
E = H’/Hmax
where H’ is the Shannon diversity index and Hmax is log Richness or the maximum possible Shannon diversity index for that sample calculated as ln(S), where S is the number of species. Finally, we also calculated Simpson’s diversity index as
D = 1 − S (pi × pi).
We first summarize plant species richness and percentages of cover, frequency, bare soil, rock, and plant litter by crop type. We then compare means among field types for species richness and diversity, evenness, similarity, and percent cover for each species group using one-way ANOVA’s and Student’s “t” test.

3. Results and Discussion

3.1. Vegetation

We first characterize differences in vegetation among native VWR, corn, and alfalfa crops. In the VWR field, native forbs and grasses account for 9/15 of the plant species (60%) and 57% of the total importance values (See Supplemental Data in Table S2). Not surprisingly, VWR had the highest abundance of plant species present (25%). Fine litter and bare soil covered 85% and 15% of the field, respectively.
In the alfalfa crop, perennial alfalfa (Medicago sativa L.) dominated in both cover (71%) and importance (66.7%; See Supplemental Data in Table S3). Another non-native, orchard grass (Dactylis glomerata L.), covered 2.4% while the native annual ragweed (Ambrosia artemisifolium L.) covered 1.2%. Bare soil covered 26% of this field with no fine litter.
Corn dominated the corn crop type, as expected, with 39% of the IV. Two native weeds (ragweed, Ambrosia artemisiifolium L., and pigweed, Amaranthus retroflexus L.) and three non- natives (foxtail grass, Setaria faberi L., crabgrass Digitaria sanguinalis L., and flower-of-an-hour, Hibiscus trionum L.) had a combined IV of 6.6% (See Supplemental Data in Table S4). Bare soil covered 52% of the ground with no fine litter.
We used PCA to characterize differences in plant species composition and cover among the corn, alfalfa, and VWR crop types. The first component accounts for 79% of the variation among these simple communities with the 2nd component accounting for the rest (Figure 3). The corn sites were distinguished by the presence of corn, pigweed (Amaranthus retroflexus L.), and high bare soil cover. The alfalfa is distinguished by alfalfa and orchard grass (Dactylis glomerata L.). The VWR crop type was dominated by VWR (Elymus virginicus L.) with seedlings of giant foxtail (Setaria faberia L.) and small ragweed (Ambrosia artemiisifolium L.). Bare soil cover distinguished these three crop types along the first axis as well.
The VWR field had higher plant species richness, evenness, and diversity than the corn and alfalfa crop types (Table 2, Figure 4). The alfalfa type had the fewest species and lowest diversity. Within individual quadrats, however, all crop types had similar mean species richness and cover although cover varied more among quadrats in the corn and alfalfa crop types (Table 2). One-way analysis of variance revealed that species richness varied significantly among the crop types (F-ratio = 42.2, p < 0.0001, R2 = 59%).

3.2. Faunal Communities

The VWR crop type also harbored the highest abundance and diversity of bird, butterfly, and dragonfly species (Table 3). The corn crop type harbored the least with the alfalfa type intermediate.
Averaged bird abundance and diversity in the VWR crop type were higher than in the alfalfa and corn type (e.g., 2.8 individuals per species in VWR type per survey vs. 1.8–2.0 and richness of 37 vs. 11–12 species, Table 3). These peaked in the VWR crop type and were lowest in corn (Table 4). VWR type shared one obligate grassland bird species with the alfalfa type: the dickcissel.
Some generalist bird species (common grackle, red-winged blackbird, house sparrow, and American crow) visited all crop types. American goldfinch overflew the corn and alfalfa crop types to forage on thistles found along fence rows and rights-of-way borders but did not use either crop type. In contrast, observed foraging goldfinches directly consumed VWR seeds and insects in the VWR crop type.
As with plants and birds, the VWR crop type supported more individuals (n = 106) and a higher diversity (4 species) of dragonflies than the other crop types. Although the most abundant species (a highly mobile species, the green darner) also occurred in the corn and alfalfa crop types, it was sporadic (1–3 individuals vs. 4, Table 4).
Butterfly diversity also peaked in the VWR crop type where six species occurred (five native and the cabbage butterfly; Table 4). However, the alfalfa type had the most butterflies and highest mean number of individuals per survey where two non-native species dominated: the white cabbage looper and yellow sulfur butterfly. These species were both only observed once in the VWR crop type. Overall, these differences were significant and accounted for 14.9% of the variation observed (ANOVA, F = 5.0, p < 0.01).

