Biodiversity and Conservation (2023) 32:987–1004

https://doi.org/10.1007/s10531-022-02534-2

ORIGINAL RESEARCH

Standardized butterfly surveys: comparing transect counts

and area-time counts in insect monitoring

Friederike Barkmann1 · Peter Huemer2  · Ulrike Tappeiner1,3  · Erich Tasser3  ·

Johannes Rüdisser1

Received: 3 May 2022 / Revised: 21 December 2022 / Accepted: 21 December 2022 /

Published online: 6 January 2023
© The Author(s) 2022

Abstract
The observed insect decline, which threatens agricultural productivity and ecosystem sta-
bility, calls for comprehensive international insect monitoring. Monitoring implementation
demands standardisation and the integration of new and innovative methods. Therefore,
we compared two quantitative butterfly survey methods – the commonly applied transect
counts (or ‘Pollard walks’) and more extensive area-time counts. We evaluated the influ-
ence of the two methods on the estimation of biodiversity variables such as species rich-
ness and species abundance to examine whether they could be applied alternatively for
the calculation of butterfly trend indicators. During 576 surveys we conducted 5-minute
transect counts and 25-minute area-time counts simultaneously at 144 different sites in
Western Austria. The estimated relative butterfly abundance of the two methods for 119
observed species showed a strong linear relationship. While we found 2.4 times more
species per site with the more extensive area-time counts than with the transect counts,
we also observed a strong correlation between estimates of local abundance (Pearson’s
r = 0.85) and observed species richness (Pearson’s r = 0.81) based on the two methods.
Area-time counts provide thorough assessments on a given location, enabling a close
connection to specific habitat types and facilitating comparability with other plot-based
biodiversity assessments. They are more suitable than transect counts when aiming to
analyse the drivers of temporal and spatial variability in butterfly occurrence. Furthermore,
area-time counts can be used synergistically for the calculation of international butterfly
abundance trends (e.g., European butterfly indicators), as we found strong linear relation-
ships for all applied indicators with both methods.

Keywords  Austria · Biodiversity · Lepidoptera · Insect sampling · Monitoring methods ·

Pollard walks

Extended author information available on the last page of the article

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Introduction

Insects are a pre-requisite for many ecosystem functions (Yang and Gratton 2014) and
related ecosystem services that are essential for human welfare (Losey and Vaughan 2006;
Macadam and Stockan 2015; Rader et al. 2016). Somewhat surprisingly, recent reports on
insect decline (e.g., Hallmann et al. 2017; Sánchez-Bayo and Wyckhuys 2019; van Klink
et al. 2020) have attracted broader public attention to the global biodiversity crisis and
its consequences (Cardoso et al. 2020). A remarkable decline in insect abundance and
diversity is observed in many different regions and terrestrial habitats (Lister and Garcia
2018; Wepprich et al. 2019; Wagner 2020), including protected areas (Rada et al. 2019).
This is of special concern as it threatens the functioning of ecosystems and the supply of
important ecosystem services, such as pollination, food supply, biological control, and soil
fertility regulation, and of diverse cultural ecosystem services (Noriega et al. 2018). The
documented decline is severe and relevant enough to demand deliberate and consequential
political action (Donkersley et al. 2022). Nevertheless, data on most insect populations and
their development are still inadequate (Eggleton 2020; Goulson 2019).
Long-term data about population trends are missing for most insect species (IPBES
2019). To date, only two out of 26 European biodiversity indicators are directly based on
the abundance and distribution of select taxa: the common bird index and the grassland but-
terfly index (EEA - European Environment Agency 2012). Butterfly monitoring, with divers
methodological approaches, has a long history, with the first schemes starting in Europe and
North America in the 1970s (Brereton et al. 2010; Taron and Ries 2015). Due to extensive
and constantly growing monitoring networks such as the European butterfly monitoring
scheme (eBMS) (Sevilleja et al. 2020), butterflies are one of the best-studied insect groups.
Similar schemes for other insects are lacking even though the importance of such programs
is widely acknowledged by science (e.g., Magurran et al. 2010; Dicks et al. 2016; Breeze
et al. 2021). Recent developments such as the EU pollinator initiative, which proposed a
monitoring mechanism as part of the EU Biodiversity Strategy (Potts et al. 2021) and the
development of the European soil monitoring (Montanarella and Panagos 2021) show the
growing political interest in the monitoring of insect biodiversity.
For the international mapping and monitoring of species populations, the integration of
regional and national observation data is needed (Jetz et al. 2019). The combination of data
from different monitoring programs can provide benefits such as a broader database with
extended spatial and temporal coverage and the potential to draw more robust and general
conclusions (Henry et al. 2008). However, data integration can also be challenging due to
variations in methods, spatiotemporal and taxonomic scope, collection purposes, and ter-
minologies (Guralnick et al. 2018; König et al. 2019). Due to the high value of long-term
monitoring data (Lindenmayer et al. 2012), keeping methods constant over time is often
considered more important than uniform methods in the monitoring network when integrat-
ing existing schemes. Even if monitoring schemes are newly established, other objectives of
the regional programs or differences in local conditions can oppose the use of standardised
methods in a monitoring network (Parr et al. 2002). When integrating monitoring schemes,
the applied methods should be evaluated regarding their compatibility to ensure that reliable
conclusions can be drawn from the data. This is also of importance when studies on differ-
ent species or taxonomic groups combine different methods depending on the appropriate
survey design for the targeted taxa (Montgomery et al. 2021).

