Journal of Insect Conservation (2020) 24:765–777

https://doi.org/10.1007/s10841-020-00250-2

ORIGINAL PAPER

Insect diversity in vineyards, almond orchards, olive orchards, alfalfa

fields, and pastures in the San Joaquin Valley of California
Mark S. Sisterson1   · Donal P. Dwyer1 · Sean Y. Uchima1

Received: 29 April 2019 / Accepted: 10 June 2020 / Published online: 19 June 2020
© This is a U.S. government work and not under copyright protection in the U.S.; foreign copyright protection may apply 2020

Abstract
For many agricultural systems, limited data is available on abundance and diversity of insects that are not crop pests or their
natural enemies. As recent studies suggest that insect abundance and diversity is declining, there is a need to quantify insect
diversity within crop fields to determine what role crop fields play in maintaining diversity. In this study, alfalfa fields, almond
orchards, vineyards, olive orchards, and pastures located in the San Joaquin Valley of California were sampled. Ground sweep
samples were collected from all five habitats. In addition, foliar beat samples were collected from almond orchards, vine-
yards, and olive orchards. In total, ~ 240,000 arthropods were collected with the majority identified to family. Across crops
and sampling methods, 20 arthropod orders and 202 insect families were observed. Hemiptera was the most abundant order
of insect collected, representing an average of 61% of all arthropods collected. Diptera was the most diverse order of insect
collected, with a total of 59 dipteran families observed across crops. Of the 202 insect families observed, 85 were observed
in all 5 habitats (42%), whereas 48 families were observed in only one habitat (24%). Families observed in a single habitat
were often represented by only a few individuals. Principal component analysis indicated that the communities present in
the understory of vine and tree crops were more similar to each other than to the communities observed in pastures or alfalfa
fields. Much of the total insect biomass belonged to a few families that included known agricultural pests. In contrast, most
of the diversity was made up of families present in low to moderate abundance.

Keywords  Community ecology · Agroecosystem · Biological control · Pollination

Introduction diversity of insects that serve as natural enemies or pollina-

Recent studies suggest that insect abundance and diversity
has declined, with agricultural intensification hypothesized
as one of many possible contributing factors (Hallmann et al.
2017; Leather 2018; Sánchez-Bayo and Wyckhuys 2019).
While crop fields are routinely sampled to monitor abun-
dance of economically damaging pests and beneficial insects
(e.g., Landis et  al. 2000; Gurr et  al. 2017; Shackelford
et al. 2013), insects with no clear economic value often go
uncounted. For example, studies quantifying abundance and
Electronic supplementary material  The online version of this
article (https​://doi.org/10.1007/s1084​1-020-00250​-2) contains
supplementary material, which is available to authorized users.
* Mark S. Sisterson
mark.sisterson@ars.usda.gov
1
San Joaquin Valley Agricultural Sciences Center, USDA,
Agricultural Research Service, 9611 South Riverbend
Avenue, Parlier, CA 93648‑9757, USA
tors are common (e.g., Costello and Daane 1999; Benhadi-
Marin et al. 2011; Klein et al. 2012; Saunders et al. 2013;
Shackelford et al. 2013). By comparison, fewer studies have
quantified diversity of non-pest herbivores or detritivores. As
a result, estimates of abundance and diversity of a consider-
able portion of the insect community inhabiting crop fields
is lacking for many agroecosystems.
California has the largest agricultural economy in the
United States, with 6 of the top ten producing counties
located in the San Joaquin Valley (San Joaquin, Stanislaus,
Merced, Fresno, Tulare, and Kern Counties; CDFA 2017).
Much of the agricultural land in the San Joaquin Valley is
occupied by perennial crops that provide temporally stable
habitats that are disrupted by pesticide applications. For
example, grapevines, almonds, and olives have stand life-
times that typically exceed 20 years (Klonsky et al. 1997;
Yaghmour et al. 2016; Fidelibus et al. 2018) and collec-
tively cover more than 380,000 hectares in the San Joaquin
Valley (CDFA 2016, 2018a). Alfalfa fields and pastures are

