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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Anim. Sci.</journal-id>
<journal-title>Frontiers in Animal Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Anim. Sci.</abbrev-journal-title>
<issn pub-type="epub">2673-6225</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fanim.2023.1083272</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Animal Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Classification of behaviors of free-ranging cattle using accelerometry signatures collected by virtual fence collars</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Versluijs</surname>
<given-names>Erik</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1351093"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Niccolai</surname>
<given-names>Laura J.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2072863"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Spedener</surname>
<given-names>M&#xe9;lanie</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zimmermann</surname>
<given-names>Barbara</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/760705"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hessle</surname>
<given-names>Anna</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1602070"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Tofastrud</surname>
<given-names>Morten</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Devineau</surname>
<given-names>Olivier</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1498714"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Evans</surname>
<given-names>Alina L.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/623225"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Inland Norway University of Applied Sciences, Department of Forestry and Wildlife Management</institution>, <addr-line>Campus Evenstad, Koppang</addr-line>, <country>Norway</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Swedish University of Agricultural Sciences, Department of Animal Environment and Health</institution>, <addr-line>Skara</addr-line>, <country>Sweden</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Inland Norway University of Applied Sciences, Department of Agricultural Sciences</institution>, <addr-line>Campus Bl&#xe6;stad, Hamar</addr-line>, <country>Norway</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Carolina Pugliese, University of Florence, Italy</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Martin Komainda, Institute of Grassland Science, University of G&#xf6;ttingen, Germany; Gamaliel Simanungkalit, University of New England, Australia; Said Benaissa, Ghent University, Belgium</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Erik Versluijs, <email xlink:href="mailto:erik.versluijs@inn.no">erik.versluijs@inn.no</email>
</p>
</fn>
<fn fn-type="equal" id="fn003">
<p>&#x2020;These authors have contributed equally to this work and share first authorship</p>
</fn>
<fn fn-type="other" id="fn002">
<p>This article was submitted to Precision Livestock Farming, a section of the journal Frontiers in Animal Science</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>14</day>
<month>04</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>4</volume>
<elocation-id>1083272</elocation-id>
<history>
<date date-type="received">
<day>28</day>
<month>10</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>27</day>
<month>03</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Versluijs, Niccolai, Spedener, Zimmermann, Hessle, Tofastrud, Devineau and Evans</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Versluijs, Niccolai, Spedener, Zimmermann, Hessle, Tofastrud, Devineau and Evans</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Precision farming technology, including GPS collars with biologging, has revolutionized remote livestock monitoring in extensive grazing systems. High resolution accelerometry can be used to infer the behavior of an animal. Previous behavioral classification studies using accelerometer data have focused on a few key behaviors and were mostly conducted in controlled situations. Here, we conducted behavioral observations of 38 beef cows (Hereford, Limousine, Charolais, Simmental/NRF/Hereford mix) free-ranging in rugged, forested areas, and fitted with a commercially available virtual fence collar (Nofence) containing a 10Hz tri-axial accelerometer. We used random forest models to calibrate data from the accelerometers on both commonly documented (e.g., feeding, resting, walking) and rarer (e.g., scratching, head butting, self-grooming) behaviors. Our goal was to assess pre-processing decisions including different running mean intervals (smoothing window of 1, 5, or 20 seconds), collar orientation and feature selection (orientation-dependent versus orientation-independent features). We identified the 10 most common behaviors exhibited by the cows. Models based only on orientation-independent features did not perform better than models based on orientation-dependent features, despite variation in how collars were attached (direction and tightness). Using a 20 seconds running mean and orientation-dependent features resulted in the highest model performance (model accuracy: 0.998, precision: 0.991, and recall: 0.989). We also used this model to add 11 rarer behaviors (each&lt; 0.1% of the data; e.g. head butting, throwing head, self-grooming). These rarer behaviors were predicted with less accuracy because they were not observed at all for some individuals, but overall model performance remained high (accuracy, precision, recall &gt;98%). Our study suggests that the accelerometers in the Nofence collars are suitable to identify the most common behaviors of free-ranging cattle. The results of this study could be used in future research for understanding cattle habitat selection in rugged forest ranges, herd dynamics, or responses to stressors such as carnivores, as well as to improve cattle management and welfare.</p>
</abstract>
<kwd-group>
<kwd>free-ranging cattle</kwd>
<kwd>behavioral classification</kwd>
<kwd>animal behavior</kwd>
<kwd>accelerometry</kwd>
<kwd>virtual fence collars</kwd>
</kwd-group>
<contract-num rid="cn001">302674</contract-num>
<contract-sponsor id="cn001">Norges Forskningsr&#xe5;d<named-content content-type="fundref-id">10.13039/501100005416</named-content>
</contract-sponsor>
<counts>
<fig-count count="4"/>
<table-count count="5"/>
<equation-count count="8"/>
<ref-count count="75"/>
<page-count count="13"/>
<word-count count="7327"/>
</counts>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Livestock grazing, whether in intensive (feed lots and pastures) or extensive (free-range) systems is a traditional practice that has persisted in our modern society to cope with increasing food production demands (<xref ref-type="bibr" rid="B40">Michalk et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B32">Komarek et&#xa0;al., 2021</xref>). Within this tradition, the ever-expanding development of modern technology has allowed for the growth of precision livestock farming management and research (<xref ref-type="bibr" rid="B19">Eastwood et&#xa0;al., 2017</xref>). Indeed, this management approach focuses on the fine-scale monitoring of individuals&#x2019; health and food intake (<xref ref-type="bibr" rid="B56">Schellberg et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B72">Werkheiser, 2018</xref>). While it remains relatively easy to implement in barn and pasture settings, free-range farming presents additional challenges; herd supervision can become more difficult as cattle are not contained. Additionally, external factors affecting cattle such as exposure to climatic extremes, parasitic load, untreated diseases, accidents, and potential carnivore effects are complex to monitor (<xref ref-type="bibr" rid="B28">Hutchings et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B60">Silanikove, 2000</xref>; <xref ref-type="bibr" rid="B57">Sevi et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B43">Nedeva, 2020</xref>). Technological advances such as biosensors, camera-equipped drones and GPS collars have offered scientists, managers and farmers tools to address those challenges (<xref ref-type="bibr" rid="B25">Herlin et&#xa0;al., 2021</xref>).</p>
<p>Notably, modern GPS collars, which often contain accelerometry sensors, provide researchers with an opportunity for non-invasive, low maintenance and remote monitoring of livestock and wildlife. If properly calibrated, this allows for the study of fine-scale animal behavior, activity budgets and energy expenditure on an individual level (<xref ref-type="bibr" rid="B68">V&#xe1;zquez Diosdado et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B46">O&#x2019;Leary et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B41">Mulvenna et&#xa0;al., 2022</xref>). Even though accelerometry data has largely increased our understanding and knowledge of livestock behavior (<xref ref-type="bibr" rid="B64">Theurer et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B67">Uenishi et&#xa0;al., 2021</xref>), most behavioral studies using accelerometry sensors focus on a few key behaviors such as grazing, resting, and walking (<xref ref-type="bibr" rid="B52">Rob&#xe9;rt et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B26">Homburger et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B5">Benaissa et&#xa0;al., 2019</xref>), and they are conducted in controlled settings such as barns and pastures (<xref ref-type="bibr" rid="B26">Homburger et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B24">Hendriks et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B53">Rodriguez-Baena et&#xa0;al., 2020</xref>).</p>
