Introduction
Bats are the second most diverse group of mammals (Burgin et al., 2018) with 1,500 species
described (Simmons & Cirranello, 2020), and display a wide variety of complex vocal
behaviours. They actively sense their surroundings using echolocation during both individual
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and social foraging (Jones & Holderied, 2007; Petrites et al., 2009; Egert-Berg & Hurme et
al., 2018; Krivoruchkoetal et al., 2024), and display complex vocalizations related to social
structure and mating (Knörnschild et al., 2017; Knörnschild et al., 2019; Smotherman et al.,
2016). Bats are keystone species that provide important ecological services such as pest
control, soil fertilisation, seed dispersal and flower pollination (Ghanem & Voigt, 2012;
Ancillotto et al., 2017;Ramírez‐ fráncel et al., 2021). Loss of bat ecosystem services can
have direct and profound effects on human health (Frank, 2024). Globally, numerous bat
species are currently threatened by extinction (IUCN, 2024) and all European bat species
are listed on Annex IV of the European Union’s Habitats Directive (92/43/EEC), requiring
member states to monitor them and ensure favourable population status. There is increasing
evidence that migrating bat species face increased mortality from expanding wind farms
(Arnett et al., 2015; Voigt et al., 2022), as countries around the world have to balance the
conservation of flying species against the need to shift energy production to renewable
sources. Other threats that bat species face include climate change (Sherwin et al., 2012),
disease (Hoyt et al., 2021) and declining insect populations (Forister et al., 2019).
Addressing these challenges requires the development of robust methods for monitoring bat
behaviour, activity, and populations.
Recording behaviour of bats which fly in the dark is challenging and monitoring population
sizes becomes especially difficult for bats that roost and breed in cavities and potentially
migrate long distances (Russo et al., 2018; Martínez-Fonseca et al., 2024; Rebolo et al.,
2024). The most cost-efficient way to record bat activity on a large scale is the use of
passive acoustic monitoring (PAM), which has become a well established method (Biffi et al.,
2024, Roemer et al., 2025, Toshkova et al., 2025). Most bat species produce ultrasonic
echolocation calls and social vocalisations that may span the audible and ultrasonic range.
Echolocation calls can be assigned to three main categories. Search phase calls are usually
species-specific but adaptable to the environment, with longer calls characteristic for open
space foraging and commuting and shorter call duration typical in cluttered environments
(Kalko and Schnitzler, 2001; Brinkløv et al., 2010). Approach calls are similar to calls
produced in clutter, but might shift in frequency. The call interval (also called pulse interval) is
also much reduced to yield quicker updates about prey or obstacle position. Finally, buzzes
are produced during the very end of a prey interception, with much reduced call amplitude
and call intervals. Social vocalisations are much more variable in some species and consist
of longer multi-node sequences resembling bird song (Smotherman et al., 2016; Springall et
al., 2019). Species can in some cases be recognised by the frequency range and
modulation, the duration and the intervals between calls, while specific behaviours can
sometimes be inferred from the type of call: buzzes mostly indicating feeding, social calls
indicating varying social behaviours such as mate attraction, territorial defence and individual
recognition (Knörnschild et al., 2017; Knörnschild et al., 2019; Smotherman et al., 2006).
While PAM is already effectively supplying vast amounts of audio data, the processing
workflow often lags in efficiency. Approaches range from manual detection and classification
of bat calls (Sugai et al., 2018) to more or less automated pipelines with or without manual
validation of software performance. The applicability of commercial software is challenged by
price and decreased transparency. Open source software also exists, but often comes
trained for a limited geographic range focused outside the Global South, where bat species
diversity is highest, and might not be easy to adapt and use without a computer science
background. An exception to this is BatDetect2 (Aodha et al., 2022b), which contains
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pre-trained models for several countries, including two in the Global South. It does, however,
require some familiarity running scripts from the console, especially if fine-tuning of the
model is needed. For all these software, retrainability remains an issue. If used in a
soundscape with new noise sources or bat species, performance is likely to drop. Retraining
without coding experience is often not possible. Finally, all but one of the open source
solutions have focused on search phase calls, with the exception of Jameson (2024), who
developed a buzz detector: buzzfindr for Canadian species. This lack of tools to detect social
calls and buzzes globally hinders bat research, since foraging or social activity are often
important behavioural components in a monitoring context (Dietzer et al., 2024).