3.3. Overall Faunal Comparison

Summing across all three fauna groups yields even higher differences in diversity (richness) among the crop types with 7.2 species per survey in the VWR type vs. 2.8 in the alfalfa type and 1.1 in the corn crop type. These differences are highly significant (F = 70.2, R2 = 71%) as are differences in total faunal abundance (summed counts, F = 68.3, R2 = 68%, both p < 0.0001 by ANOVA). This difference in faunal composition is also evident in the DFA where crop type accounts for 77.8% of the variance along the first canonical axis (Figure 4). The first canonical axis cleanly distinguishes the VWR type from corn and alfalfa reflecting the presence there of grassland and wet grassland birds (vesper sparrow, savannah sparrow and sora rail), the blue dasher dragonfly, and the tiger swallowtail butterfly. Most birds occurred more frequently in the VWR crop type, but blue jays and ruby-throated hummingbirds occurred at least once in the corn and alfalfa types.
Bird, dragonfly and butterfly richness, diversity and relative abundances differed among the three cropping systems with VWR consistently scoring higher than either annual corn or perennial alfalfa types. Given that all three had very similar histories of cultivation, these differences in plant and animal diversity and abundance are thought to reflect recent differences in site preparation, planting, crop type structural diversity, and harvest systems. The starkest contrast occurred between the two domestic crops and VWR. These conspicuous differences are believed to stem from differences in tillage, biocide, and fertilizer use, and how fields were harvested relative to the VWR type. These results demonstrate how quickly planting a native perennial crop like VWR can improve biodiversity, including faunal groups with species currently in decline. Given the recent dramatic declines in birds [17,50], dragonflies [24], and butterflies [25] in temperate North America and Europe, our results suggest plantings of a perennial native grassland crop such as VWR may slow or help reverse declines.
Although declines in grassland birds are best documented, dragonflies and butterflies of grasslands and agricultural landscapes have also experienced sharp declines across the U.S. Midwest. The iconic North American migrant monarch butterfly (Danaus plexippus L.) is currently being considered for listing under the U.S. Endangered Species Act reflecting its continental-scale declines. The use of biocides like neonicotinoids and expanded soil tillage disrupting many fencerow habitats have reduced their populations. Disruptive grazing practices have also diminished faunal populations around the globe.