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also common in the San Joaquin Valley, covering an area
exceeding 195,000 hectares (CDFA 2018b). Stand lifetimes
of alfalfa fields are 3–5 years (Canevari and Putnam 2007)
and stand lifetimes for pastures are presumably measured
in years. Further, analysis of geographic information sys-
tems maps on crop distributions indicates that vineyards,
almonds, and alfalfa are frequently planted in proximity (i.e.,
within 1.5 km of each other) in the San Joaquin Valley of
California (Sisterson et al. 2010).
In the case of tree and vine crops, the insect community
consists of two portions: the portion inhabiting crop foliage
and the portion inhabiting plants growing in the understory
(e.g., Rieux et al. 1999; Simon et al. 2010). Ground cover
present on orchard or vineyard floors consists of weeds
(Hanson et al. 2014) and in some cases cultivated ground
cover (e.g. Daane and Costello 1998). Presence of vege-
tation in the understory of tree and vine crops may affect
insect diversity in two ways. First, presence of ground cover
increases plant diversity, which may affect insect diversity
(e.g., Moreira et al. 2016). Second, as weed species found in
almond orchards, vineyards, and olive orchards are likely to
be similar (Hanson et al. 2014), the communities present on
ground cover in tree and vine crops may be similar.
To describe the insect community found in the San
Joaquin Valley agroecosystem, alfalfa fields, almond
orchards, vineyards, olive orchards, and pastures were sam-
pled. The approach used here was to intensively sample a
limited number of sites of each crop and identify all col-
lected insects to the family level. By intensively sampling
sites, the goal was to account for all families commonly
observed in these five crops based on the sampling meth-
ods used. Emphasis was placed on documenting presence
and abundance of families that are not considered pests
and whose contribution to ecosystem services are poorly
described. Describing the insect community present in the
San Joaquin Valley agroecosystem is important for two rea-
sons. First, surveys provide baseline data for comparison
to future studies. Second, surveys may benefit agricultural
production by documenting presence of insects that provide
ecosystems services affecting crop yield, but whose contri-
butions are overlooked.
Materials and methods
Field sites
Field sites were located in Fresno and Tulare counties,
California, USA (Table 1). In 2012, 4 vineyards, 3 almond
orchards, 2 olive orchards, 3 alfalfa fields, and 4 pastures
were selected for sampling. Sites were sampled for 2 years
or until a site was removed from production. For sites that
were removed from production at the end of the first year, an
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Journal of Insect Conservation (2020) 24:765–777
alternative site with the same crop was selected for sampling
in the second year. Over the course of the study, 5 vineyards,
4 almond orchards, 2 olive orchards, 4 alfalfa fields, and
5 pastures were sampled, with sampling duration ranging
between 9 and 24 months. Field sites were managed accord-
ing to common grower practices. At the end of the study,
growers were contacted and asked to provide information
on application of insecticides, fungicides, and herbicides.
While pastures are spot-sprayed with herbicide (Forero et al.
2015), insecticides are typically not applied. Accordingly,
pesticide records were not requested from pasture managers.
For approximate locations of field sites and a description of
neighboring crops see Supplementary Table S1. For pesti-
cide use history see Supplementary Table S2.
Sampling
In alfalfa fields and pastures, arthropods were collected by
conducting 25 ground sweeps using a 40 cm diameter sweep
net, with 2 to 3 sets of 25-sweeps collected per field on each
sampling date. All collected arthropods and plant debris
were placed into a sealable plastic bag and returned to the
laboratory for processing. In vineyards, almond orchards,
and olive orchards samples were collected using two meth-
ods. First, 25 sweeps of ground cover present in almond
orchards, vineyards, and olive orchards were collected as
was done in alfalfa fields and pastures. At each sampling
location, the sampler began at the edge of the vineyard or
orchard and moved inward ~ 1 m with each pendulum swing
of the net. Specifically, 6–8 sets of 25-sweeps were collected
per orchard or vineyard on each sampling date, with sweep
locations uniformly distributed across each side of the field.
Ground sweeps were not collected if ground cover was
absent due to tillage or herbicide treatment or too short to
sweep. For statistical purposes, ground sweeps not collected
due to absence of ground cover were treated as zeros as the
insects being sampled are unlikely to inhabit bare ground.
In contrast, ground sweeps that were not collected due to
ground cover being too short to sweep were treated as miss-
ing values as no assessment could be made.
To directly sample almond, grape, and olive foliage, beat
samples were collected. Foliage beats were collected by
beating the foliage with a stick while holding the sweep net
beneath the crop canopy. The sampler started at the edge of
the orchard or vineyard and beat the foliage ten times while
walking into the vineyard or orchard. Beats of foliage were
taken on each visit to a vineyard, almond orchard, and olive
orchard, with 6–8 sets of beat samples collected per orchard
or vineyard on each sampling date. Beat sample locations
were uniformly distributed across each side of the field. In
winter when grapevines and almond trees are dormant, the
content of the sweep net was bagged only if arthropods were
present. Sampling began in January of 2012 and sweep and
Journal of Insect Conservation (2020) 24:765–777 767