To address this, we have trained several versions of BatSpot, which is based on
ANIMAL-SPOT, a convolutional neural network designed to detect and classify animal
vocalisations (Bergler et al., 2022). We present detection models for bat echolocation search
phase calls, buzzes and social calls, as well as a classification model for the search phase
calls. We compare the performance of these models to several commercial and open source
solutions on a validation dataset. We provide examples on how to validate the models for a
new location and how to retrain the model with a reduced amount of training data. The
model also allows for transfer learning, adding additional classes to the classification model.
Finally, we provide a GUI (see Figure 1) and a README on how to train, retrain,
transfer-train and validate models.
Figure 1 Screenshot of the GUI.
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Methods
Call detector
To detect search phase calls in continuous recordings, we trained a binary version of
BatSpot (hereafter called ‘call detector’). We used 12,396 noise examples and 10,766 target
examples (search phase calls) from Denmark. We also used additional 2,000 noise
examples for noise augmentation (these are added randomly in the background to increase
generalisation ability). These were chosen randomly from the original noise set (with 14,396
noise examples). The model was trained on clips of 20 ms duration. Longer examples were
randomly cut, shorter examples were zero-buffered. The frequency range was 1-95 kHz, this
covers the high-energy part of all calls and is enough for detection. We report the validation
and test accuracy, which is the proportion of true positives and true negatives out of the total
number of validation/test examples. The reported validation accuracy is the highest accuracy
during the training process, where the validation set is used to test if the model should stop
training. The test accuracy is the accuracy on the test set, which is not shown to the model
until after training has stopped.
To validate the model performance on full recordings we used 210 recordings from Denmark
of variable duration. None of these had been used to generate training data. All search
phase calls (including faint calls) were manually annotated. The call detector was run on
windows of 20 ms, with 10 ms overlap between windows. The threshold for considering a
detection true was 0.5. For each detection BatSpot gives a probability that it is correct in the
prediction and by choosing 0.5 we allow only predictions where this probability is at least 0.5.
Since the call detector and classifier are often used together, validation results are also
presented together. For more detailed information, see below.
Call classifier
To classify the detections from the call detector, we trained a multi-class version of BatSpot
(hereafter called ‘call classifier’) including call examples of 12 species(-complexes). We used
1,161 examples of Barbastella barbastellus, 2,840 examples of Eptesicus serotinus (now
known as Chenaphaeus serotinus), 1,217 examples of Myotis brandtii/mystacinus, 1,503
examples of Myotis dasycneme, 2,401 examples of Myotis daubentonii, 1,325 examples of
Myotis nattereri, 2,397 examples of Nyctalus noctula, 982 examples of Plecotus auritus,
1,226 examples of Pipistrellus nathusii, 1,145 examples of Pipistrellus pipistrellus, 2,374
examples of Pipistrellus pygmaeus and 1,517 examples of Vespertilio murinus. Examples
were drawn from the recordings made during the Danish national monitoring programme,
where the focal individual is either observed in the field during recording or, if data quality
allows, annotated by at least two experts. We also included 551 examples of buzzes, 899
examples of approach phase calls and 439 examples of social calls. These categories were
not annotated to species level and considered as noise, since the call detector model was
only supposed to detect search phase calls. To get optimal performance the approach phase
call category was encoded as the noise category, serving as the default, if none of the other
categories is predicted with high enough probability. The model was trained on clips of 20
ms duration. Longer examples were randomly cut, shorter examples were zero-buffered.
The frequency range was 10-125 kHz, which meant that we had to use a slightly modified
version of BatSpot, where the files were not downsampled to 192 kHz during processing, but
rather to 250 kHz. This was done, since some species have calls extending beyond 96 kHz,
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information that can be used by the model during classification. We report the validation and
test accuracy, which is the proportion of correct classifications out of the total number of
validation/test examples.
To validate the model on detections from full recordings, we used the same validation set as
for the call detector, and ran the call classifier on all the detections from the call detector.
Since several Myotis species are notoriously difficult to discriminate from each other, we
pooled both the validation and classifications from the model into the genus level label M.