4. Crop Management Affects Vegetative Structure and Ecological Integrity

Corn crops are grown after fields are prepared by first applying pre-emergent herbicides followed by tilling the soil (unless no-till) and seeding typically on 32–36 in. (81–91 cm) centers with corn stems spaced every 7–10 in. (17.8–25 cm) in each row. By the end of our surveys, corn plants were 8–12 feet (244–366 cm) tall with fully developed leaves spaced about every 6 in. (15 cm) from ground level to just below the corn tassel at the top. Heavy shade beneath these plants restricted the growth of ground story vegetation to margins of the crop field, and “skips” where individual plants did not germinate or grow. Ground story vegetation occurred in these gaps, and other than where crop residue from the previous crop year persisted, most of the crop type was dominated by bare soil. Such soils were hard-packed and vulnerable to erosion. The corn field was devoid of dragonflies, butterflies, and bird life except for a few species flying over the field.
Alfalfa crops are also intensely managed typically tilling the soil to generate a fine seed bed which is then drilled on 6–9 in. (15.2–23 cm) row centers. Alfalfa develops into dense 0.5 to ~1 m height plants (61–91 cm) plants followed by 3–4 hay harvests each growing season for dairy cow feed. Few other plant species persist under this management, and it also reduces insect and bird abundance and diversity. Although we observed yellow sulfur and white cabbage looper butterflies emerging and some adults actively foraging on alfalfa, this was temporary and usually lasted for only a week that coincided with alfalfa flowering. Butterfly abundance then declined to zero after haying.
VWR was no-till drilled on 9 in. (23 cm) centers into soy stubble with 2–4 in. (5–10 cm) spacing along rows. After germination, plants produced 9–37 stems (mean = 17) that grew to 2–4 ft. (61–122 cm). As plants grow, lower leaves yellow and wither, allowing some sunlight to reach the ground. This supports a continuous persistent ground cover composed of other grasses and some forbs. Species include non-natives like foxtail grasses (Setaria viridis L., S. faberii L., S. glauca L.) and dandelion (Taraxacum officinale L.) as well as natives like horsetail (Conyza canadensis), small ragweed (Artemisia artemisifollium L.), and some sedges (Carex spp). These species, in turn, support a diversity of insect fauna including grasshoppers, leaf hoppers, soldier beetles, small moths, and other species observed foraging and pollinating the ground story vegetation. The low-growing understory did not compete with VWR, and grew as a living green cover over the ground, ameliorating soil exposure and temperatures in ways that reduced erosion and retained more soil moisture. Although all three crop species dominated their respective fields, only VWR fields supported non-weedy native plant species.

Faunal-Crop Relationships

Corn is harvested for grain feed and alfalfa is grazed, mowed, bailed for hay, and fermented. Few native plant species tolerate the intensive management these two crops require, generating biological and physical conditions that also restrict their use by birds, dragonflies, and butterflies (plus many other taxa and groups of species not assessed). Altering crop management practices can affect how animals use fields. Ref. [51] found that organic practices, smaller fields, and more hedgerows increased bird species abundance and often richness reflecting the higher plant species richness and structural diversity present. Ref. [52] found that adaptive multi-paddock grazing in perennial grasslands boosted both vegetation biomass and bird species richness and abundance. This included rare obligate-grassland breeding birds, aligned with this study finding higher bird, dragonfly and butterfly richness and abundance in VWR fields relative to corn and alfalfa. Collectively, these studies support the idea that vegetation structural diversity is necessary to support many bird species [53,54]. This helps explain why fields that are hayed, mowed, or continuously grazed show declines in bird populations [55,56,57].
Declines in vegetative species and structural diversity may also lead to insect declines as insects appear to respond similarly to birds in cropped systems. That dragonflies and butterflies used the VWR field may be because it emulates native upland grasslands more than corn or alfalfa fields. Faunal groups have distinct nutritional needs during breeding, brood rearing, overwintering, and migration. Each life cycle stage may require different habitat elements to support growth.
Droughts, floods, or other changes in meteorological conditions can make habitats unsuitable, reducing survival and reproduction. In addition, corn and alfalfa cultivation involve using biocides and repeated hay harvests for alfalfa which reduce patch and structural heterogeneity, nectar resources, and seeds, further limiting insect and bird abundance. Whole-plant harvests in corn (such as for silage) and alfalfa further reduce habitat that can support insects that overwinter in soil or dead plant stems.
Crop production is often driven by economic factors which also vary, contributing to dynamic bird, butterfly, and dragonfly population changes [58]. Simple habitat improvements, like hedge rows, uncut grass fields, or prairie strips benefit faunal populations as agriculture intensifies [35,59,60]. Crop diversity and habitat heterogeneity in the UK have proved essential for sustaining bird and insect diversity [61,62,63,64]. The same processes of agricultural intensification, habitat losses, and landscape homogenization appear to be driving butterfly declines in North America and Europe [25,27,28,65]. Declines in monarch butterflies (Danaus plexippus L.) reflect habitat loss and biocide use [66]. Limiting biocide use and enhancing open land habitats, such as roadside verges and hedgerows, will be crucial for maintaining these species. Crop production that avoids biocides [67] could become even more important as habitat heterogeneity and cropland diversity decline [60,68,69,70].
Concerns about dragonfly and damselfly declines have grown as land use changes threaten their aquatic habitats and life cycle stages [24]. Conservation strategies for these species emphasize habitat diversity across larger landscape spatial scales [71]. Supporting insects like dragonflies and butterflies require maintaining continuous habitat gradients from water to uplands [72].
Managing agricultural landscapes wholistically is essential to reverse these declines [26]. Insects provide crucial services for food systems such as pollinators [73], pest controllers, and prey for birds and dragonflies [74]. Agricultural intensification and biocide use are linked to declines in insect, bird, and plant populations that threaten these services [75,76,77]. Climate change, pollution, and invasive species contribute further to these declines [77,78].
Although we did not study bird nesting success, research suggests that successful reproduction also depends on vegetation structure. Dickcissels, eastern meadowlarks, and other grassland birds thrive in open, taller vegetation [79]. We found that obligatory grassland birds used VWR fields more than corn and alfalfa, reflecting such habitat heterogeneity and longevity. Dickcissels perched on corn plants near alfalfa, marginally using both habitats. Grasshopper and savannah sparrows favor habitats with ground litter and grass [79], as found in the VWR field. Taller vegetation and plant diversity supports insects like leaf hoppers, grasshoppers, micro-moths, and bees [80]. In contrast, corn and alfalfa fields lack the vegetation needed by these insect and bird species [80]. While not directly measured, following grain harvests in VWR fields, straw and basal leaves persist in VWR fields as in native perennial grasslands; this type of living and dead standing plant material can provide overwintering habitat for birds, butterflies, and other insects and wildlife [81].
In addition to its advantages for insect, bird, and plant diversity documented here, perennial native grasses also improve soil carbon [52,82,83], forage stocks, and soil-water conditions [84,85]. Perennial native grass crops thereby contribute to soil and water conservation and help ameliorate greenhouse gas emissions [86,87,88,89]. Such fields also support winter livestock grazing, mimicking native grasslands and providing benefits for livestock health and farm management [6,10,18,90,91,92,93]. Such benefits incentivize farmers to adopt native perennial grassland plantings. Even smaller farms could play important roles by providing key habitats for birds and insects foraging near waterbodies, wetlands, and corridors [94,95]. Integrating these efforts at larger scales would also support multiple cropping systems, enhancing landscape-level (gamma) diversity [96,97].