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Butterflies are one of the few taxa for which a standardized quantitative sampling method
is commonly applied in many different countries – Pollard walks, a transect count method
(Pollard and Yates 1993). Pollard walks have long been established and used in the eBMS
(Schmucki et al. 2015), North American butterfly monitoring networks (Taron and Ries
2015) and many other studies on butterflies with different objectives (e.g., Bruppacher et
al. 2016; Dániel-Ferreira et al. 2020; Kolkman et al. 2021). The observation protocols in
general follow the original recommendation of Pollard and Yates (1993) regarding weather
conditions and the distance from the observer at which butterflies are counted. The length of
transects differs substantially and ranges from short transects of 50 m (Loos et al. 2014) to
a few kilometres (Herrnando et al. 2016).
Apart from transect counts, area and point counts are widely applied visual survey
methods in the monitoring of butterflies and other insects that can be easily identified in the
field (Montgomery et al. 2021). In contrast to the mostly standardised transect counts, there
are varying applications of the area and point counts. For point counts, observers count all
butterflies around them during a defined period of time. Point counts are now commonly
applied in citizen science projects such as the Big Butterfly Count (Dennis et al. 2017) or
can be applied when sites are difficult to cross, e.g., due to dense vegetation or sensitive
habitats (Montgomery et al. 2021). Implementations of the method differ substantially in
the time spent surveying, the distance from the surveyor at which butterflies are counted
and the shape of the surveyed area (e.g., full or semi-circle) (Henry et al. 2015; Dennis et
al. 2017; Lang et al. 2019).
Counts in a defined area also show varying sampling designs. The main difference in
area counts is that while transect counts have a fixed path that is walked by the observers,
area counts have no such fixed path. In some studies, large areas are surveyed while record-
ing the survey effort (time) but are not covered systematically (e.g., the 4th of July butterfly
count in the US on survey circles with a 7.5-mile radius; Swengel 1990). Area counts as
defined by Hardersen and Corezzola (2014) as plot-based surveys are a more standardized
approach, as a defined amount of time is spent on each survey site. From here on, we use
the term area-time count for such standardized approaches where a patch of a defined size
is surveyed for a defined amount of time. Area-time counts were used for butterfly counts in
studies that considered different taxonomic groups on the same survey sites (Su et al. 2004;
Grill et al. 2005; Marini et al. 2009) or assessed the impact of environmental factors on but-
terfly communities (Debinski et al. 2001; Fiedler et al. 2017). Area-time counts provide a
thorough assessment of species richness at a specific site and are therefore advantageous in
heterogeneous landscapes with small habitat patches. Furthermore, they provide better cov-
erage of rare species, facilitating the analysis of drivers of spatial and temporal patterns in
butterfly abundance and richness. However, transect counts can be implemented more easily
in a citizen science context, as has been done for many decades in different countries (van
Swaay et al. 2020), as transects can very often follow existing trails, and hence, disturbance
of grassland sites can be minimized.
International biodiversity monitoring approaches such as the envisaged EU Pollinator
Monitoring Scheme (Potts et al.2021) call for a standardization of methods as well as an
integration of different methods. Ensuring data comparability is crucial when methodolo-
gies differ (Parr et al. 2002). We therefore compared two commonly applied butterfly survey
methods: the traditional Pollard and Yates (1993) transect counts and area-time counts. For