Table 1  Description of field Cropa Cultivar(s) Date planted Period sampled

sites and sampling duration
Grape
­A1 French colombard and Zinfandel 1985 2012-Jan to 2013-Mar
­A2 Cabernet Sauvignon and Rhone types 1993/1998 2013-Apr to 2013 Dec
C Thompson seedless 1979/2011 2012-Jan to 2013-Dec
D Red Globe 1996 2012-Jan to 2013-Dec
E Thompson seedless 1962 2012-Jan to 2013-Sep
Almond
B Nonpareil, Aldridge 2010/2011 2012-Jan to 2013-Dec
C Nonpareil, Monterey 2011/2012 2012-Jan to 2013-Dec
D Nonpareil, Aldridge, Monterey 2007 2012-Jan to 2013-Dec
F Nonpareil, Sonora, Carmel 1987 2013-Feb to 2013-Dec
Olive
A Koroneiki, Arbequina, Arbosona 2003 2012-Jan to 2013-Nov
B Arbequina, Arbosona 2009 2012-Jan to 2013-Dec
Alfalfa
A Roundup ready NK 2011 2012-Jan to 2013-Jun
B 57N58 2010 2012-Feb to 2012-Dec
E Pioneer 5728 2009 2012-Jan to 2013-Dec
F Dairyland Magnum 80 2012 2013-Jan to 2013-Dec
Pasture
A Grasses NA 2012-Jan to 2013-Dec
B Grasses NA 2012-Jan to 2013-Dec
­C1 Grasses NA 2012-Jan to 2013-Mar
­C2 Grasses NA 2013-Apr to 2013-Dec
D Grasses NA 2012-Jan to 2012-Dec
a
 Crops with the same letter were < 2 km apart. Distances between sites with different letters were substan-
tially > 2 km

beat samples were collected monthly until the end of the Samples are stored at the San Joaquin Valley Agricultural
study in December of 2013. In total ~ 2100 sweep/beat sam- Sciences Center in Parlier, CA.
ples were collected, resulting in the collection of ~ 240,000
arthropods. Analysis

Sample processing
On return to the laboratory all samples were placed into
a −20 °C freezer. To process each sample, the contents
were emptied into a 15 cm diameter petri dish and exam-
ined under a dissecting microscope. Non-insect arthropods
were identified to order (Acari and Araneae). Adult and
juvenile insects were identified to family using Triplehorn
and Johnson (2005). Some juvenile insects can be difficult
to identify to family (e.g., some Diptera, Lepidoptera, and
Coleoptera larvae). For such specimens, identification was
limited to order. In total, 98% of insects were identified to
family. Ideally, Thysanoptera, Collembola, and Pscoptera
should be slide mounted for identification which was not
practical given the quantity collected. Accordingly, family
identification of these three orders is tentative. After iden-
tification all arthropods were preserved in 70% ethanol.
Average abundance per sweep or beat sample was deter-
mined for each insect family. As our goal was to document
presence and distribution across crops, principal component
analyses were used to identify latent variables that distin-
guished between the insect communities observed in each
crop at the order and family level. JMP stats (SAS, Cary,
NC) was used to complete principal component analyses.
The intent was to intensively sample a limited num-
ber of sites to determine total families present based on
the sampling methods used. To determine if sampling
achieved this goal, rarefaction was conducted. If rarefac-
tion curves approach a horizontal asymptote, there is con-
fidence that most insect families that may be captured
via ground sweeps or foliage beats were observed. Con-
ducting rarefaction also provides the opportunity to com-
pare diversity among crops while controlling for num-
ber of insects observed (Gotelli and Colwell 2001). The

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768 Journal of Insect Conservation (2020) 24:765–777

rarefaction program was coded in C++ (Microsoft Visual

C++; Redmond, WA). For each run of the rarefaction
program, a specified number of arthropods was randomly
selected from the pool of all arthropods observed in each
habitat without replacement and number of families
observed recorded. For each specified number of arthro-
pods, 1000 repetitions of the rarefaction program were
completed and 95% confidence intervals determined.
To compare number of families observed in each order
among crops, an approach based on rarefaction was used to
control for differences in number of insects collected in each
crop. To accomplish this, number of families in each order
collected via ground sweeps or foliage beats was estimated
for a standardized number of arthropods. For ground sweep
samples, number of families in each order was estimated
by randomly selecting 5000 arthropods from the pool of
all arthropods collected via grounds sweeps in each crop
and determining number of families observed in each order.
The process was repeated 1000 times and mean number of
families observed per order was determined and 95% con-
fidence intervals constructed. For beat samples, number of
families per order was determined by randomly selecting
1700 arthropods.