5. Summary

In sum, fields supporting native perennial grassland crops like Viginia wild rye can help reverse declines in bird, dragonfly, and butterfly populations. While further research is needed (e.g., on nesting and life-cycle success for various species), it is already clear that VWR fields offer several ecological and economic benefits over conventional row-crop corn or hayed alfalfa field cropping systems.
We hope the results reported here plus growing concerns for conservation will spark shifts in how farms and agricultural landscapes are managed to better protect biodiversity while sustaining soil health and other ecosystem services. Using native perennial crops to restore grassland bird, dragonfly, and butterfly populations would align conservation with agricultural goals. Expanding these efforts to encompass many to most farms in a watershed could scale-up the value and scope of these restoration efforts to encompass entire agricultural landscapes.
Transitioning to more diverse cropping systems could thus help unite farmers, ranchers, and consumers to better protect biodiversity and ecosystem services while sustaining healthy watersheds and agricultural productivity and economics.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agriculture15151666/s1, Table S1. Birds, dragonflies, and butterflies detected at survey points in taxonomic order; Table S2. Plant species summary in Virginia wild rye field. Based on sampling of 1 m2 quadrats (n = 20); Table S3. Summary vegetation and plant species analysis in the alfalfa field study areas; Table S4. Vegetation summary in corn field study site based on 1 m2 sampled random quadrats (n = 20).