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this purpose, we analysed data from 144 sites from a newly established butterfly monitoring
scheme in western Austria where both methods were applied simultaneously.
To evaluate whether transect counts and area-time counts could be applied alternatively
for the calculation of trend indicators in the context of long-term butterfly abundance moni-
toring, we analysed whether and how the two methods influence the estimation of different
biodiversity variables. At the site level, we used variables such as species richness, spe-
cies abundance and butterfly habitat quality. At the regional level, we used variables that
describe the regional population state (and trend) of individual species, such as relative
abundance (summed over all sites) and site occupancy (the number of sites on which a spe-
cies was detected).

Methods

Study area and survey sites

The study area comprises the Austrian federal states of Vorarlberg and Tyrol in the Eastern
Alps, with an area of 15.241 km2. In each state, 100 survey sites were designated in grass-
land habitats using a stratified random and spatially balanced sampling design. The sam-
pling design aims to representatively survey all grassland habitats of the study region and to
select homogenous habitats at individual sites. This approach allows for the implementation
of probability-based designs – in our case, a stratified random sampling – while maintaining
a spatial balance among the selected locations (Theobald et al. 2007). With a spatially bal-
anced sampling design, the sampling locations are evenly spread over the entire survey area
(or strata). A spatially balanced sampling hence supports efficient surveys when response
variables have a spatial trend, even if the spatial pattern of this trend is not known before the
sample is drawn (Kermorvant et al. 2019). The additional use of strata enables the applica-
tion of variable sampling density if the area covered by the different strata is of substantially
different size, shape or habitat quality. The applied strata should ideally cover more or less
homogenous habitat types or homogenous response units delimited by basic landscape (e.g.,
relief) and land use or land cover (LULC) characteristics (Havlík et al. 2011; Schirpke et al.
2020). This enables us to better account for LULC changes when calculating butterfly trend
indicators, supports a space-for-time substitution to model past or future trajectories of but-
terfly communities where historic baseline data are missing (Blois et al. 2013; Montgomery
et al. 2020) and facilitates future habitat analysis and modelling.
Therefore, we divided all grassland habitats of the federal states of Tyrol und Vorarlberg
into four strata: (1) flat valley meadows with a slope gradient of less than 15%, (2) hillside
meadows with a slope gradient of 15% or more, (3) high mountain to alpine meadows - basi-
cally all meadows above the current timberline, and (4) all grasslands in protected Natura
2000 sites (European Commission 2020). Slope gradient (to differentiate (1) and (2)) was
used as a proxy for management intensity, as the intensity of management practices gener-
ally decreases with increasing slope gradient (Rüdisser et al. 2015). In each federal state, 25
sites (covering all strata) are surveyed per year. Therefore, four years are needed to survey
all sites, and repetition starts in the fifth year. In Tyrol, data are already available from all
100 sites surveyed in the years 2018–2021. In Vorarlberg, monitoring started in 2020 and
hence covered 50 sites from 2020 to 2021.

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Fig. 1  Map showing the location