Results

Description of the arthropod community in ground

sweep samples at the level of order

A principal component analysis of ground sweep samples

conducted at the order level resulted in 6 components with
eigenvalues > 1 (Fig. 1a). Across crops, 20 arthropod orders
were observed in ground sweep samples, with 12 orders
observed in all habitats (Fig. 2a; Hemiptera, Diptera, Thy-
sanoptera, Hymenoptera, Coleoptera, Aranea, Lepidoptera,
Collembola, Pscoptera, Orthroptera, Neuroptera, and Acari).
Eight orders were not observed in all habitats (Fig. 2a; Man-
todea, Odonata, Ephemeroptera, Dermaptera, Blatodea,
Isopoda, Embidiina, and Strepsiptera).
Score plots of components 1 and 2 provided the most
informative separation of ground foliar communities at
the level of order (Fig.  1a, Supplementary Table  S3).
Ten orders had loadings on component 1 that were > 0.5
(Fig. 2a; Hemiptera, Diptera, Thysanoptera, Hymenoptera,
Coleoptera, Araneae, Lepidoptera, Psocoptera, Neuroptera,
and Odonata). As observations from alfalfa fields had the
greatest component 1 scores (Fig. 1a), the ten orders with
high loadings on component 1 were observed in greater
abundance in alfalfa fields compared to the four other sam-
pled habitats (Fig. 2a). In fact, component 1 was correlated
with total arthropod abundance (F = 85.3, df = 1, 18, P <
0.0001, r2 = 0.85), indicating that the primary difference
Fig. 1  Principal component score plots for ground sweeps (a) and
foliage beats (b) conducted at the order level. For ground sweeps,
component 1 had an eigenvalue of 7.3 and accounted for 36.6% of the
variation among sampling locations and component 2 had an eigen-
value of 3.3 and accounted for 16.7% of the variation among sam-
pling locations. For foliage beats, component 1 had an eigenvalue of
4.1 and accounted for 25.4% of the variation among sampling loca-
tions and component 2 had an eigenvalue of 3.3 and accounted for
20.6% of the variation among sampling locations. Orders with load-
ings > 0.50 on component 1 or 2 are shown in Fig.  2. Sampling of
almond orchards, olive orchards, vineyards, pastures, and alfalfa
fields during 2012–2013
in the community at the order level between alfalfa fields
and the other sampled crops was abundance. Component 1
and 2 scores were similar for vineyards, almond orchards,