Author Contributions

Conceptualization, data analysis, writing and editing was conducted by co-authors S.I.A. and D.M.W. S.M.L., L.D., K.G., M.B. and G.P. contributed field vegetation data collection under the direction of S.I.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

Archived data is available by request from the corresponding author. Data is provided as private-for-peer review by request with 3 tables provided as supplemental data submitted with the Ms.

Acknowledgments

Interest in native perennial plants for grain production was first brought to the corresponding author’s attention by Wes Jackson at The Land Institute. We thank Wes for the inspiration and decades of research on native plants. We also thank our Indigenous colleagues throughout the Americas who have told us of their rich relationships with native plants as foodstuffs, medicines, ceremonials, and other culturally significant materials. We hope that what we have learned in these studies will contribute to recovering and restoring our collective future legacy of past relationships needed to rekindle, perpetuate, and celebrate native plants and animals.

Conflicts of Interest

Steven Apfelbaum is a volunteer founder of the non-profit, non-commercial research institute, Applied Ecological Institute, Inc. and declares this research was conducted with no commercial or financial interest that could be construed as a potential conflict of interest.

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Figure 1. General location of the study sites in the dashed oval, with color coding denoting crop field types (lavender = alfalfa; tan = corn). Dark green are forested parcels; lighted green and blue are wetlands and waterways, and intermediate green are mixed shrub-dominated fallowed fields, in south central Wisconsin, Green County, USA.
Figure 1. General location of the study sites in the dashed oval, with color coding denoting crop field types (lavender = alfalfa; tan = corn). Dark green are forested parcels; lighted green and blue are wetlands and waterways, and intermediate green are mixed shrub-dominated fallowed fields, in south central Wisconsin, Green County, USA.
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Figure 2. Bird, butterfly and dragonfly sampling occurred in each of the numbered Corn (C1–C5), Alfalfa (A1–A5) and Virginia wild rye (V1–V5) study sites by crop type. Vegetation sampling occurred at the points A-3, C-3 and V3 as randomly selected points.
Figure 2. Bird, butterfly and dragonfly sampling occurred in each of the numbered Corn (C1–C5), Alfalfa (A1–A5) and Virginia wild rye (V1–V5) study sites by crop type. Vegetation sampling occurred at the points A-3, C-3 and V3 as randomly selected points.
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Figure 3. Principal components analysis for total plant cover and plant species richness clearly separated the three crop types. Coded names (AMARET-Amaranthus retroflexus L. SETFAB-Setaria faberi L. BIDFRO-Bidens frondosa L. AMBART-Ambrosia artemisifolium L. DACGLO-Dactylis glomerata L.) are further defined in Table S1 (See Supplemental Information).
Figure 3. Principal components analysis for total plant cover and plant species richness clearly separated the three crop types. Coded names (AMARET-Amaranthus retroflexus L. SETFAB-Setaria faberi L. BIDFRO-Bidens frondosa L. AMBART-Ambrosia artemisifolium L. DACGLO-Dactylis glomerata L.) are further defined in Table S1 (See Supplemental Information).
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Figure 4. Discriminant function analysis of vegetation and faunal associations.
Figure 4. Discriminant function analysis of vegetation and faunal associations.
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Table 1. Farm management details for each crop study field.
Table 1. Farm management details for each crop study field.
ActivityCrop
VWRCornAlfalfa
PlantingMay 2023AnnuallyApril 2021