<p>Very few studies examine the behavior of free-ranging cattle in remote areas, where cattle are difficult to monitor and observe due to them moving over large areas and using dense vegetation types (<xref ref-type="bibr" rid="B65">Tofastrud et&#xa0;al., 2019</xref>). Yet, in these conditions, behaviors such as vigilance, social behaviors or grooming can be representative of stress, as cows have been reported to increase vigilance when stressed (<xref ref-type="bibr" rid="B71">Welp et&#xa0;al., 2004</xref>), and to decrease milk production when separated from the herd and not allowed to perform social behaviors (<xref ref-type="bibr" rid="B23">Hedlund and L&#xf8;vlie, 2015</xref>). In addition, self and allogrooming are reported to be frequent as maintenance behaviors and shown to be an important proxy for welfare (<xref ref-type="bibr" rid="B31">Kohari et&#xa0;al., 2007</xref>). While these behaviors are important to monitor, they are especially difficult to observe. <xref ref-type="bibr" rid="B66">Tofastrud et&#xa0;al. (2018)</xref> used two-axial accelerometry data at five-minute intervals on free-ranging cattle to study resting, grazing and movement activity patterns. Although this allows for insight into general free-ranging cattle behavioral habits in remote areas, it lacks the ability to precisely quantify additional, rarer behaviors and restricts the amount of information capable of being calibrated and further studied. Continuous, high resolution (10Hz) tri-axial accelerometry data can potentially increase the number of behaviors that can be classified and provide more detailed information about cattle behavior in large rugged, forested ranges (<xref ref-type="bibr" rid="B27">Hounslow et&#xa0;al., 2019</xref>). Such studies could contribute to the improvement of farmer monitoring systems related to high precision farming in outfields and offer the possibility to study effects from external factors, such as carnivores, on cattle behavior while monitoring welfare.</p>
<p>There are numerous ways to analyze accelerometry data, ranging from simple decision trees to complex neural networks (<xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al., 2022</xref>). Most studies utilize supervised machine learning methods such as random forest (<xref ref-type="bibr" rid="B17">de Weerd et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B75">Williams et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al., 2022</xref>), while others use unsupervised machine learning such as hidden Markov Models (<xref ref-type="bibr" rid="B36">Leos-Barajas et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B14">Chimienti et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B48">Rautiainen et&#xa0;al., 2022</xref>). Supervised methods provide the advantage of allowing for accelerometry data to be calibrated on actual behavioral observations, which then allows for prediction of behaviors based on collected data.</p>
<p>Feature selection is the first step in this process. Features can be calculated and extracted from raw accelerometry data and be used for behavioral classification. Orientation dependent features, such as the mean and variance of the raw x, y, z values, body pitch roll, yaw and dynamic acceleration (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>) remain sensitive to the sensor&#x2019;s orientation (<xref ref-type="bibr" rid="B1">Abell et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B5">Benaissa et&#xa0;al., 2019</xref>). For example, even though collars are assumed to be stationary positioned on the animal, there can be noise related to rotation, collar deployment errors, and other causes. This may result in additional variability in orientation-dependent features, which may render these features unusable without correcting for orientation (<xref ref-type="bibr" rid="B74">Williams et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B4">Barker et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B30">Kamminga et&#xa0;al., 2018</xref>). This sensitivity can be challenging when standardizing accelerometer sensor placement during animal handling, and recapturing individuals to manually fix issues is difficult (<xref ref-type="bibr" rid="B11">Chakravarty et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B10">Cade et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B48">Rautiainen et&#xa0;al., 2022</xref>). To remedy this problem, orientation independent features can be utilized. Models can then account for displacement of the sensor&#x2019;s orientation and make obsolete the need to correct for orientation.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Position of the GPS collar below the cows&#x2019; neck. Black lines indicate the direction of the three axes (X, Y, Z) and arrows indicate on which axis the roll and pitch were calculated. Art by Saskia H. Wulff.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fanim-04-1083272-g001.tif"/>
</fig>
<p>High resolution (&gt; 1Hz) accelerometry data is usually smoothed with a running average over a given time window (<xref ref-type="bibr" rid="B58">Shepard et&#xa0;al., 2008a</xref>). The chosen window length can result in the loss of certain less frequent and shorter behaviors that might not be detectable, while longer lasting, more common behaviors increase the accuracy and precision of the predictions (<xref ref-type="bibr" rid="B39">Mansbridge et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B12">Chang et&#xa0;al., 2022</xref>). Therefore, clear study goals become essential in accelerometry analysis, as the study aim will determine different trade-offs and decisions for modeling (<xref ref-type="bibr" rid="B12">Chang et&#xa0;al., 2022</xref>).</p>
<p>For instance, some authors suggest that averaging values in the sensor provides an opportunity to increase data collection capacity, and thus allows for live monitoring of behaviors with computationally simple and cost-efficient features and algorithms, which reduce battery usage (<xref ref-type="bibr" rid="B29">Kamminga, 2020</xref>; <xref ref-type="bibr" rid="B45">Nuijten et&#xa0;al., 2020</xref>). Study questions will thus determine data collection methods, which resolution the data needs to be collected in, as well as which steps should be included in the pre-processing. This will induce trade-offs which will determine what type and number of behaviors that can be analyzed (<xref ref-type="bibr" rid="B30">Kamminga et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al., 2022</xref>).</p>
<p>In this paper, we aim to calibrate high-resolution accelerometry data collected by commercially available livestock collars deployed on free-ranging cattle in remote areas. Additionally, we aim to investigate the effect of collar placement during deployment of collar and how it might affect accelerometry data. Finally, we attempt to address the gap in knowledge concerning the classification of less frequent cattle behaviors to accelerometry data, as these behaviors can represent behavioral changes or even be indicators of stress.</p>
<p>Our classification study contributes to potential research on cattle social interactions, behavioral responses to carnivores, and energy expenditure of free-ranging cattle in remote forested areas. Additionally, this study has the potential to develop tools for improved monitoring systems for farmers, and therefore to contribute to the practice of agroforestry and precision livestock farming.</p>
<p>We first hypothesized that the model performance to predict general behaviors such as walking (locomotion), foraging, vigilance, standing, laying/resting, or ruminating, would be affected by the choice of features included in the models (orientation-dependent versus orientation-independent features) and by the direction of the collar position on the neck of the animals (as we wanted to account for collar placement variation by the farmers on cattle) (H1). Models with orientation-independent features might perform better than those using orientation-dependent features, as orientation-independent features can account for potential rotation or tightness differences of collars when individuals navigate through rugged terrain. Additionally, models including orientation-dependent features corrected for direction might perform better than those without correction.</p>
<p>Secondly, we hypothesized that less frequent behaviors such as social interactions and body care movements would be impacted by the smoothing of the data (H2). Data smoothed with a short running mean would allow for the detection of rarer behaviors, but at the cost of a loss in the overall model&#x2019;s performance as a short running window mean can induce more noise during the analysis.</p>
</sec>
<sec id="s2">
<label>2</label>