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Fig. 2  Mean (± SE) abundance of each arthropod order per sample
for ground sweeps (a) and foliage beats (b). For ground sweeps, a
sample consisted of 25 pendulum swings of the net. For foliage beats,
a sample consisted of ten beats of the foliage. Numbers above bars
indicates that an order had a loading of > 0.50 on principal compo-
nent 1 or 2 (see Fig. 1). Sampling of almond orchards, olive orchards,
vineyards, pastures, and alfalfa fields during 2012–2013
olive orchards, and pastures, suggesting little difference in
abundance of each order (Figs. 1a, 2a).
Description of the arthropod community in foliage
beat samples at the level of order
A principal component analysis on foliage beat samples con-
ducted at the order level had 6 components with eigenvalues
> 1 (Fig. 1b). Across vineyards, almond orchards, and olive
orchards, 16 arthropod orders were observed in foliage beat
samples. Ten orders were observed in beats samples col-
lected from all three habitats (Fig. 2b; Hemiptera, Diptera,
Thysanoptera, Hymenoptera, Araneae, Coleoptera, Neurop-
tera, Psocoptera, Collembola, and Lepidoptera). Six orders
(Fig.  2b; Acari, Mantodea, Orthoptera, Ephemeroptera,
Dermaptera, and Odonata) were observed in one or two
of the three habitats. Orders that were not observed across
all three habitats were generally found in low abundance
(Fig. 2b).
Score plots of components 1 and 2 indicated minor differ-
ences in community composition at the order level (Fig. 1b,
Supplementary Table S4). Observations from olive orchards
had the greatest component 1 scores. Four orders had posi-
tive loadings on component 1 that were > 0.5 (Fig. 2b;
Acari, Collembola, Mantodea, and Odonata), suggesting
greater abundance in olive orchards. Three orders had nega-
tive loadings < − 0.5 on component 1 (Fig. 2b; Coleop-
tera, Dermaptera, and Ephemeroptera), suggesting lower
abundance in olive orchards. Observations from vineyards
had greater component 2 scores than almond orchards. Five
orders had positive loadings on component 2 that were >
0.50 (Fig. 2b; Aranea, Diptera, Hemiptera, Lepidoptera, and
Orthoptera) suggesting greater abundance in vineyards than
in almond orchards. Thysanoptera had a negative loading on
component 2 that was < -0.50 suggesting greater abundance
in almond orchards than in vineyards (Fig. 2b).
Insect diversity at the family level in ground sweeps
and foliage beat samples
Across beat and sweep samples, 202 insect families were
observed, with 85 families (42%) observed in all five sam-
pled habitats. In ground sweeps, 18% of families observed in
each crop were represented by a single individual (Fig. 3a).
In foliage beats, 32% of families observed in each crop were
represented by a single individual (Fig. 3b). Averaged across
crops, 5% of families were represented by > 750 individu-
als in ground sweeps and 1% of families were represented
by > 750 individuals in beats of foliage (Fig. 3). Families
observed in high abundance included insects that are eco-
nomically important pests (e.g., Aphididae, Cicadellidae,
Membracidae, Miridae, Thysanoptera, Curculionidae) and
natural enemies (e.g., Anthocoridae and Braconidae). Most
of the insect biomass observed belonged to a few families,
whereas most of the diversity was comprised of families pre-
sent in low to moderate abundance (Fig. 3). The aforemen-
tioned pattern is typical of a wide range of systems (McGill
et al. 2007).
Rarefaction curves for ground sweep and foliage beats
were decelerating (Fig. 4), suggesting that common families
were observed, and a few rare families were not observed.
On a per individual basis, family diversity in ground sweep
samples was greatest in pastures, intermediate in vineyards,
almond orchards, and olive orchards, and lowest in alfalfa
fields (Fig. 4a). As rarefaction measured diversity on a per
individual basis, family diversity in alfalfa fields appeared
low as Aphididae represented 50% of all arthropods col-
lected in alfalfa. Comparison of rarefaction curves for beat
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770
Fig. 3  Histogram of the number of individuals observed per insect
family for ground sweeps (a) and foliage beats (b). Sampling of
almond orchards, olive orchards, vineyards, pastures, and alfalfa
fields during 2012–2013. Bin ranges were as follows: 1 (1), 5 (2 to
27), 50 (28 to 275), 500 (276 to 750), > 750 (> 750)
samples suggested that family diversity was greatest on olive
foliage, intermediate on almond foliage, and least on grape
foliage. One a per individual basis, family diversity on grape
foliage appeared low as 58% of all arthropods collected in
beats of grape foliage belonged to the Cicadellidae.
Patterns of family diversity within orders was simi-
lar for ground sweep and foliar beat samples and similar
across crops (Fig. 5). Across crops, the greatest diversity
was observed in the Diptera with an average of 32 dipteran
families in a random sample of 5000 arthropods collected
via ground sweeps and an average of 26 dipteran families
observed in a random sample of 1700 arthropods collected
via foliage beats. Diversity at the family level was moderate
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Journal of Insect Conservation (2020) 24:765–777
Fig. 4  Rarefactions curves indicating the number of insect families
observed based on the number of arthropods collected for ground
sweeps (a) and foliage beats (b). Error bars represent 95% confidence
intervals. Sampling of almond orchards, olive orchards, vineyards,
pastures, and alfalfa fields during 2012–2013
for Hymenoptera, Hemiptera, and Coleoptera. For ground
sweep samples, an average of 17 hymenopteran, 15 hemip-
teran, and 10 coleopteran families were observed in a ran-
dom sample of 5000 arthropods. For foliage beat samples, an
average of 14 hymenopteran, 15 hemipteran, and 8 coleop-
teran families were observed in a random sample of 1700
arthropods. The remaining orders were represented by an
average of 3 or fewer families in random samples of 5000 or
1700 arthropods for sweep and beat samples, respectively.
Description of the insect community in ground
sweep samples at the level of family
A principal component analysis conducted at the fam-
ily level for ground sweep samples had 18 components
with eigenvalues > 1. Score plots of component 1 and
component 2 were most informative (Fig. 6a). In total, 65
Journal of Insect Conservation (2020) 24:765–777 771
Fig. 5  Number of insect families observed in each order for ground
sweeps (a) and foliage beats (b). To eliminate effects of insect abun-
dance on family diversity, number of families observed in each order
was estimated by randomly selecting 5000 arthropods from sweep
sample data sets for each crop or 1700 arthropods from beat sample
data sets for each crop and determining number of families observed
in each order. This process was repeated 1000 times and mean num-
ber of families observed in each order was determined. Error bars
represent 95% confidence intervals. Orders represented by a single
family were omitted from the figure (e.g., Dermaptera: Forficulidae,
Mantodea: Mantidae etc.) as were orders that were sufficiently rare
as to not typically be observed in the 5000 or 1700 arthropod sam-
ple (e.g., Ephemeroptera, Odonata, Strepsiptera). Sampling of almond
orchards, olive orchards, vineyards, pastures, and alfalfa fields during
2012–2013
Fig. 6  Principal component score plots for ground sweeps (a) and
foliage beats (b) conducted at the Family level. For ground sweeps,