of the 144 survey sites in the
Austrian states of Tyrol and
Vorarlberg

Butterfly surveys

The butterfly survey took place from 2018 to 2021. To optimize the sampling effort in rela-
tion to the additional information gain, we opted to survey each site four times in one year
(as described above) between mid-May and early September (Barkmann 2020; Hardersen
and Corezzola 2014; Lang et al. 2016; Roy et al. 2007). Observers selected survey dates
flexibly to optimize weather conditions (see next paragraph) and to achieve on average
three to four weeks and a minimum of at least one week between two consecutive surveys.
This approach aims to broadly capture the butterfly species composition of the entire season
(Stewart et al. 2020; Hardersen and Corezzola 2014).
At six of the study sites, only three surveys took place due to unfavourable weather
conditions at high elevations. These six sites were excluded from the analysis, leaving 144
sites (Fig. 1).
The butterfly surveys were carried out between 10:00 and 17:00 in warm (13 C or more),
dry and calm windy conditions (Beaufort scale: 0–3) following Pollard and Yates (1993). At
temperatures below 17 °C, surveys were only conducted in full sunshine. At temperatures
over 17 °C, cloud cover of up to 40% was permitted. To allow a comparison of the two
methods, all surveys were carried out in two phases:
First, a 50-m transect was sampled as described by Pollard and Yates (1993) by walking
slowly at a speed of approximately 10 m min− 1, sampling all sitting and passing butterflies
in an imaginary rectangle of 2.5 m on the sides and 5 m in front. In exceptional cases, the
transect was unilateral, i.e., 5 m on one side. This variant was chosen, for example, if there
was no suitable path through the area and the transect was only possible at the edge of the
area. The resulting walking time per 50-m transect was approx. 5 min.
Second, an area-time count was carried out for an additional 25 min. For the area-time
count, the area around the original transect was extended to 1000 m², i.e., the area 10 m to
the left and right of the transect. During the area-time count, the area was walked slowly in
winding lines, avoiding multiple counting of the same individuals.
Double-counting during the survey cannot be completely ruled out but was avoided as
best as possible. Individuals that were already detected during the transect count were there-
fore not recorded again during the following 25-min area-time count. Although the com-

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plete area-time count of the survey sites comprises both phases of the survey, the two phases
were separated for the data analysis presented herein to obtain two independent datasets.
All butterflies were identified either in-flight or caught and identified in-hand before
release. Animals were caught only if necessary to ensure species identification. If this was
the case, the observer interrupted the survey time to either identify the butterfly, take a
picture or collect for later identification and then resumed the survey where they had inter-
rupted it. Only in exceptional cases were individuals collected for later identification. The
cryptic species pairs Aricia agestis/artaxerxes, Colias hyale/alfacariensis, Leptides sinapis/
realis and Pyrgus malvae/malvoides are difficult to differentiate in the field and were there-
fore treated as species complexes.

Indicators and data analysis

Data analysis aimed to understand and describe the influence of the two different survey
methods on butterfly abundance and diversity estimates. For this purpose, we compared the
results of the 50 m transect counts, which took 5 min, with the results of the 25-min area-
time counts for species richness, butterfly abundance, and butterfly habitat quality (BHQ) at
the site level and regarding individual species abundance and site occupancy at the regional
level. BHQ is an indicator that combines min-max normalized abundance and species rich-
ness and is a suitable metric to compare different sites regarding their general habitat quality
for butterflies (Rüdisser et al. 2017). BHQ was calculated as

(Si − SMin) (Ai − AMin)

BHQi = ∗  (Eq. 1)
SMax − SMin AMax − AMin

where Si is the species richness at site i and Ai is the abundance at the corresponding site.
SMin and SMax are the minimal and maximal species richness of all sites, and AMin and
AMax are the minimal and maximal species abundance of all sites.
For all analyses, data from the four visits per site were summed. For individual species
abundance at the regional level, all counts were summed, and site occupancy was deter-
mined by identifying the number of sites where the specific species occurred at least once.
For each of the abovementioned variables, the correlation between the data from the
transect count and from the area-time count was calculated using the Pearson correlation
coefficient. For analysis of BHQ, the square root of the indicator was used for both methods
for a less skewed distribution of the data. There were some outliers with a high number of
individuals for the abundance data for both the individual species and survey sites. Thus, the
data from both methods were square-root transformed, and the correlation was calculated
for both the original and the transformed data.
Additionally, we compared the individual-based species accumulation curves of both
survey methods for all four strata. The species abundance data from all sites within a stratum
were summed, and species richness was interpolated and extrapolated as described in Chao
et al. (2014) with 50 bootstrap replications. The data analysis was conducted for all sites and
additionally for the different strata individually.
Data analysis was conducted in R (Core Team 2021). The iNEXT package (Hsieh et al.
2016) was used to calculate rarefaction and extrapolation curves, and the ggplot2 package
(Wickham 2019) was used for data visualisation.