<title>Methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Study area</title>
<p>We collected data in three summer grazing ranges in the Innlandet county of Norway. This region of Norway belongs to the boreal forest biome and is dominated by coniferous forest, mires and lakes, and only about 4% is covered by agricultural fields. Many beef cattle breeders release their suckler cows with their calves into the forest during the summer months, to make use of outfield grazing resources and to spare the fields close to the farm for winter forage production. The summer ranges included in this study (Steinvik 27.2 km<sup>2</sup>, long = 11.28&#xb0;, lat = 61.23&#xb0;, Deset west 16.4 km<sup>2</sup>, long = 11.42&#xb0;, lat = 61.29&#xb0;, T&#xf8;rberget 5.8 km<sup>2</sup>, long = 12.29&#xb0;, lat = 61.08&#xb0;) consisted of a patchwork of forest stands of different age classes due to clearcutting practices, often followed by soil scarification, thinning and other silvicultural practices used to increase timber production. Forest stands were either dominated by Norway spruce (<italic>Picea abies</italic>) or Scots pine (<italic>Pinus silvestris</italic>), interspersed with birch (<italic>Betula pendula</italic>, <italic>B. pubescens</italic>) and other deciduous species. Only minor parts of the ranges were covered by bogs and old grazing meadows. The terrain was rugged and covered an elevational gradient of 300 &#x2013; 640&#xa0;m above sea level. A network of forest roads connected the forest stands. Earlier studies in similar habitat have shown that cattle prefer to graze in young forest stands (<xref ref-type="bibr" rid="B65">Tofastrud et&#xa0;al., 2019</xref>), where there is access to graminoids of different species (<xref ref-type="bibr" rid="B63">Spedener et&#xa0;al., 2019</xref>). However, clearcutting and soil scarification induced varied landscapes that force wildlife and free-ranging livestock to walk on uneven, rugged terrain with obstacles such as fallen trees, stumps, and tree residuals after logging.</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Study animals</title>
<p>All suckler cows belonging to four farms were fitted with virtual fence collars (<xref ref-type="bibr" rid="B44">Nofence, 2022</xref>) in May 2021. The farmers trained the cows for virtual fencing while still at the farm, following the instructions given by Nofence (<xref ref-type="bibr" rid="B44">Nofence, 2022</xref>). In end of May and beginning of June, the cows were released into their summer grazing ranges (45 cows Steinvik, 21 cows in Deset West, and 13 cows in T&#xf8;rberget), along with their (uncollared) calves. The grazing ranges were delimited by virtual fencing, and range size did not change much during the summer season. In this study, we included data from 38 cows (4 &#x2013; 16 individuals per farm) of the following breeds: Hereford (n = 16), Limousine (n = 5), Charolais (n = 4), and 13 crossbred individuals including the beef breeds Simmental, Hereford and the dual-purpose breed Norwegian Red (NRF) (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;1</bold>
</xref>).</p>
<p>Nofence collars with virtual fencing technology (<xref ref-type="bibr" rid="B9">Brunberg et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B72">Werkheiser, 2018</xref>; <xref ref-type="bibr" rid="B55">S&#xf8;raa and Vik, 2021</xref>; <xref ref-type="bibr" rid="B69">Verdon et&#xa0;al., 2021</xref>) triangulate the positions of animals (1 position every 5 to 15 minutes) through the GNSS (Global Navigation Satellite Systems), as well as record movement activity with a motion sensor that yields high-resolution tri-axial accelerometry data (10Hz). The battery is designed to last for at least three months and to be continuously recharged through solar panels. Each animal carried a total weight of 1446g. As cows weighted between 500 and 900 kg depending on the breed and age, collars made up about 0.3-0.5% of the body weight. These devices fell under the recommended threshold of 3-5% of an animal&#x2019;s body mass (<xref ref-type="bibr" rid="B3">Arnemo et al., 2011</xref>; <xref ref-type="bibr" rid="B62">Soulsbury et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B22">Hamidi et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B61">Sonne et&#xa0;al., 2022</xref>).</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Accelerometry sensor activation</title>
<p>Accelerometry sensors on the Nofence collars were remotely activated to continuously sample and transmit data during bouts of 48h. Bouts were distributed throughout the grazing season at intervals of minimum three weeks between bouts per cow, to enable the solar-powered batteries to recharge. This resulted in 1-5 sampling bouts per monitored cow. The order of activation followed a somewhat opportunistic design, depending on where the cows were in relation to each other and in relation to the habitat type. Cows in dense forest were difficult to observe, and we therefore activated mostly collars of cows in more open habitat types.</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Video data</title>
<p>A team of two people located and filmed adult individual cattle in-field while the collars&#x2019; accelerometers were activated (<xref ref-type="bibr" rid="B2">Arablouei et&#xa0;al., 2021</xref>). To ensure a maximum number of behaviors was captured, we filmed the cattle throughout the summer, in varying weather conditions, times of day, and terrains. The team located the cattle with the Nofence app (<xref ref-type="bibr" rid="B44">Nofence, 2022</xref>), which displays the latest positions of individuals.</p>
<p>Carrying a video camera (Canon XA40, Canon Inc.), the team approached the herds as quietly as possible (min. 10 - 15&#xa0;m distance) to minimize disturbance and stress. When the habitat was open (e.g., clearcuts), the team filmed the entire herd, but when the vegetation was dense, the team focused on filming single individuals. Individual cows were identified by their earmarks and color patterns, using direct observation, binoculars or camera zoom. Video clips lasted between 24 seconds to 48 minutes and were downloaded from the internal memory card every evening, to be stored on a One Drive folder for later use.</p>
<p>All video footage was then viewed and tagged in the software BORIS (<xref ref-type="bibr" rid="B20">Friard and Gamba, 2016</xref>), an open-source platform for behavioral coding of video/audio files, to identify and label individual cattle behavior (done multiple times for the same video if multiple individuals in the footage). This was done using an ethogram with a total of 42 behaviors and postures in the following categories: alert, body care, excretion, intake, locomotion, posture, posture transition and social interaction. This ethogram was constructed to be as inclusive as possible of all cattle behaviors (<xref ref-type="bibr" rid="B34">Langford et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B38">MacKay et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B47">Petherick et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B66">Tofastrud et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B42">Navarro et&#xa0;al., 2019</xref>) and was further detailed with field observations (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;2</bold>
</xref>). The principal investigator of the fieldwork and video analysis labeled ~95% of the video material and trained and supervised three students for the remaining videos, thus minimizing observer bias.</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Pre-processing of the accelerometry data</title>
<p>The software R, version 4.2.1 (<xref ref-type="bibr" rid="B49">R Core Team, 2022</xref>) with the Rstudio interface (<xref ref-type="bibr" rid="B54">R Studio Team, 2021</xref>) was used for data pre-processing, analysis, and visualization. The tri-axial accelerometry data was matched with the corresponding video data. First, the approximate time stamps of the tri-axial accelerometry data were calculated in decimal seconds. Raw data files have rounded timestamps, up to 32 observations per timestamp, while the data was collected at 10Hz. We converted this to unique timestamps with decimal seconds by subtracting the amount of time that had passed since the last rounded timestamp (i.e., when the time stamp was 13:02:02, and it was the 5th observation, we subtracted 2.7 seconds ((32-5)/10) from the original time stamp). Double observations were then removed, resulting in a data set of approximately 10Hz with an error of &#xb1;0.5 seconds. Due to this rounding error, possible time drift in video recordings, and to reduce observer bias in video analysis, we excluded any behavior which had a shorter length than five seconds.</p>
<p>In the next step, the behavioral data was matched with the accelerometry data by timestamp using the function &#x2018;<italic>foverlaps</italic>&#x2019; from the <italic>Data.table</italic> package (<xref ref-type="bibr" rid="B18">Dowle and Srinivasan, 2019</xref>). For the initial match, all behaviors were used (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;2</bold>