Depending on context, the same may be true for the low frequency species E. serotinus, N.
noctula and V. murinus, thus these were pooled into an ENV complex. Calls assigned to B.
barbastellus were considered noise, since they should not occur in the area where the
validation data was collected. If calls could not be assigned to a species (-complex) or
multiple calls were overlapping, these were excluded from validation. The call classifier was
run on windows of 20 ms, with 15 ms overlap between windows. The overlap was increased
to increase the chance of the classifier getting a window with the whole call included. The
threshold for considering a detection true was 0.5. To quantify performance we created a
confusion matrix and computed performance statistics. For the detection step we computed:
(1) recall = tp/(tp+fn), which gives the fraction of calls that were detected out of the total
number of calls; (2) precision = tp/(tp+fp), which gives the fraction of correct detections out of
all detections; and (3) F1 = 2 * (recall*precision)/(recall+precision), which gives the average
across the two; where tp = number of true positives, tn = number of true negatives, fp =
number of false positives and fn = number of false negatives. For the classification step we
computed: accuracy = correct classifications / (correct classifications + mistakes), which also
included classifying the false positives of the detection model (where the correct
classification was noise). For the overall performance, we computed: accuracy = correct
classifications / (correct classifications + mistakes + misses), where the misses were the
false negatives from the detection model. Since some projects only need to know if a
species is present in a file or not, we also summarised the output per file by taking the
species with most calls predicted and created a confusion matrix at file level, where we
excluded 10 out of the 210 files, as they contained calls that could not be assigned within
Pipistrellus (for the call level performance only these few calls were excluded and all other
calls in the file were still used). We opted not to include area-under-the-curve values for
either analysis, as these are harder to interpret.
To compare the combined performance of the BatSpot call detector and classifier to existing
software, we ran the open source software BSG-BAT (Meramo et al., 2025), BatDetect2
(Aodha et al., 2022) and BAT (Fundel et al., 2023), as well as the commercial softwares
Sonochiro (version 4.1.4, Biotope, France, biotope.fr), Kaleidoscope (version 5.8.1, Wildlife
Acoustics Inc, USA, wildlifeacoustics.com) and BTO bioacoustics pipeline (BTO Acoustic
Pipeline, n.d.). We used the same validation set as for the validation of BatSpot. In files with
one or multiple calls that could not be assigned to a species (-complex), it would not be
possible to assign a label at the file level. For each software we used default settings and
followed best practise described in the manual. For BatDetect2, we changed the threshold
for accepting a detection to 0.4 to balance the recall and precision. For each software, we
compared the primary detection and classification to the ground truth (manual annotation by
an expert) at file level. Where the software made multiple predictions, only the prediction
with highest probability was used, since each of the validation files only contained a single
species. If the software predicted species not present in the study area or did not classify the
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detection, the file was considered to contain only noise. For each software we show the
confusion matrix and compute the recall, precision and F1 score.
Buzz detector
To detect buzzes in continuous recordings, we trained a binary version of BatSpot. We used
2,293 noise examples and 1,288 target examples from Denmark, Germany and Panama.
We also used 200 additional noise examples for noise augmentation. These were chosen
randomly from the original noise set (of 1,488 noise examples). The model was trained on
clips of 200 ms duration. Longer examples were randomly cut, shorter examples were
zero-buffered. The frequency range was 1-95 kHz, which contains the part of the buzz with
the most energy for most species. We report the validation and test accuracy, which is the
proportion of true positives and true negatives out of the total number of validation/test
examples.
To validate model performance we used 30 recordings from Denmark of variable duration
(3-15 s), 100 recordings of 55 s from Germany and 90 recordings from Panama of variable
duration (0.2-0.9 s). None of these had been used to generate training data. All buzzes
(including faint and incomplete buzzes) were manually annotated. The buzz detector was
run on windows of 200 ms, with 100 ms overlap between windows. The threshold for
considering a detection true was 0.5 for Denmark and Panama, but was set to 0.9 for
Germany to balance recall and precision. For validation we computed: recall = tp/(tp+fn),
precision = tp/(tp+fp), F1 = 2 * (recall*precision)/(recall+precision), where tp = number of true
positives, tn = number of true negatives, fp = number of false positives and fn = number of
false negatives.
To compare our buzz detector to existing tools we ran the function buzzfindr from the R
package buzzfindr (Jameson, 2024) on the same validation recordings using the default
settings. We show the confusion matrix and compute the recall, precision and F1 score.