component 1 had an eigenvalue of 39.9 and accounted for 20.7% of
the variation among sampling locations and component 2 had an
eigenvalue of 22.3 and accounted for 11.5% of the variation among
sampling locations. For foliage beats, component 1 had an eigenvalue
of 27.5 and accounted for 20.9% of the variation among sampling
locations and component 2 had an eigenvalue of 22.6 and accounted
for 17.1% of the variation among sampling locations. Families in
the Diptera, Hymenoptera, Hemiptera, and Coleoptera with loadings
having absolute values > 0.50 on components 1 or 2 are shown in
Figs.  7, 8, 9, and 10. Sampling of almond orchards, olive orchards,
vineyards, pastures, and alfalfa fields during 2012–2013
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families had loadings on component 1 with an absolute
value > 0.50 and 27 families had loadings on component
2 with an absolute value > 0.50 (for loadings see Sup-
plementary Table S5; for raw counts see Supplementary
Table  S6). As observations from alfalfa fields had the
largest component 1 scores (Fig. 6a), families with high
Fig. 7  Mean (± SE) abundance of families belonging to Diptera in
beat or sweep samples. For sweep samples (a), families with an aver-
age abundance across crops of < 0.05 per sample were omitted. For
beat samples (b), families with an average abundance across crops
of < 0.002 per sample were omitted. Numbers above bars indicates
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Journal of Insect Conservation (2020) 24:765–777
positive loadings on component 1 were generally in greater
abundance in alfalfa fields than in the other sampled crops
(Figs. 7, 8, 9, 10). Likewise, as pastures had the largest
component 2 scores (Fig. 6a), families with high positive
loadings on component 2 were generally found in greater
abundance in pastures than in the other sampled habitats
that a family had a loading on component 1 or 2 that had an abso-
lute value > 0.50 (see Fig. 6). Positive values indicate positive load-
ings, negative values indicate negative loadings. Sampling of almond
orchards, olive orchards, vineyards, pastures, and alfalfa fields during
2012–2013
Journal of Insect Conservation (2020) 24:765–777 773
Fig. 8  Mean (± SE) abundance of families belonging to Hymenop-
tera in beat or sweep samples. For sweep samples (a), families with
an average abundance across crops of < 0.05 per sample were omit-
ted. For beat samples (b), families with an average abundance across
crops of < 0.002 per sample were omitted. Numbers above bars
indicates that a family had a loading on component 1 or 2 that had
an absolute value > 0.50 (see Fig.  6). Positive values indicate posi-
tive loadings, negative values indicate negative loadings. Sampling
of almond orchards, olive orchards, vineyards, pastures, and alfalfa
fields during 2012–2013
(Figs. 7, 8, 9, 10). Component 1 and 2 scores were simi-
lar for observations from vineyards, almond orchards, and
olive orchards, indicating that the communities present in
the understory of vine and tree crops were more similar to
each other than to the communities observed in pastures
or alfalfa fields (Fig. 6a).
A total of 55 dipteran families were observed across
crops, with 29 families (53%) observed in all five sampled
crops (Fig. 7a). A total of 23 dipteran families had loadings
with an absolute value > 0.50 on component 1 and 9 families
had loadings with absolute values > 0.50 on component 2.
A total of 41 hymenopteran families were observed across
crops, with 15 families (37%) observed in all five sampled
Fig. 9  Mean (± SE) abundance of families belonging to Hemiptera in
beat or sweep samples. For sweep samples (a), families with an aver-
age abundance across crops of < 0.05 per sample were omitted. For
beat samples (b), families with an average abundance across crops
of < 0.002 per sample were omitted. Numbers above bars indicates
that a family had a loading on component 1 or 2 that had an abso-
lute value > 0.50 (see Fig. 6). Positive values indicate positive load-
ings, negative values indicate negative loadings. Sampling of almond
orchards, olive orchards, vineyards, pastures, and alfalfa fields during
2012–2013
habitats (Fig. 8a). Fifteen hymenopteran families had load-
ings with an absolute value > 0.50 on component 1 and 6
families had loadings with absolute values > 0.50 on com-
ponent 2. A total of 30 hemipteran families were observed
across crops, with 13 families (43%) observed in all five
sampled habitats (Fig. 9a). Twelve hemipteran families had
loadings with an absolute value > 0.50 on component 1
and 3 families had loadings with an absolute value > 0.50
on component 2. A total of 30 coleopteran families were
observed, with 8 families (27%) observed in all five sampled
habitats (Fig. 10a). In total, 7 families had loadings with
absolute values > 0.50 on component 1 and 4 families had
loadings with absolute values > 0.50 on component 2.
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Fig. 10  Mean (± SE) abundance of families belonging to Coleop-
tera in beat or sweep samples. For sweep samples (a), families with
an average abundance across crops of < 0.05 per sample were omit-
ted. For beat samples (b), families with an average abundance across
crops of < 0.002 per sample were omitted. Numbers above bars indi-
cates that a family had a loading on component 1 or 2 with an abso-
lute value > 0.50 (see Fig. 6). Positive values indicate positive load-
ings, negative values indicate negative loadings. Sampling of almond
orchards, olive orchards, vineyards, pastures, and alfalfa fields during
2012–2013
Description of the insect community in foliage beat
samples at the level of family
A principal component analysis conducted at the family
level for samples collected via foliage beats from vineyards,
almond orchards, and olive orchards required 10 compo-
nents to explain 100% of the variance among observations.
Component 1 scores tended to be greater in vineyards than
in almond orchards and olive orchards, whereas component
2 scores tended to be greater in olive orchards than in vine-
yards and almond orchards (Fig. 6b). In total, 33 families
had loadings with an absolute value > 0.50 on component 1
and 24 families had loadings with an absolute value > 0.50
13
Journal of Insect Conservation (2020) 24:765–777
on component 2 (for loadings see Supplementary Table S7;
for raw counts see Supplementary Table S8).
Across crops, 40 families of Diptera were observed in
beat samples, with 19 dipteran families (46%) observed in
beat samples from all three crops (Fig. 7b). Sixteen dipteran
families had loadings with absolute values > 0.5 on com-
ponent 1 and 8 dipteran families had loadings with absolute
values > 0.50 on component 2. A total of 26 hymenopteran
families were observed in beats samples across crops, with
8 families (31%) observed in samples from all three crops
(Fig. 8b). In total, 3 families of Hymenoptera had loadings
with absolute values > 0.5 on component 1 and 3 families
had loadings with absolute values > 0.5 on component 2.
Across crops, 21 hemipteran families were observed, with
12 families (55%) observed in all three crops (Fig. 9b). Five
hemipteran families had loadings with absolute values >
0.50 on component 1 and 4 hemipteran families had loadings
with absolute values > 0.50 on component 2. Across crops,
16 families of Coleoptera were observed in beat samples,
with 5 families (31%) observed in all three crops (Fig. 10b).
Four coleopteran families had loadings with absolute values
> 0.50 on component 1 and one coleopteran family had a