</xref>). However, as some behaviors overlapped with each other and we could only keep one single behavior per observation for the analyses, we introduced decision rules based on cattle ecology and body movement (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure&#xa0;1</bold>
</xref>, <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;3</bold>
</xref>). For example, grazing behavior was prioritized over walking, and vigilant behavior was considered only when no other behaviors were shown, to avoid noise in the accelerometry signature from the other behaviors. Behaviors that made up less than 1% of all observation time were pooled into one class named &#x2018;other&#x2019;.</p>
</sec>
<sec id="s2_6">
<label>2.6</label>
<title>Features calculation</title>
<p>We used the raw accelerometry data to calculate orientation dependent and independent features to avoid additional complex pre-processing steps and to reduce the need for computing power. The mean and the standard deviation were calculated along the three axes (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>) using 1, 5, and 20 seconds running means, following previous work with accelerometry data on cattle (<xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al., 2022</xref>). The pitch is the angle of the collar in degrees (&#xb0;) along the x axis (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>, Equation 1) where the collar is facing upwards for a +90&#xb0; angle and downwards for a -90&#xb0; angle. Similarly, the roll is the angle of the collar along the y axis (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>, Equation 2) (<xref ref-type="bibr" rid="B59">Shepard et&#xa0;al., 2008b</xref>; <xref ref-type="bibr" rid="B15">Chimienti et&#xa0;al., 2016</xref>). Thereafter, we calculated the mean and standard deviation of the pitch and roll across the three-running means (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Overview of the features used in the models: orientation dependent (three axis, body pitch, and body) and orientation independent (overall dynamic body acceleration (ODBA), vector of dynamic body acceleration (VEDBA), and magnitude of acceleration (AMAG)).</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Orientation</th>
<th valign="middle" align="center">Features</th>
<th valign="middle" align="center">Definition</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="6" align="left">Dependent</td>
<td valign="middle" align="left">Mean X, Y, Z</td>
<td valign="middle" align="left">Mean acceleration recorded from the accelerometer (g) across the running mean for all three axis (X,Y,Z)</td>
</tr>
<tr>
<td valign="middle" align="left">Standard deviation X, Y, Z</td>
<td valign="middle" align="left">Standard deviation of the acceleration recorded from the accelerometer (g) across the running mean for all three axis (X,Y,Z)</td>
</tr>
<tr>
<td valign="middle" align="left">Mean body pitch</td>
<td valign="middle" align="left">Mean angle of the collar along the x axis (degrees) across the running mean.</td>
</tr>
<tr>
<td valign="middle" align="left">Standard deviation body pitch</td>
<td valign="middle" align="left">Standard deviation of the angle of the collar along the x axis (degrees) across the running mean</td>
</tr>
<tr>
<td valign="middle" align="left">Mean body roll</td>
<td valign="middle" align="left">Mean angle of the collar along the y axis (degrees) across the running mean.</td>
</tr>
<tr>
<td valign="middle" align="left">Standard deviation body roll</td>
<td valign="middle" align="left">Standard deviation of the angle of the collar along the y axis (degrees) across the running mean</td>
</tr>
<tr>
<td valign="middle" rowspan="6" align="left">Independent</td>
<td valign="middle" align="left">Mean ODBA</td>
<td valign="middle" align="left">Overall Dynamic Body Acceleration &#x2013; Measure of general effort across three axis averaged across the running mean</td>
</tr>
<tr>
<td valign="middle" align="left">Standard deviation ODBA</td>
<td valign="middle" align="left">Standard deviation of the ODBA across the running mean</td>
</tr>
<tr>
<td valign="middle" align="left">Mean VEDBA</td>
<td valign="middle" align="left">Vector of Dynamic Body Acceleration - Vector of the general effort across three axis averaged across the running mean</td>
</tr>
<tr>
<td valign="middle" align="left">Standard deviation VEDBA</td>
<td valign="middle" align="left">Standard deviation of the VEDBA across the running mean</td>
</tr>
<tr>
<td valign="middle" align="left">Mean AMAG</td>
<td valign="middle" align="left">Magnitude of acceleration - Measure of the magnitude of effort across three axis averaged across the running mean</td>
</tr>
<tr>
<td valign="middle" align="left">Standard deviation AMAG</td>
<td valign="middle" align="left">Standard deviation of the AMAG across the running mean</td>
</tr>
</tbody>
</table>
</table-wrap>
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<p>For the overall dynamic body acceleration (ODBA) and the vector of dynamic body acceleration (VEDBA) we first subtracted the mean acceleration from the raw acceleration for each axis to calculate the dynamic acceleration (respectively noted <italic>dx, dy, dz for each axis</italic>). The sum of the absolute values from the dynamic acceleration was then used to calculate the ODBA (Equation 3), and subsequently the mean and standard deviation of ODBA across the running means (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). The VEDBA was calculated by taking the square root of the squared dynamic acceleration for the three axes (Equation 4), again followed by calculation of mean and standard deviation across the running mean (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). Finally, the magnitude of acceleration (AMAG) was calculated by taking the square root of the squared acceleration for the three axes (Equation 5), with mean and standard deviation across the running mean (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>).</p>
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<label>(5)</label>
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<p>During the analyses, we noticed that about half of the collars were placed in the reverse direction on the cow. We identified these based on the pitch angle from &#x201c;foraging_low&#x201d; behavior (a positive angle suggests a reversed collar), and by looking at the video data (the collars have a small mark to distinguish between left and right side). The data was then corrected by reversing both x and y axis. The orientation-dependent features were calculated for both the original and the corrected data sets (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Flow chart showing the modeling process. For each feature category (orientation dependent, orientation dependent corrected, orientation independent), we ran three models, one for each running mean (1, 5, and 20 seconds). Furthermore, after assessing the models performance we reran the best model including more behaviors.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fanim-04-1083272-g002.tif"/>
</fig>
</sec>
<sec id="s2_7">
<label>2.7</label>
<title>Random forest models</title>
<p>We prepared data for orientation-dependent, orientation-dependent with correction, and orientation-independent features (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Within each category, we ran three models on the three different running means (1, 5, and 20 seconds). All features were standardized before running the models. The data were grouped by behavior and split with 80% used for training and 20% for validation. Thereafter, random forest models were run using the function &#x2018;<italic>h2o.randomforest&#x2019;</italic> from the <italic>H2O</italic> package version 3.36.1.2 (<xref ref-type="bibr" rid="B35">LeDell et&#xa0;al., 2021</xref>). We chose the random forest algorithm for its versatility, and because it has good predictive power for its computation time (<xref ref-type="bibr" rid="B6">Biau &amp; Scornet, 2016</xref>). Models were run with 150 trees, as with this number of trees, the log-loss of the model became stable. We used 5-fold cross-validation, and we added a weight to each class (N of rarest behavior divided by the N of the behavioral class) to account for class imbalance and potential over-fitting (<xref ref-type="bibr" rid="B16">Cutler et&#xa0;al., 2012</xref>). Additionally, we checked for individual variation by running an individual-based cross-validation (without data split by behavior). The package&#x2019;s default settings were used for the other hyperparameters (max tree depth = 20, mtries = square root of the number of features). We visualized model results using the <italic>DALEX</italic> package (<xref ref-type="bibr" rid="B7">Biecek, 2018</xref>) and the <italic>ggplot2</italic> package (<xref ref-type="bibr" rid="B73">Wickham, 2016</xref>). Furthermore, the model accuracy, precision, and recall were calculated to compare model performance (Equations 6, 7, and 8, respectively) (<xref ref-type="bibr" rid="B29">Kamminga, 2020</xref>). Based on the best performing model, we ran the last model including more behaviors to test if models including more behaviors performed similarly well (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>). Here, the threshold for a behavior to be included was minimum 0.1% of the total observation time.</p>