Fertilizer UseThrough 2022 crop yearThrough 2024 crop yearThrough 2021 crop year
Biocide UseOnly herbicide for site preparation prior to planting VWRAnnually for decades for weed control Fungicide applied annuallyAnnually for decades during the corn
rotation period. None applied since 2021
conversion to alfalfa
HarvestSeed. Fall 2024Entire plant annually for corn seed crop and stoverThree times annually
Prior Year’s CropCornCorn/soybeansCorn
Table 2. Comparison of mean plant species importance value (sum of relative % cover and % frequency—see methods) and overall vegetation richness (S), evenness (E), and diversity (H’ & D (Simpsons Index), from the three crop treatments.
Table 2. Comparison of mean plant species importance value (sum of relative % cover and % frequency—see methods) and overall vegetation richness (S), evenness (E), and diversity (H’ & D (Simpsons Index), from the three crop treatments.
Field TypeMeanStand. Dev.SEH’D
Corn4.36513.23970.5431.0570.5740
Alfalfa4.35215.44040.5310.7370.4331
VWR4.3618.43916 0.708 1.963 0.8008
Table 3. Bird, dragonfly, and butterfly abundance and diversity compared among the three crop types.
Table 3. Bird, dragonfly, and butterfly abundance and diversity compared among the three crop types.
Variable# of SurveysBirdsButterfliesDragonflies
Field Type: CornAlfalfaVWRCornAlfalfaVWRCornAlfalfaVWR
Total Individuals20571456082301013106
Mean observations
(s.d.)		201.78
(1.64)		1.99
(2.26)		2.84
(3.42)		12.73
(1.62)		1.67
(0.82)		113.93
(5.11)
Species Richness20121137226114
Table 4. Faunal group summary of mean, min, max and Standard Deviation for species richness (S), evenness (E), diversity (H’), and Simpson Diversity Index (D) based on 20 surveys in each crop type.
Table 4. Faunal group summary of mean, min, max and Standard Deviation for species richness (S), evenness (E), diversity (H’), and Simpson Diversity Index (D) based on 20 surveys in each crop type.
Birds: N = 38 Species
NumberNameMeanStand. DevSumMinimumMaximumSEH’D
1Corn1.4473.67455.00000.00017.000110.8051.9290.8086
2Alfalfa3.7118.810141.00000.00031.000110.8031.9270.8292
3VWR15.71136.368597.00000.000193.000360.6542.3420.8364
Dragonfly’s: N = 4 Species
NumberNameMeanStand. DevSumMinimumMaximumSEH’D
1Corn0.5001.0002.00000.0002.0001NaN0.0000.0000
2Alfalfa0.5001.0002.00000.0002.0001NaN0.0000.0000
3VWR26.50037.899106.00002.00083.00040.5210.7220.3665
Butterfly’s: N = 7 Species
NumberNameMeanStand. DevSumMinimumMaximumSEH’D
1Corn4.2868.97630.00000.00024.00020.7220.5000.3200
2Alfalfa0.2860.4882.00000.0001.00021.0000.6930.5000
3VWR1.2861.1139.00002.0003.00050.9461.5230.7654
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Apfelbaum, S.I.; Lehnhardt, S.M.; Boston, M.; Daly, L.; Pinnow, G.; Gillespie, K.; Waller, D.M. Native Grass Enhances Bird, Dragonfly, Butterfly and Plant Biodiversity Relative to Conventional Crops in Midwest, USA. Agriculture 2025, 15, 1666. https://doi.org/10.3390/agriculture15151666

AMA Style

Apfelbaum SI, Lehnhardt SM, Boston M, Daly L, Pinnow G, Gillespie K, Waller DM. Native Grass Enhances Bird, Dragonfly, Butterfly and Plant Biodiversity Relative to Conventional Crops in Midwest, USA. Agriculture. 2025; 15(15):1666. https://doi.org/10.3390/agriculture15151666

Chicago/Turabian Style

Apfelbaum, Steven I., Susan M. Lehnhardt, Michael Boston, Lea Daly, Gavin Pinnow, Kris Gillespie, and Donald M. Waller. 2025. "Native Grass Enhances Bird, Dragonfly, Butterfly and Plant Biodiversity Relative to Conventional Crops in Midwest, USA" Agriculture 15, no. 15: 1666. https://doi.org/10.3390/agriculture15151666

APA Style

Apfelbaum, S. I., Lehnhardt, S. M., Boston, M., Daly, L., Pinnow, G., Gillespie, K., & Waller, D. M. (2025). Native Grass Enhances Bird, Dragonfly, Butterfly and Plant Biodiversity Relative to Conventional Crops in Midwest, USA. Agriculture, 15(15), 1666. https://doi.org/10.3390/agriculture15151666

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