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Fig. 2  Scatterplots of the regional indicators for 119 species obtained with data from the transect counts
and with data from the area-time counts; r = Pearson’s correlation coefficient; dashed line: 1:1 reference
line; solid line: linear regression line; p value < 0.001 for both. a: square root of the abundance per species
over all survey sites; b: site occupancy (number of sites a species was detected on)

Results

During the 576 surveys, we made 9817 butterfly detections and observed 119 species. A
total of 7499 butterflies were counted during the area-time counts and 2318 during the tran-
sect counts. The time between two consecutive surveys at the same site ranged from 7 to
63 days, with a mean of 23.9 days (SD: 8.9 days). For 95% of two consecutive surveys, the
time between the visits ranged from 11 to 42 days.
The butterfly abundances detected per site with the area-time counts ranged from two
to 211 (mean: 54.2, SD: 41.9) and with the transect counts from one to 103 (mean: 16.6,
SD: 15.7). With the area-time counts one to 30 (mean: 12.8, SD: 6.2) and with the transect
counts one to 18 (mean: 6.4, SD: 3.8) species were detected per site. Individual species were
detected on average at 2.3 times more sites with the area-time counts than with the transect
counts. On average, 2.4 times more species per site were detected with the area-time counts
than with the transect counts (mean of the proportion calculated for individual sites).
Individual species abundances at the regional level, and hence over all survey sites,
ranged from zero to 905 (mean: 63.0, SD: 119.4) individuals detected with the area-time
counts and from zero to 292 (mean: 19.5, SD: 37.1) individuals detected with the transect
counts. With the area-time counts, individual species were detected at zero to 106 (mean:
15.5, SD: 19.7) sites, and with the transect counts, they were detected at zero to 64 (mean:
7.8, SD: 11.6) sites (Supplementary information Table S1). Three species were detected dur-
ing transect counts only, and 20 species were detected during area-time counts only.
The correlation analysis for the data at the regional level showed a strong linear rela-
tionship between the area-time counts and the transect counts, with a Pearson correlation
coefficient of r = 0.96 for the species site occupancy, r = 0.96 for the untransformed species
abundance and r = 0.96 for the square-root transformed species abundance (Fig.  2). The
correlation coefficients for the individual strata ranged from r = 0.77 to r = 0.94 for site occu-
pancy, from r = 0.70 to r = 0.98 for untransformed abundance and from r = 0.84 to r = 0.96

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Fig. 3  Comparison of the species accumulation curves of the two survey methods (area- time count and
transect count) based on the number of individuals recorded. a: flat valley meadows with a slope gradient
of less than 15%; b: hillside meadows with a slope gradient of 15% or more; c: high mountain to alpine
meadows - basically all meadows above the current timberline; d: grasslands in Natura 2000 sites, e: all
sites

for the square-root transformed abundance. All correlations were significant with p < 0.001.
The abundances of two species showed a noticeable deviation: Pieris rapae was counted
more often and Cupido minimus less often on the area-time counts compared to the transect
counts than would be expected from the linear correlation between the two variables.
The rarefied species accumulation curves based on the data of the two survey methods
consistently overlapped for all strata except ‘high mountain to alpine meadows’. Species
accumulation curves based on the area-time count data were largely saturated across all
strata, while those based on the transect data were not saturated (Fig. 3).
At the site level, the correlation analysis also showed a clear linear relationship between
the data from the transect counts and the area time counts, with Pearson correlation coeffi-
cients of r = 0.80 for butterfly abundance per site, r = 0.85 for square-root-transformed abun-
dance, r = 0.81 for species richness, and r = 0.86 for the square root of BHQ (Fig.  4a-c).

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Fig. 4  Scatterplots of the biodiversity indicators over 144 survey sites obtained with data from the transect
counts compared with data from the area-time counts; p value < 0.001 for all correlations. r = Pearson’s
correlation coefficient; dashed line: 1:1 reference line; solid line: linear regression line; a: butterfly abun-
dance at the individual survey sites; b: species richness at the individual survey sites; c: square root of the
butterfly habitat quality indicator (BHQ) at the individual survey sites

The correlations at site level within the individual strata ranged from r = 0.72 to r = 0.87 for
untransformed abundance, from r = 0.79 to r = 0.85 for the square root transformed abun-
dance, from r = 0.73 to r = 0.81 for species richness and from r = 0.80 to r = 0.88 for the
square root of the BHQ. All correlations were significant at p < 0.001.