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</mml:mrow>
</mml:mfrac>
</mml:mrow>
</mml:math>
</disp-formula>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<p>We annotated 6 898 behavioral observations based on video analysis and matched them with accelerometry data, resulting in a total of 1 240 588 observations for 31 behaviors (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;2</bold>
</xref>). Visual inspection showed appropriate matching, i.e. the accelerometry data showed different signatures for different behaviors (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Example of four behaviors and their raw accelerometry signature for the duration of 30 seconds (300 observations). For the behaviors; <bold>(A)</bold> Walking, <bold>(B)</bold> Ruminating laying, <bold>(C)</bold> Vigilance, and <bold>(D)</bold> Foraging low. Line color indicate the different axis; blue: z axis, red: <italic>x</italic> axis, green: <italic>y</italic> axis. The proportion of observations for each of these behaviors is 9%, 23%, 8%, 33%, respectively. See the full overview of behaviors in <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;3</bold>
</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fanim-04-1083272-g003.tif"/>
</fig>
<p>Overall, the models with 20 seconds running mean performed better than those with shorter running means (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Orientation-independent models showed a lower performance than orientation-dependent models. Corrected orientation-dependent models performed similar to uncorrected models (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Both orientation-dependent models with 20 seconds running mean (models C and F) had average accuracy, precision and recall &gt; 0.96. The orientation-independent model I had an average precision of 0.88, recall 0.93 and accuracy 0.99 (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Precision averaged 0.62 and 0.63 for the 1 second running mean in the orientation-dependent and corrected orientation-dependent models, respectively (models A and D, <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>), and averaged 0.37 and 0.64 for the 1 second and the 5 seconds running means, respectively in the models with orientation-independent features (models G and H, <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Similarly, the recall averaged 0.69 for the 1 second running mean in the orientation-dependent and corrected orientation-dependent models (models A and D, <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). The orientation-independent models G and H had an average recall of 0.39 and 0.72 for 1 and 5 seconds running mean, respectively (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>). Prediction performance varied across behaviors for recall, accuracy and precision (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>, and <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figures&#xa0;2</bold>
</xref>, <xref ref-type="supplementary-material" rid="SM1">
<bold>3</bold>
</xref>, respectively). Behaviors in models with a shorter running mean varied more in recall than in models with a longer running mean, e.g., &#x2018;foraging_high&#x2019; and &#x2018;other&#x2019; in the orientation-dependent models A and D were below 0.4 (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). Those behaviors were correctly classified in less than 40% of their occurrence. Orientation-dependent models using 5 seconds running means performed better with lowest recall of 0.8 (model B, <xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). For models with 20 seconds running mean, the recall was never below 0.96, showing that the lowest performing behavior had a classification success of 0.96. The performance of the orientation-independent models showed overall lower recall values for each behavior. Still, increased running means in the orientation-independent models helped to improve prediction and decreased the number of classification errors.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Overview of the nine models with the average and range for accuracy (as the ratio of correct predictions out of all predictions), precision (as the ratio of true positives over the sum of false positives and true negatives), and recall (as the ratio of correct predicted outcomes to all predictions).</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Model</th>
<th valign="bottom" align="center">Accuracy</th>
<th valign="bottom" align="center">Precision</th>
<th valign="bottom" align="center">Recall</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">(A) Orientation dependent (1 sec)</td>
<td valign="bottom" align="center">0.954 (0.907-0.989)</td>
<td valign="bottom" align="center">0.617 (0.325-0.894)</td>
<td valign="bottom" align="center">0.687 (0.473-0.842)</td>
</tr>
<tr>
<td valign="top" align="left">(B) Orientation dependent (5 sec)</td>
<td valign="bottom" align="center">0.985 (0.961-0.997)</td>
<td valign="bottom" align="center">0.854 (0.645-0.972)</td>
<td valign="bottom" align="center">0.923 (0.839-0.982)</td>
</tr>
<tr>
<td valign="top" align="left">(C) Orientation dependent (20 sec)</td>
<td valign="bottom" align="center">0.997 (0.991-0.999)</td>
<td valign="bottom" align="center">0.961 (0.875-0. 995)</td>
<td valign="bottom" align="center">0.985 (0.965-0.999)</td>
</tr>
<tr>
<td valign="top" align="left">(D) Orientation dependent corrected (1 sec)</td>
<td valign="bottom" align="center">0.957 (0.908-0.990)</td>
<td valign="bottom" align="center">0.629 (0.337-0.898)</td>
<td valign="bottom" align="center">0.693 (0.480-0.848)</td>
</tr>
<tr>
<td valign="top" align="left">(E) Orientation dependent corrected (5 sec)</td>
<td valign="bottom" align="center">0.986 (0.959-0.998)</td>
<td valign="bottom" align="center">0.868 (0.648-0.975)</td>
<td valign="bottom" align="center">0.922 (0.826-0.978)</td>
</tr>
<tr>
<td valign="top" align="left">(F) Orientation dependent corrected (20 sec)</td>
<td valign="bottom" align="center">0.997 (0.990-1.000)</td>
<td valign="bottom" align="center">0.964 (0.872-0.995)</td>
<td valign="bottom" align="center">0.985 (0.953-0.999)</td>
</tr>
<tr>
<td valign="top" align="left">(G) Orientation independent (1 sec)</td>
<td valign="bottom" align="center">0.918 (0.834-0.974)</td>
<td valign="bottom" align="center">0.371 (0.108-0.755)</td>
<td valign="bottom" align="center">0.392 (0.080-0.741)</td>
</tr>
<tr>
<td valign="top" align="left">(H) Orientation independent (5 sec)</td>
<td valign="bottom" align="center">0.956 (0.901-0.985)</td>
<td valign="bottom" align="center">0.642 (0.437-0.881)</td>
<td valign="bottom" align="center">0.721 (0.498-0.849)</td>
</tr>
<tr>
<td valign="top" align="left">(I) Orientation independent (20 sec)</td>
<td valign="bottom" align="center">0.988 (0.970-0.997)</td>
<td valign="bottom" align="center">0.884 (0.798-0.973)</td>
<td valign="bottom" align="center">0.931 (0.891-0.981)</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>The r<italic>ecall (classification success)</italic> for each model, specified by behavior and based on the model&#x2019;s validation data. Rows indicate orientation-dependent features, corrected orientation-dependent features, and orientation-independent features, respectively.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fanim-04-1083272-g004.tif"/>
</fig>
<p>The orientation-dependent model with 20 seconds running mean had a few occasions where behaviors were misclassified (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). However, the confusion matrix for the orientation-independent model with 20 seconds running mean showed a higher degree of misclassification (<xref ref-type="table" rid="T4">
<bold>Table&#xa0;4</bold>
</xref>), especially for behaviors that were more similar in their acceleration signature (e.g. &#x2018;foraging_low&#x2019; and &#x2018;walking&#x2019;, or &#x2018;ruminating_standing&#x2019; and &#x2018;ruminating_laying&#x2019;). The confusion matrices for all other models are presented in <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Tables&#xa0;4&#x2013;10</bold>
</xref>.</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Confusion matrix for the orientation dependent features without collar correction with 20 seconds running mean.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center"/>
<th valign="middle" colspan="11" align="center">Predicted behaviors</th>
</tr>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center"/>
<th valign="middle" align="center">allogrooming</th>
<th valign="middle" align="center">calf_suckle</th>
<th valign="middle" align="center">foraging_high</th>
<th valign="middle" align="center">foraging_low</th>
<th valign="middle" align="center">laying_resting</th>
<th valign="middle" align="center">other</th>
<th valign="middle" align="center">ruminating_laying</th>
<th valign="middle" align="center">ruminating_standing</th>
<th valign="middle" align="center">standing_resting</th>
<th valign="middle" align="center">vigilance</th>
<th valign="middle" align="center">walking</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="11" align="left">
<bold>Actual behaviors</bold>
</td>
<td valign="bottom" align="left">allogrooming</td>