loading with an absolute value > 0.50 on component 2.
Discussion
There is increasing recognition that adoption of cropping
practices that conserve insect diversity may benefit crop
production through improved biological control, pollination
services, and decomposition/nutrient cycling (e.g., Simon
et al. 2010; Kleijn et al. 2011; Lichtenberg et al. 2017; Win-
ter et al. 2018). Designing strategies that conserve insect
diversity within agricultural systems requires an under-
standing of community structure at the agroecosystem level.
Here, the insect communities found in alfalfa fields, almond
orchards, vineyards, olive orchards, and pastures located
in the San Joaquin Valley of California were described. A
total of 205 insect families were observed, with 85 families
observed in all 5 habitats. The insect communities present in
the five sampled habitats had several similarities and some
differences.
Members of the Hemiptera were the most abundant
across crops and sampling methods. High abundance of
Hemiptera was primarily due to presence of herbivores,
particularly from the families Aphididae and Cicadelli-
dae. Diptera was the second most abundant insect order
representing 22% of all arthropods observed and harbored
the greatest diversity at the family level. High relative
abundance and/or diversity of the Diptera at the family
level has been observed in numerous cropping systems
(Sáenz-Romo et al. 2019; Ruano et al. 2004; Prischmann
et al. 2005; Ponce et al. 2011; Poveda et al. 2012; Pocock
Journal of Insect Conservation (2020) 24:765–777 775
and Jennings 2008). Despite high relative contribution of
Diptera to total insect biomass and diversity in agricul-
tural systems, the role of Diptera in agricultural systems
is understudied. For example, a metanalysis examin-
ing effects of diversified farming on insect diversity by
Lichtenberg et al. (2017) reviewed 62 studies, with only
15 studies quantifying abundance of at least a single dip-
teran family. Further, a recent review on the decline of
the entomofauna by Sánches-Bayo and Wyckhuys (2019)
implies that few studies have been conducted on members
of the Diptera with studies on Diptera primarily focusing
on a single family (Syrphidae).
Natural enemies including spiders (Araneae), lacewings
(Neuroptera: Chysopidae), damsel bugs (Hemiptera: Nabi-
dae), minute pirate bugs (Hemiptera: Anthocoridae), big
eyed bugs (Hemiptera: Geocoridae), ladybeetles (Coleop-
tera: Coccinellidae), and parasitoid wasps (e.g., Hyme-
noptera: Braconidae, Ichneumonidae, Eulophidae, and
Pteromalidae) were common and observed in all habitats.
Identification of spiders was limited to order and there is
likely considerable diversity within the Araneae in the San
Joaquin Valley agroecosytem. Costello and Daane (1999)
observed 6 families of spiders and 5 families of predaceous
insects in Californian vineyards. Benhadi-Marin et al. (2011)
observed 14 families of spiders in almond orchards in Por-
tugal. Many of the natural enemy families observed in high
abundance are recognized as important natural enemies in
agricultural systems. By comparison, the role of predatory
Diptera has received less attention (Pfister et al. 2017). Here,
relatively high abundance of five families of predacious/par-
astic Diptera was observed (Dolichopodidae, Empididae,
Tachinidae, Syrphidae, and Pipunculidae). Soares et  al.
(2016) also observed relatively high abundance of Dolicho-
podidae in an experimental vineyard in Brazil and Rieux
et al. (1999) reported that Empididae were common in sam-
ples collected from ground cover in pear orchards in France.
In this study, almond was the only crop that required an
insect pollinator (Yaghmour et al. 2016). The only fam-
ily of native bees observed in almond orchards during our
study belonged to the Halictidae and observations of Hal-
ictidae occurred outside of bloom. Many species of Dip-
tera are associated with flower and may serve as pollinators
(Ssymank et al. 2008; Rader et al. 2016). Several of the
dipteran families observed in almond orchards during this
study are associated with flowers (e.g. Syrphidae, Heleo-
myzidae, Empididae, Drosophilidae, Anthomyiidae, Chloro-
pidae, Ceratopogonidae, Cecidomyiidae, Muscidae; Larson
et al. 2001; Wardhaugh 2015; Hahn and Brühl 2016). Two
observational studies on almond pollination determined
that 17.4% of visits to almond flowers in California (Klein
et al. 2012) and 4.1% of visit to almond flowers in Mallorca
(Spain; Alomar et al. 2018) were by members of the Diptera.
While Alomar et al. (2018) documented that 12 families
of Diptera visited almond flowers, to our knowledge, the
only dipteran family that has been shown to successfully
pollinate almonds are members of the Syrphidae (Henselek
et al. 2018).
One approach to conserving diversity in orchard and vine
crops is maintenance of plants in the understory (e.g., Daane
and Costello 1998; Daane et al. 2018; Winter et al. 2018).
Here, a diverse group of insects was observed on weeds nat-
urally occurring in the understory of vine and orchard crops.
Principal component analyses suggested that the insect com-
munities present on weeds in the understory of vineyards,
almond orchards, and olive orchards were similar. As weed
species present in vineyards, almond orchards, and olive
orchards are likely to be similar (Hanson et al. 2014), it is
reasonable to expect the community inhabiting those weeds
to be similar. Rarefaction indicated that on a per individual
basis family richness in sweeps of ground cover was greater
in pastures, vineyards, almond orchards, and olive orchards
compared to alfalfa fields. As an increase in plant diver-
sity is anticipated to increase insect diversity (e.g., Moreira
et al. 2016), higher family richness on a per individual basis
may be explained by greater plant diversity in pastures and
in the understory of vineyards, almond orchards, and olive
orchards compared to alfalfa fields. Nonetheless, arthro-
pod abundance was greatest in alfalfa fields and abundance
of families common to all crops was generally greatest in
alfalfa fields. Accordingly, it is possible that alfalfa fields
play an important role in the persistence of much of the
diversity present among crops. In fact, alfalfa has previously
been referred to as a ‘field insectary’ which produces large
numbers of generalist predators and parasitoids (Summers
1998).
In this study, ~ 240,000 arthropods were collected, with
the majority identified to family. As sampling methodology
affects the types of insects captured, alternative sampling
methods may yield insect families that were not observed.
For example, pitfall trapping is likely to yield families
that live in close association with the soil surface that are
unlikely to be captured by sweep net. Similarly, insects that
infest fruit, bore into plants, or make galls may be under-
represented in ground sweeps or foliar beats. Thus, our
assessment quantifies insects found on the foliage of ground
cover and crop vegetation. In addition, the field sites used in
this study were in areas with at least two other crops in the
vicinity. Analysis of GIS maps indicates that crop diversity
varies throughout the San Joaquin Valley (e.g., Sisterson
et al. 2008, 2010). As landscape effects are documented to
influence insect abundance (e.g., Shackelford et al. 2013;
Pfister et al. 2017), sampling in areas with higher or lower
crop diversity may be associated with changes in the insect
communities in each crop.
For identification, the keys provided by Triplehorn and
Johnson (2005) were used. Identification of many insect
13