Discussion

In the face of the observed insect decline, the importance of sound insect monitoring is being
increasingly emphasized. The envisaged establishment of EU-wide pollinator monitoring
(Potts et al. 2021) could be a major and important step towards better insect conservation
(Samways et al. 2020; Warren et al. 2021) and the establishment of comprehensive Euro-
pean biodiversity monitoring. Although there already exists broad experience in butterfly
monitoring, the development and establishment of transcontinental pollinator monitoring is

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still in its infancy. Standardization of methods as well as the development and integration
of new innovative approaches are essential for successful and sustainable implementation.
For this purpose, butterfly survey methods must be compared to assess their efficiency and
mutual compatibility. We added to this evaluation with an extensive comparison between
transect and area-time counts.
Not surprisingly, we observed more species per site with the more extensive area-time
counts than with the transect counts. The 2.4-times higher species richness per site for the
area-time counts also resulted in 2.3-times the number of sites where a species was detected.
This led to a more complete coverage of the butterfly community composition in the differ-
ent strata. The species accumulation curves were saturated across all strata when based on
area-time count data. The species accumulation curves for transect count data were not yet
saturated due to the lower sampling effort, but were very similar to the curves for the area-time
counts for all but one stratum. For the alpine meadows the curves diverge slightly. This might
be caused by the high habitat diversity combined with naturally lower butterfly abundance in
this stratum. However, we assume that species accumulation curves for transect counts would
saturate too with increased sampling effort (more sites or higher sampling frequency).
Both methods – transect counts and area-time counts – led to surprisingly consistent
proportional results regarding relative butterfly abundance (r = 0.96) and species occupancy
(r = 0.96) on the regional scale. The observed strong linear relationship between the local
diversity estimates of abundance (r = 0.80), species richness (r = 0.81), and BHQ (r = 0.86)
based on the transect counts and on the more comprehensive area-time counts is very good
news. This confirms both the robustness of the transect counts and their compatibility with
the more comprehensive area-time counts.
While the transect counts with 50-m-long transects and four surveys resulted in a cumu-
lative survey time of approximately 20  min per transect and year – covering an area of
250 m² – the area-time counts resulted in a cumulative sampling time of 100 min per site and
year – covering an area of 1000 m². However, the strong linear correlation between the data
from the transect counts and from the far more comprehensive area-time counts indicate that
even a few surveys with relatively low effort can be highly informative. This is true for both
population monitoring at the regional scale and for local site comparison regarding diversity
and overall habitat quality.
While documented differences in sampling effort can be accounted for when combining
data from multiple monitoring schemes, differences in the detectability of distinct species
can lead to severe biases (Kellner and Swihart 2014). It is well known that visual survey
methods can underestimate the presence of cryptic or sedentary species (Dennis et al. 2006).
Distance sampling and mark-recapture studies revealed that butterfly detectability during
transect counts varies substantially among species and is influenced by colour, wingspan,
and behaviour (Isaac et al. 2011; Pellet et al. 2012). Therefore, the data obtained from active
visual counting methods do not reflect true butterfly abundance but rather species-specific
relative abundances (van Swaay et al. 2008). In contrast to the transect counts, the area-time
counts do not follow a fixed path and hence could lead to a changed detectability of specific
species via disturbance or better capturing of the spatiotemporal variation of resources (e.g.,
nectar availability, microclimate, water) (Dennis et al. 2006). We showed that this was not
the case. This is in line with Kadlec et al. (2012), who compared area-time counts and tran-
sect counts at ten sites in the Czech Republic and found no difference between the detect-
ability of mobile and imperceptible species between the two methods.