<td valign="bottom" align="center">
<bold>3564</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">4</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">9</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">calf_suckle</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>4151</bold>
</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">4</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">foraging_high</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">
<bold>7444</bold>
</td>
<td valign="bottom" align="center">10</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">2</td>
</tr>
<tr>
<td valign="bottom" align="left">foraging_low</td>
<td valign="bottom" align="center">59</td>
<td valign="bottom" align="center">26</td>
<td valign="bottom" align="center">815</td>
<td valign="bottom" align="center">
<bold>80698</bold>
</td>
<td valign="bottom" align="center">20</td>
<td valign="bottom" align="center">92</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">26</td>
<td valign="bottom" align="center">297</td>
<td valign="bottom" align="center">74</td>
<td valign="bottom" align="center">364</td>
</tr>
<tr>
<td valign="bottom" align="left">laying_resting</td>
<td valign="bottom" align="center">14</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">8</td>
<td valign="bottom" align="center">
<bold>21485</bold>
</td>
<td valign="bottom" align="center">40</td>
<td valign="bottom" align="center">150</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">40</td>
<td valign="bottom" align="center">46</td>
<td valign="bottom" align="center">3</td>
</tr>
<tr>
<td valign="bottom" align="left">other</td>
<td valign="bottom" align="center">11</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">7</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">
<bold>6446</bold>
</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">11</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">7</td>
<td valign="bottom" align="center">14</td>
</tr>
<tr>
<td valign="bottom" align="left">ruminating_laying</td>
<td valign="bottom" align="center">30</td>
<td valign="bottom" align="center">17</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">67</td>
<td valign="bottom" align="center">156</td>
<td valign="bottom" align="center">
<bold>56607</bold>
</td>
<td valign="bottom" align="center">152</td>
<td valign="bottom" align="center">68</td>
<td valign="bottom" align="center">48</td>
<td valign="bottom" align="center">17</td>
</tr>
<tr>
<td valign="bottom" align="left">ruminating_standing</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">
<bold>5537</bold>
</td>
<td valign="bottom" align="center">4</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
</tr>
<tr>
<td valign="bottom" align="left">standing_resting</td>
<td valign="bottom" align="center">52</td>
<td valign="bottom" align="center">7</td>
<td valign="bottom" align="center">44</td>
<td valign="bottom" align="center">48</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">103</td>
<td valign="bottom" align="center">22</td>
<td valign="bottom" align="center">32</td>
<td valign="bottom" align="center">
<bold>17892</bold>
</td>
<td valign="bottom" align="center">31</td>
<td valign="bottom" align="center">61</td>
</tr>
<tr>
<td valign="bottom" align="left">vigilance</td>
<td valign="bottom" align="center">10</td>
<td valign="bottom" align="center">70</td>
<td valign="bottom" align="center">40</td>
<td valign="bottom" align="center">44</td>
<td valign="bottom" align="center">17</td>
<td valign="bottom" align="center">58</td>
<td valign="bottom" align="center">71</td>
<td valign="bottom" align="center">96</td>
<td valign="bottom" align="center">197</td>
<td valign="bottom" align="center">
<bold>18069</bold>
</td>
<td valign="bottom" align="center">45</td>
</tr>
<tr>
<td valign="bottom" align="left">walking</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">123</td>
<td valign="bottom" align="center">340</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">57</td>
<td valign="bottom" align="center">24</td>
<td valign="bottom" align="center">20</td>
<td valign="bottom" align="center">129</td>
<td valign="bottom" align="center">20</td>
<td valign="bottom" align="center">
<bold>21166</bold>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Columns represent the predicted behaviors; rows represent the actual observed behavior the validation data set.Bold values are the number of observations correctly predicted by the model.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>Confusion matrix for the orientation independent features without collar correction with 20 seconds running mean.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center"/>
<th valign="middle" colspan="11" align="center">Predicted behaviors</th>
</tr>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center"/>
<th valign="middle" align="center">allogrooming</th>
<th valign="middle" align="center">calf_suckle</th>
<th valign="middle" align="center">foraging_high</th>
<th valign="middle" align="center">foraging_low</th>
<th valign="middle" align="center">laying_resting</th>
<th valign="middle" align="center">other</th>
<th valign="middle" align="center">ruminating_laying</th>
<th valign="middle" align="center">ruminating_standing</th>
<th valign="middle" align="center">standing_resting</th>
<th valign="middle" align="center">vigilance</th>
<th valign="middle" align="center">walking</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="11" align="left">
<bold>Actual behaviors</bold>
</td>
<td valign="bottom" align="left">allogrooming</td>
<td valign="bottom" align="center">
<bold>3407</bold>
</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">38</td>
<td valign="bottom" align="center">7</td>
<td valign="bottom" align="center">14</td>
<td valign="bottom" align="center">18</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">44</td>
<td valign="bottom" align="center">4</td>
<td valign="bottom" align="center">35</td>
</tr>
<tr>
<td valign="bottom" align="left">calf_suckle</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">
<bold>4079</bold>
</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">8</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">25</td>
<td valign="bottom" align="center">14</td>
<td valign="bottom" align="center">14</td>
<td valign="bottom" align="center">9</td>
<td valign="bottom" align="center">3</td>
</tr>
<tr>
<td valign="bottom" align="left">foraging_high</td>
<td valign="bottom" align="center">30</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">
<bold>6721</bold>
</td>
<td valign="bottom" align="center">427</td>
<td valign="bottom" align="center">10</td>
<td valign="bottom" align="center">14</td>
<td valign="bottom" align="center">22</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">70</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">153</td>
</tr>
<tr>
<td valign="bottom" align="left">foraging_low</td>
<td valign="bottom" align="center">308</td>
<td valign="bottom" align="center">113</td>
<td valign="bottom" align="center">1120</td>
<td valign="bottom" align="center">
<bold>77405</bold>
</td>
<td valign="bottom" align="center">64</td>
<td valign="bottom" align="center">312</td>
<td valign="bottom" align="center">143</td>
<td valign="bottom" align="center">68</td>
<td valign="bottom" align="center">516</td>
<td valign="bottom" align="center">126</td>
<td valign="bottom" align="center">2317</td>
</tr>
<tr>
<td valign="bottom" align="left">laying_resting</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">67</td>
<td valign="bottom" align="center">36</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">
<bold>20589</bold>
</td>
<td valign="bottom" align="center">58</td>
<td valign="bottom" align="center">436</td>
<td valign="bottom" align="center">92</td>
<td valign="bottom" align="center">229</td>
<td valign="bottom" align="center">237</td>
<td valign="bottom" align="center">40</td>
</tr>
<tr>
<td valign="bottom" align="left">other</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">9</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">78</td>
<td valign="bottom" align="center">38</td>
<td valign="bottom" align="center">
<bold>6124</bold>
</td>
<td valign="bottom" align="center">47</td>
<td valign="bottom" align="center">31</td>
<td valign="bottom" align="center">88</td>
<td valign="bottom" align="center">10</td>
<td valign="bottom" align="center">72</td>
</tr>
<tr>
<td valign="bottom" align="left">ruminating_laying</td>
<td valign="bottom" align="center">69</td>
<td valign="bottom" align="center">267</td>
<td valign="bottom" align="center">23</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">493</td>
<td valign="bottom" align="center">262</td>
<td valign="bottom" align="center">
<bold>54773</bold>
</td>
<td valign="bottom" align="center">544</td>
<td valign="bottom" align="center">451</td>
<td valign="bottom" align="center">246</td>
<td valign="bottom" align="center">56</td>
</tr>
<tr>
<td valign="bottom" align="left">ruminating_standing</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">16</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">132</td>
<td valign="bottom" align="center">
<bold>5271</bold>
</td>
<td valign="bottom" align="center">65</td>