Social call detector
To detect social calls in continuous recordings, we trained a binary version of BatSpot. We
used 1,296 noise examples and 1,412 target examples from Denmark. We did not use noise
augmentation. The model was trained on clips of 75 ms duration. Longer examples were
randomly cut, shorter examples were zero-buffered. The frequency range was 1-60 kHz. We
report the validation and test accuracy, which is the proportion of true positives and true
negatives out of the total number of validation/test examples.
To validate the model performance we used 19 recordings from Denmark of variable
duration. None of these had been used to generate training data. All social calls (including
faint calls) were manually annotated. If a social call was not clearly a single unit (parts
separated by ca. 50 ms), each part was separately annotated. The social call detector was
run on windows of 75 ms, with 50 ms overlap between windows. The threshold for
considering a detection true was 0.9. For validation we computed: recall = tp/(tp+fn),
precision = tp/(tp+fp), F1 = 2 * (recall*precision)/(recall+precision), where tp = number of true
positives, tn = number of true negatives, fp = number of false positives and fn = number of
false negatives.
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Retraining
Retraining is the use of a previously trained model to speed up and/or improve performance
of a new model. For retraining, the weights of the old model are imported after which new
training examples are used to (slightly) update these weights. To illustrate how retraining
increases the performance, we ran the existing call detector on a validation set of 24 files of
5 s each from Germany. We then created a training set (using different recordings)
consisting of 290 noise examples and 477 target examples from Germany. We also used an
additional 50 noise examples for noise augmentation. These were chosen randomly from the
original noise set (of 340 noise examples). With that we first trained a model from scratch
using the same settings and then trained a model using retraining from the Danish call
detector model. We restricted the number of learning rounds (epochs) to 100, but did not
change any other settings. To compare we present the recall, precision and F1 for all three
models (baseline, retrained and trained from scratch).
Results
Call detector and classifier
After 150 epochs (the maximum) the call detector had a validation accuracy of 0.98 and a
test accuracy of 0.97. The call classifier trained for 81 epochs (after no improvement for 20
epochs) and achieved a validation accuracy of 0.88 and a test accuracy of 0.87. For
validation of the models on full recordings, we annotated 6,477 search phase calls. Of these,
40 could not be assigned a species, 74 were overlapping, 193 were either P. nathusii or P.
pipistrellus and 490 were either P. pipistrellus or P. pygmeus, and were therefore excluded
from validation. The detector performed well on the validation set, which balanced recall
(how well the model finds calls) and precision (how many detections are correct) (see Figure
2) and showed no strong bias towards missing certain species. The classification model also
performed well, with most errors being due to confusion of P. nathusii and P. pipistrellus or
assigning false positives to ENV or Myotis, likely due to low frequency and click-like noise
both confusing the call detector and classifier.
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Figure 2 Overall confusion matrix for the call detector and classifier. Rows are the manual
annotations, columns are the results from the models for each call. Ppyg = Pipistrellus
pygmeus, Ppip = P. pipistrellus, Pnat = P. nathusii, Paur = Plecotus auritus, M = Myotis sp.,
ENV = Eptesicus serotinus/Nyctalus noctula/Vespertillio murinus, -noise- = calls annotated
as noise or classified as buzz/approach/social/Bbar, -missed- = calls not detected by the call
detector.
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Overall BatSpot outperformed all other tested software (see Figure 3 and 4). BatDetect2 and
the BTO bioacoustics pipeline came close in balancing the recall and precision as well as
accurately classifying the files with detections (see Table 1).
Figure 3 Overall confusion matrix at file level for the call detector and classifier. Rows are
the manual annotations, columns are the results from the models for each call. Ppyg =
Pipistrellus pygmeus, Ppip = P. pipistrellus, Pnat = P. nathusii, Paur = Plecotus auritus, M =
Myotis sp., ENV = Eptesicus serotinus/Nyctalus noctula/Vespertillio murinus, -noise- = calls
annotated as noise or classified as buzz/approach/social/Bbar, -missed- = calls not detected
by the call detector.