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There was a strong linear relationship between the species abundance counts from the
two methods. Only two species showed a considerable deviation from the linear correlation
(Fig. 2a). For Pieris rapae, more individuals were detected with the area-time counts than
would have been expected from the abundance counted on the transects. In contrast, Cupido
minimus was counted in proportionally higher abundance on the transect counts. This devia-
tion is likely due to mud-puddling, a behaviour that is often observed in adult butterflies and
can lead to aggregations of individuals (Downes 1973; Boggs and Dau 2004). Such aggre-
gations can cause high local densities of adult butterflies on open spots such as paths. This
was observed on one transect in early July, during which 70 individuals of Cupido minimus
were counted on a transect. On the subsequent area-time count, only two individuals were
detected. Consequently, the respective site also deviates from the linear correlation for on-
site butterfly abundance (Fig. 4a). In this case, the transect count likely overestimates abun-
dance, while the area-time count provides a more realistic assessment. In this context, note
that we separated the data from the transect counts and the consecutive area-time counts to
obtain independent variables for the presented methodological analysis. This might have
contributed to an underestimation of the butterfly abundance and species richness estimates
based on area-time counts in relation to the estimates based on transect counts.
The similar detectability of butterfly species with both methods supports the integration
of data from area-time counts and transect counts for the analysis of trends in butterfly abun-
dance. Differences in the time spent surveying and the area of survey sites do not hinder a
comparison and combination of abundance data because, unlike species richness (Gotelli
and Colwell 2001), species abundance is scale-independent. Due to this and other favour-
able mathematical characteristics, such as monotonicity and proportionality to species’
declines and increases (van Strien et al. 2012), the geometric mean of species abundance is
often used for the calculation of indicators. These indicators, such as the living planet index
(Loh et al. 2005) and the UK wild bird indicator (Gregory et al. 2008), generally assess and
combine population trends of multiple species. The EU butterfly indicator for grassland spe-
cies (van Swaay et al. 2019) and additional indicators developed in the eBMS framework
(Schmucki et al. 2015; van Swaay et al. 2020) are also based on changes in the geometric
mean of species abundance.
For the calculation of international butterfly indicators, density estimates of the con-
tributing national (or regional) butterfly monitoring schemes are combined (van Swaay et
al. 2020). A comparison of methods – as was done here – allows us to assess the factor by
which the detected species densities differ between methods and hence enables the conver-
sion of density estimates and facilitates the calculation of collated indicators.
Depending on the aims and requirements of a monitoring scheme, area-time counts can be a
reasonable alternative to the commonly applied transect counts. Transect counts can be imple-
mented much easier in a citizen science context, as has been done for many decades in differ-
ent countries (van Swaay et al. 2020), because transects can follow existing trails and hence
disturbance of grassland sites can be minimized. Shorter transects that stay within the same
LULC class can give a good impression of butterfly habitat quality, as shown in this study, but
more detailed information on butterfly fauna, such as data on rare species, requires more exten-
sive surveys. Area-time counts provide a more complete assessment of species richness and
community composition at a specific site and are therefore especially useful in small-structured
anthropogenic-influenced landscapes. Next to the information gain, area-time counts are also
more efficient regarding the time spent travelling and surveying. Much effort in systematic moni-

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toring does not result solely from the mere survey time but rather from time travelling to sites and
documenting site and weather conditions. Area-time counts allow more thorough assessments
of a site when focusing surveys on a given location. They enable a closer connection to specific
habitat types and better comparability with other plot-based monitoring schemes (Levanoni et
al. 2011; Hardersen and Corezzola 2014). Especially in heterogeneous landscapes, this approach
facilitates the selection of sites that cover a homogenous LULC class and are influenced by the
same local environmental factors. In contrast, transects with a comparable survey effort often
lead through diverse habitats (Caritg et al. 2011; Taron and Ries 2015), which makes it more
challenging to relate monitoring data to environmental variables. In some monitoring schemes,
transects are divided into shorter sections that lead through homogenous habitats to overcome
this obstacle (e.g., Luppi et al. 2018). However, sections of uniform length do not often match
changes in habitat, while sections with variable length impede the comparison of data from
different habitat types due to differences in survey effort. Area-time counts are therefore better
suited than transect counts when aiming to analyse the drivers of temporal and spatial varia-
tions in butterfly species richness, abundance and community composition, particularly on small
scales (Guariento et al. 2022). Going beyond the detection of population trends and investigat-
ing the causes of declines in butterfly abundance is crucial for taking adequate conservation
measures.