<td valign="bottom" align="center">9</td>
<td valign="bottom" align="center">15</td>
</tr>
<tr>
<td valign="bottom" align="left">standing_resting</td>
<td valign="bottom" align="center">133</td>
<td valign="bottom" align="center">107</td>
<td valign="bottom" align="center">129</td>
<td valign="bottom" align="center">301</td>
<td valign="bottom" align="center">145</td>
<td valign="bottom" align="center">191</td>
<td valign="bottom" align="center">293</td>
<td valign="bottom" align="center">130</td>
<td valign="bottom" align="center">
<bold>16389</bold>
</td>
<td valign="bottom" align="center">166</td>
<td valign="bottom" align="center">309</td>
</tr>
<tr>
<td valign="bottom" align="left">vigilance</td>
<td valign="bottom" align="center">38</td>
<td valign="bottom" align="center">137</td>
<td valign="bottom" align="center">115</td>
<td valign="bottom" align="center">79</td>
<td valign="bottom" align="center">203</td>
<td valign="bottom" align="center">93</td>
<td valign="bottom" align="center">305</td>
<td valign="bottom" align="center">199</td>
<td valign="bottom" align="center">557</td>
<td valign="bottom" align="center">
<bold>16809</bold>
</td>
<td valign="bottom" align="center">182</td>
</tr>
<tr>
<td valign="bottom" align="left">walking</td>
<td valign="bottom" align="center">77</td>
<td valign="bottom" align="center">50</td>
<td valign="bottom" align="center">242</td>
<td valign="bottom" align="center">1378</td>
<td valign="bottom" align="center">74</td>
<td valign="bottom" align="center">101</td>
<td valign="bottom" align="center">98</td>
<td valign="bottom" align="center">59</td>
<td valign="bottom" align="center">256</td>
<td valign="bottom" align="center">63</td>
<td valign="bottom" align="center">
<bold>19502</bold>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>Columns represent the predicted behaviors; rows represent the actual observed behavior the validation data set.Bold values are the number of observations correctly predicted by the model.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>The individual-based cross validation model using a 20 second running mean and orientation-dependent data indicated that rarer behaviors were more difficult to predict for individuals with less observations, but overall, the model performed similar to the model using the 80-20 data split, because there were enough observations of these rare behaviors across all individuals. The averaged accuracy was 0.997, the averaged precision was 0.965, and the averaged recall was 0.987 (See <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table&#xa0;11</bold>
</xref> for the confusion matrix).</p>
<p>The model including more behaviors had an average accuracy of 0.99, precision of 0.71, and recall of 0.96. The classification success for the behaviors that were also present in the other models performed similarly well (<xref ref-type="table" rid="T3">
<bold>Tables&#xa0;3</bold>
</xref>&#x2013;<xref ref-type="table" rid="T5">
<bold>5</bold>
</xref>). However, rarer behaviors had a high classification success with almost no misclassifications, e.g., in the behaviors &#x2018;stretching&#x2019; and &#x2018;throw_head&#x2019; (<xref ref-type="table" rid="T5">
<bold>Table&#xa0;5</bold>
</xref>).</p>
<table-wrap id="T5" position="float">
<label>Table&#xa0;5</label>
<caption>
<p>Confusion matrix for orientation dependent features with 20 seconds running mean including all behaviors below 0.1% proportionally of the data set.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center"/>
<th valign="middle" colspan="20" align="center">Predicted behaviors</th>
</tr>
<tr>
<th valign="middle" align="left"/>
<th valign="middle" align="center">
</th>
<th valign="middle" align="center">
allogrooming
</th>
<th valign="middle" align="center">
butting
</th>
<th valign="middle" align="center">
calf_suckle
</th>
<th valign="middle" align="center">
defecating
</th>
<th valign="middle" align="center">
foraging_high
</th>
<th valign="middle" align="center">
foraging_low
</th>
<th valign="middle" align="center">
getting_up
</th>
<th valign="middle" align="center">
laying_down
</th>
<th valign="middle" align="center">
laying_resting
</th>
<th valign="middle" align="center">
other
</th>
<th valign="middle" align="center">
rub_scratch
</th>
<th valign="middle" align="center">
ruminating_laying
</th>
<th valign="middle" align="center">
ruminating_standing
</th>
<th valign="middle" align="center">
self_grooming
</th>
<th valign="middle" align="center">
shaking
</th>
<th valign="middle" align="center">
standing_resting
</th>
<th valign="middle" align="center">
throw_head
</th>
<th valign="middle" align="center">
</th>
<th valign="middle" align="center">
vigilance
</th>
<th valign="middle" align="center">
walking
</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" rowspan="20" align="left">
<bold>Actual behaviors</bold>
</td>
<td valign="bottom" align="left">allogrooming</td>
<td valign="bottom" align="center">
<bold>3538</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">8</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">7</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">5</td>
</tr>
<tr>
<td valign="bottom" align="left">butting</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>806</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">calf_suckle</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>4147</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">defecating</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>262</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">foraging_high</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>7261</bold>
</td>
<td valign="bottom" align="center">88</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">10</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">21</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">11</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">16</td>
<td valign="bottom" align="center">24</td>
</tr>
<tr>
<td valign="bottom" align="left">foraging_low</td>
<td valign="bottom" align="center">370</td>
<td valign="bottom" align="center">288</td>
<td valign="bottom" align="center">179</td>
<td valign="bottom" align="center">29</td>
<td valign="bottom" align="center">3972</td>
<td valign="bottom" align="center">
<bold>72446</bold>
</td>
<td valign="bottom" align="center">119</td>
<td valign="bottom" align="center">129</td>
<td valign="bottom" align="center">149</td>
<td valign="bottom" align="center">231</td>
<td valign="bottom" align="center">136</td>
<td valign="bottom" align="center">204</td>
<td valign="bottom" align="center">351</td>
<td valign="bottom" align="center">209</td>
<td valign="bottom" align="center">58</td>
<td valign="bottom" align="center">962</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">24</td>
<td valign="bottom" align="center">569</td>
<td valign="bottom" align="center">2055</td>
</tr>
<tr>
<td valign="bottom" align="left">getting_up</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>589</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">laying_down</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>490</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">laying_resting</td>
<td valign="bottom" align="center">57</td>
<td valign="bottom" align="center">11</td>
<td valign="bottom" align="center">48</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">61</td>
<td valign="bottom" align="center">40</td>
<td valign="bottom" align="center">38</td>
<td valign="bottom" align="center">59</td>
<td valign="bottom" align="center">
<bold>19881</bold>
</td>
<td valign="bottom" align="center">41</td>
<td valign="bottom" align="center">41</td>
<td valign="bottom" align="center">553</td>
<td valign="bottom" align="center">63</td>
<td valign="bottom" align="center">66</td>
<td valign="bottom" align="center">74</td>
<td valign="bottom" align="center">207</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">24</td>
<td valign="bottom" align="center">426</td>
<td valign="bottom" align="center">105</td>
</tr>
<tr>
<td valign="bottom" align="left">other</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>1118</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">3</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
</tr>
<tr>
<td valign="bottom" align="left">rub_scratch</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">
<bold>773</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
</tr>
<tr>
<td valign="bottom" align="left">ruminating_laying</td>
<td valign="bottom" align="center">469</td>
<td valign="bottom" align="center">73</td>
<td valign="bottom" align="center">176</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">159</td>
<td valign="bottom" align="center">22</td>
<td valign="bottom" align="center">113</td>
<td valign="bottom" align="center">7</td>
<td valign="bottom" align="center">773</td>
<td valign="bottom" align="center">99</td>
<td valign="bottom" align="center">129</td>
<td valign="bottom" align="center">
<bold>52143</bold>
</td>
<td valign="bottom" align="center">981</td>
<td valign="bottom" align="center">118</td>
<td valign="bottom" align="center">460</td>
<td valign="bottom" align="center">454</td>
<td valign="bottom" align="center">8</td>
<td valign="bottom" align="center">267</td>