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Figure 4 Confusion matrices for BSG-BAT, BatDetect2, BAT, Sonochiro, Kaleidoscope and
BTO bioacoustics pipeline. Rows are the manual annotations, columns are the results from
the different software at file level. Ppyg = Pipistrellus pygmeus, Ppip = P. pipistrellus, Pnat =
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P. nathusii, Paur = Plecotus auritus, M = Myotis sp., ENV = Eptesicus serotinus/Nyctalus
noctula/Vespertillio murinus, -noise- = files annotated as containing noise or any other bat
species.
Table 1 Performance of all tested software on the Danish validation set. Recall = tp/(tp+fn),
precision = tp/(tp+fp), F1 = 2 * (recall*precision)/(recall+precision) and accuracy =
(tp+tn)/(tp+tn+fp+fn) for all files with detections; where tp = number of true positives, tn =
number of true negatives, fp = number of false positives and fn = number of false negatives.
The highest value(s) in each column is/are highlighted in bold.
Detection Classification
Software Recall Precision F1 Accuracy
BatSpot 1.00 0.95 0.97 0.95
BSG-BAT 0.91 0.87 0.89 0.80
BatDetect2 1.00 0.92 0.96 0.92
BAT 0.48 0.76 0.59 0.58
Sonochiro 0.79 0.97 0.88 0.84
Kaleidoscope 0.68 0.97 0.80 0.77
BTO 0.87 0.99 0.92 0.92
Buzz detector
After 100 epochs (the maximum) the model had a validation accuracy of 0.97 and a test
accuracy of 0.96. For the validation recordings performance was generally very high (F1:
0.95, recall: 0.96, precision: 0.93). Performance was highest for the Panamanian recordings
(F1: 0.97, recall: 0.95, precision: 0.99), intermediate for the Danish recordings (F1: 0.88,
recall: 0.85, precision: 0.92) and lowest for the German recordings (F1: 0.84, recall: 0.94,
precision: 0.77). Despite the increased threshold for accepting detections (0.9 instead of
0.5), the model detected a lot of false positives for the German recordings.
Buzzfindr had much lower performance on the same validation data (F1: 0.37, recall: 0.28,
precision: 0.54).
Social call detector
After 150 epochs (the maximum) the model had a validation accuracy of 0.99 and a test
accuracy of 0.98. For the validation recordings performance was lower but still good given
the limited dataset (F1: 0.91, recall: 0.93, precision: 0.98).
Retraining
The call detector trained on only Danish data did not perform very well on the German
validation set (F1: 0.48, recall: 0.88, precision: 0.33). Retraining clearly improved the
precision (F1: 0.79, recall: 0.79, precision: 0.78) and led to slightly better performance than
training from scratch (F1: 0.73, recall: 0.83, precision: 0.64).
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Discussion
Our aim was to develop a tool that can both detect the three most common vocalisation
types of bats (search phase calls, feeding buzzes and social calls) and classify search phase
calls, the vocalisation type most commonly used for species identification and monitoring
studies (Schnitzler & Kalko, 2001; Russo et al., 2018). We present pre-trained models with
reasonable out-of-the-box performance and the option to retrain the models with minimal
time investment. Finally, we show that the tool can also be used in the Global South. The
three detection models presented achieved good performance for the situations that they
had been trained for. The buzz detector in particular showed that a single model could detect
buzzes in mixed habitats recordings in Denmark, Germany and Panama, with very different
noise backgrounds and species compositions.
When applying the search phase call detector model trained exclusively on Danish data to
German recordings, we observed a clear drop in performance. This is to be expected, since
the noise sources can be very different, even within the same country. This is also clearly
what drove the drop in performance, with precision (the proportion of detections that
contained search phase calls) dropping much more than recall. Of course, if the detector is
used in a country with not only new species (as in Germany), but also new genera or even
families that produce search phase calls that do not resemble any of the training examples,
a drop in recall is also to be expected. In both cases, retraining will be needed until a dataset
with enough examples across the globe becomes available and a global bat detector can be
trained. Our hope is that the data presented with this paper can contribute to such a global
dataset, and that until a global model is a reality, our Danish model can serve as a basis
model to retrain detectors for new locations.
Classification is a more challenging task, and one where even experts with decades of
experience cannot always assign call sequences of good quality to a species (Jennings et al.