Conclusion

Area-time counts and transect counts resulted in similar assessments of relative abundance
and site occupancy on regional level. Both methods can therefore be used alternatively or
synergistically for the calculation of abundance-based trends. Area-time counts have many
benefits for the analysis of drivers of spatial and temporal patterns in butterfly abundance
and richness because they provide a better coverage of small habitat patches. However,
transect counts can be a valuable complementation to area-time counts in order to enhance
spatial or temporal coverage e.g., in a citizen science context.
Supplementary Information  The online version contains supplementary material available at https://doi.
org/10.1007/s10531-022-02534-2.

Acknowledgements  The Viel-Falter Butterfly Monitoring Scheme is coordinated by the Department of

Ecology at the University of Innsbruck and implemented together with the Natural History Collections of
the Tiroler Landesmuseen Betriebsgesellschaft m.b. H, inatura - Erlebnis Naturschau GmbH and EURAC
research. It is financially supported by the states of Tyrol and Vorarlberg as well as by the private Founda-
tion Blühendes Österreich. Our special thanks go to very committed co-workers and partners: Eva Benedikt,
Ingrid Felipe, Georg Friebe, Valérian Goueset, Eva Hengsberger, Anette Herburger, Kurt Lechner, Walter
Michaeler, Bernd Öggl, Alois Ortner, Constantin Pöll, Johannes Rauch, Petra Schattanek, Daniel Schmid,
Barbara Stoinschek, Ruth Swoboda, Florian Westreicher, Benjamin Wiesmair, Ronald Würflinger, and 73
anonymous volunteers. F.B., J.R., and U.T. are members of the ‘Mountain Regions’ research area at the
University of Innsbruck.

Authors contribution  Friederike Barkmann: Formal analysis, Investigation, Data Curation, Writing- Origi-
nal Draft, Visualization; Erich Tasser: Conceptualization, Methodology, Writing - Review & Editing; Peter
Huemer: Conceptualization, Methodology, Supervision; Ulrike Tappeiner: Conceptualization, Methodology,
Supervision. Johannes Rüdisser: Conceptualization, Methodology, Validation, Formal analysis, Investigation,
Data curation, Project administration, Funding acquisition, Writing- Reviewing and Editing, Supervision.

1 3
Biodiversity and Conservation (2023) 32:987–1004 999

Funding  This work was financially supported by the states of Tyrol and Vorarlberg as well as the Foundation
Blühendes Österreich.
Open access funding provided by University of Innsbruck and Medical University of Innsbruck.

Data availability The published data are available via eBMS data access (https://butterfly-monitoring.net/
ebms-data%20access) or directly upon request with the corresponding author (Johannes.Ruedisser@uibk.ac.at).

Declarations

Competing interests  The authors have no relevant financial or non-financial interests to disclose.

Compliance with ethical standards All samples were collected according to the local environmental law.
Ethical review and approval were waived for this study, as no vertebrate animals were involved.

Open Access  This article is licensed under a Creative Commons Attribution 4.0 International License, which
permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give
appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence,
and indicate if changes were made. The images or other third party material in this article are included in the
article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is
not included in the article’s Creative Commons licence and your intended use is not permitted by statutory
regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright
holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

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Authors and Affiliations

Friederike  Barkmann1 · Peter  Huemer2 · Ulrike  Tappeiner1,3 · Erich  Tasser3 ·

Johannes  Rüdisser1

Johannes Rüdisser
Johannes.Ruedisser@uibk.ac.at
Friederike Barkmann
Friederike.Barkmann@uibk.ac.at
Peter Huemer
P.Huemer@tiroler-landesmuseen.at
Ulrike Tappeiner
Ulrike.Tappeiner@uibk.ac.at

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1004 Biodiversity and Conservation (2023) 32:987–1004

Erich Tasser
Erich.Tasser@eurac.edu
1
Department of Ecology, Universität Innsbruck, Sternwartestraße. 15, 6020 Innsbruck, Austria
2
Tiroler Landesmuseen Betriebsges.m.b.H, Sammlungs- und Forschungszentrum,
Naturwissenschaftliche Sammlungen, Krajnc-Straße 1, 6060 Hall, Austria
3
Institute for Alpine Environment, Eurac Research, Viale Druso 1, 39100 Bolzano, Italy

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