776
families is straightforward. However, in some cases it can
be difficult to distinguish between closely related families,
particularly if antenna, tarsi, or wings are damaged. Given
the quantity of insects identified (~ 240,000 arthropods were
processed), it is unrealistic to assume that no misidentifica-
tions were made. If insects were misidentified, they were
likely placed in a closely related family (i.e., within the same
superfamily). In addition, Triplehorn and Johnson (2005)
may not include recent taxonomic revisions. For example,
Triplehorn and Johnson (2005) recognize only two families
of Diptera within the superfamily Empidoidea: Empididae
and Dolichopodidae, whereas Moulton and Wiegmann
(2007) provides support for dividing the superfamily into
five families (Moulton and Wiegmann 2007).
Agricultural fields are some of the most extensively
sampled habitats for insects, although sampling typically
targets a limited number of economically important taxa.
Thus, life histories and ecosystem services provided by a
few taxa found in agroecosystems are often well described
with much less information available for the remainder of
the insect community. Given concerns about the decline of
insects worldwide, documenting insect diversity in agricul-
tural areas is important in determining what role agricul-
tural fields may play in conserving diversity. As the human
population continues to increase, there is a need for research
that focuses on improving crop yields. The challenge for
agricultural entomologists is to identify insect conservation
strategies that improve crop yields by leveraging the ecosys-
tem services provided by insects.
Acknowledgements  Funding for this work was from United States
Department of Agriculture (USDA) Agricultural Research Service
appropriated Project 2034-22000-012-00D. Mention of trade names
or commercial products in this publication is solely for the purpose of
providing specific information and does not imply recommendation or
endorsement by the USDA. USDA is an equal opportunity employer.
Compliance with ethical standards  in total flying insect biomass in protected areas. PLoS ONE
Conflict of interest  The authors declare that the they have no conflict
of interest.
Ethical approval  This article does not contain any studies with human
participants or vertebrate animals. All authors have approved the manu-
script and agree to submission to Journal of Insect Conservation.
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