2008; Rydell et al. 2017). This is simply because some species produce very similar calls
when flying in similar environments, and because a given species is often able to modify
calls when flying in a different environment. We therefore opted to follow the standard
procedure of grouping calls from Myotis species into a single category and also grouping
calls from species producing lower frequency calls into an ENV species-complex. The
classification model was trained with each species separately (with the exception of M.
brandtii and M. mystacinus which could not be separated when generating our training data)
to allow the model to learn as many distinguishing features as possible. The grouping was
then done after the prediction step, since performance within groups was low as expected.
Although an overall accuracy of 0.80 seems low, it should be kept in mind that this includes
detection and classification of even the faintest of calls. For many projects (e.g.
environmental impact assessments), the aim is to detect and classify species at the file level
(as presence/absence in that file), and this is where performance is much higher (F1 = 0.97).
There are also projects (e.g. those aiming to describe the characteristics of calls under
varying conditions), in which exact detection of start and end times for individual calls is
important. Additionally, when detecting buzzes or social calls, one might want a very low
false negative rate, to ensure most target events are detected. This can be accomplished by
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lowering the detection threshold and subsequently validating all detections manually to get
rid of the false positives. Alternatively, one might want a low false positive rate, something
particularly relevant when many calls are produced in a sequence and post-hoc processing
can be done on the detections. One way to do this is to set up a decision tree, in which, for
example, a file is only considered to contain calls from a species if the classifier assigns at
least five calls to that species within a sliding window of 3 seconds. This eliminates any false
positives that occur spaced out in time.
BatSpot offers a GUI and README for how to retrain, to facilitate obtaining a model with
good performance in new recording situations, which is an important step forward and allows
researchers to optimise the models without coding experience. An example of this is the
offshore environment, where boat, buoy and water noise create challenges to both the
detection and classification step by introducing new noise types. With the increase of wind
farm development offshore, and the need for both pre- and post-construction monitoring,
models performing well in the offshore environment are particularly important. This is one of
the reasons that the search phase call detector in BatSpot was trained with a large portion of
examples from buoys and wind turbines in the Danish North Sea. That said, BatSpot is far
from the first neural network to detect and classify search phase calls, although to the best of
our knowledge, it is the first to additionally detect buzzes and social calls. BatDetect2 is a
very promising alternative for search phase calls, and achieves comparable performance
out-of-the-box. One potential reason for BatDetect2’s high performance is the inclusion of a
self-attention layer, which enables the model to integrate information of multiple calls in a
sequence to predict the species producing individual calls. With the integration into a simple
python package, BatDetect2 requires minimal coding experience, and might therefore be the
best option if there is no time for targeted model retraining.
For buzzes and social calls, BatSpot presents a much bigger step forward. The detection of
buzzes is important when looking at foraging behaviour, but has previously received almost
no attention in the literature. The R package buzzfindr (Jameson, 2024) is an important step
forward, but struggles to detect faint buzzes and buzzes in noisy environments, something
important in settings with multiple individuals flying in an environment with much insect noise
as well (as is the case for many of the German and Panamanian recordings). The fact that a
single model was able to perform well in three countries across two continents is a major
step forward. Social calls have received even less attention and are also particularly difficult
to detect automatically, because this category basically includes all vocalisations not used to
detect the surroundings. With our current model in BatSpot, it should be possible to reliably
detect longer social calls, such as the trills produced by species we have evaluated in the
Pipistrellus genus. The data presented with the paper will also serve as an important
contribution to a future comprehensive dataset with recordings from across Europe or wider.
We envision that the social call detector, once further developed, will be able to support
passive acoustic monitoring purposes by enabling automated analysis and localisation of
important roosting and mating areas.
For all three models, there are several steps to keep in mind when applying them to large
datasets. First, a validation set is necessary to quantify the recall and precision for each
location and season included in the dataset to be analysed. This does not need to be a
specific percentage of all annotated data, but should rather ensure that all variation in the full
dataset is covered and performance on it can be quantified. Validating performance is not
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preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in
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specific to BatSpot, as even established commercial software might show a strong bias
when applied in, for example, the offshore environment (Smeele et al. 2026). If retraining is
needed, this can be done by following the steps in the README (see below). This should
drastically reduce training time compared to training with all the published data plus the data
collected in the specific project (Verwimp et al., 2025). And finally, once the model performs
satisfactorily, all raw data can be processed using the scripts supplied to index and process
large datasets on a high performance computing cluster, or a local computer with GPU.
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