<td valign="bottom" align="center">514</td>
<td valign="bottom" align="center">219</td>
</tr>
<tr>
<td valign="bottom" align="left">ruminating_standing</td>
<td valign="bottom" align="center">4</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">2</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">4</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">
<bold>5453</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">12</td>
<td valign="bottom" align="center">20</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">9</td>
<td valign="bottom" align="center">8</td>
<td valign="bottom" align="center">3</td>
</tr>
<tr>
<td valign="bottom" align="left">self_grooming</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>1079</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">shaking</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>592</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">standing_resting</td>
<td valign="bottom" align="center">246</td>
<td valign="bottom" align="center">101</td>
<td valign="bottom" align="center">79</td>
<td valign="bottom" align="center">31</td>
<td valign="bottom" align="center">304</td>
<td valign="bottom" align="center">352</td>
<td valign="bottom" align="center">29</td>
<td valign="bottom" align="center">40</td>
<td valign="bottom" align="center">112</td>
<td valign="bottom" align="center">248</td>
<td valign="bottom" align="center">57</td>
<td valign="bottom" align="center">148</td>
<td valign="bottom" align="center">142</td>
<td valign="bottom" align="center">186</td>
<td valign="bottom" align="center">19</td>
<td valign="bottom" align="center">
<bold>15540</bold>
</td>
<td valign="bottom" align="center">41</td>
<td valign="bottom" align="center">29</td>
<td valign="bottom" align="center">196</td>
<td valign="bottom" align="center">393</td>
</tr>
<tr>
<td valign="bottom" align="left">stretching</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>390</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">throw_head</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">1</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">
<bold>421</bold>
</td>
<td valign="bottom" align="center">0</td>
<td valign="bottom" align="center">0</td>
</tr>
<tr>
<td valign="bottom" align="left">vigilance</td>
<td valign="bottom" align="center">78</td>
<td valign="bottom" align="center">5</td>
<td valign="bottom" align="center">241</td>
<td valign="bottom" align="center">45</td>
<td valign="bottom" align="center">277</td>
<td valign="bottom" align="center">182</td>
<td valign="bottom" align="center">24</td>
<td valign="bottom" align="center">6</td>
<td valign="bottom" align="center">172</td>
<td valign="bottom" align="center">69</td>
<td valign="bottom" align="center">60</td>
<td valign="bottom" align="center">225</td>
<td valign="bottom" align="center">300</td>
<td valign="bottom" align="center">32</td>
<td valign="bottom" align="center">47</td>
<td valign="bottom" align="center">545</td>
<td valign="bottom" align="center">9</td>
<td valign="bottom" align="center">99</td>
<td valign="bottom" align="center">
<bold>16106</bold>
</td>
<td valign="bottom" align="center">195</td>
</tr>
<tr>
<td valign="bottom" align="left">walking</td>
<td valign="bottom" align="center">105</td>
<td valign="bottom" align="center">69</td>
<td valign="bottom" align="center">40</td>
<td valign="bottom" align="center">65</td>
<td valign="bottom" align="center">733</td>
<td valign="bottom" align="center">1410</td>
<td valign="bottom" align="center">43</td>
<td valign="bottom" align="center">14</td>
<td valign="bottom" align="center">82</td>
<td valign="bottom" align="center">137</td>
<td valign="bottom" align="center">53</td>
<td valign="bottom" align="center">154</td>
<td valign="bottom" align="center">202</td>
<td valign="bottom" align="center">98</td>
<td valign="bottom" align="center">13</td>
<td valign="bottom" align="center">535</td>
<td valign="bottom" align="center">32</td>
<td valign="bottom" align="center">10</td>
<td valign="bottom" align="center">188</td>
<td valign="bottom" align="center">
<bold>17917</bold>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The columns represent the predicted behavior; rows represent the actual observed behavior.Bold values are the number of observations correctly predicted by the model.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>In this study, tri-axial accelerometer signatures were assigned to observed behaviors of free-ranging cattle based on supervised machine learning algorithms and using different accelerometer features and running mean smoothing windows. We found that (1) using a long running mean (20 seconds) translated to best model performance across all model categories, and (2) model performance remained excellent when using orientation-dependent instead of orientation-independent features, or when adding more behaviors (with accurate classification even with minority class behaviors i.e., allogrooming, suckling calf).</p>
<p>How tight and in which direction collars were deployed, and terrain ruggedness through which the cows navigated, varied widely in our study. We therefore expected that orientation-independent features derived from accelerometer data, would lead to better prediction performance than orientation-dependent features (hypothesis H1). However, models based solely on orientation-dependent features performed better than those based on orientation-independent features.</p>
<p>While our study assessed the outcome and performance of either orientation-dependent or -independent features, previous studies have combined up to 60 features of both types to increase model performance (<xref ref-type="bibr" rid="B37">Lush et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B50">Riaboff et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al., 2022</xref>). However, as our best model with solely orientation-dependent features had excellent performance (0.997 accuracy, 0.961 precision, 0.985 recall), we did not need to extend the model by including additional orientation-dependent and -independent features. We believe our model performed so well in part because the accelerometer was placed along with the battery and other collar electronics on the low side of the collar. The combined weight of the unit seems to hold the accelerometer in place, independently from collar tightness. In comparable studies on marine mammals, accelerometer placement is highly variable (<xref ref-type="bibr" rid="B59">Shepard et&#xa0;al., 2008b</xref>).</p>
<p>Moreover, and contrary to our predictions, we found that correcting for collar orientation did not improve the model performance. Indeed, both corrected and uncorrected models performed similarly. While placing the accelerometer backwards can impact axes values, the accelerometer unit remains below the neck of the animal and hangs similarly for all individuals. In fact, backwards placement of the accelerometer affects only the mean of the X and Y axes, with a more pronounced effect for the X axis, as well as the mean pitch values; the variance of the Z axis and the pitch are not affected by orientation (<xref ref-type="bibr" rid="B70">Wang et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B12">Chang et&#xa0;al., 2022</xref>). Despite variations in feature values, the amplitude and pattern of movement remain consistent, which might explain why orientation-corrected models did not outperform uncorrected models. Additionally, the sample size in our study is large enough to rule out noise in the data due to collar deployment error, making our models robust (<xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al., 2022</xref>). Furthermore, cattle are large and slow animals, and behavior-specific body movements can be more easily differentiated compared to small-sized animals. This might be more challenging for smaller, faster moving species, and placement of accelerometers in those species, and placement of accelerometers in those species is likely more important (<xref ref-type="bibr" rid="B21">Gr&#xfc;new&#xe4;lder et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B8">Brewster et&#xa0;al., 2018</xref>).</p>
<p>We initially hypothesized that the detection of less frequent behaviors, such as social interactions and body care movements, would be impacted by the smoothing of the data, as these behaviors might not be detected by long running mean windows. Contrary to our predictions, we found an increase in model performance across all categories (orientation-dependent, orientation-dependent with correction, orientation-independent) with increasing smoothing window, with highest performance when using a 20 second running mean. Other studies have shown that smoothing of the data increases the classification success by reducing noise, and that larger animals often require a longer running mean as their movements are generally slower (<xref ref-type="bibr" rid="B58">Shepard et&#xa0;al., 2008a</xref>). However, this often results in a decrease in classification success for behaviors which are rare and short in time (<xref ref-type="bibr" rid="B39">Mansbridge et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B12">Chang et&#xa0;al., 2022</xref>). In our study we did not find such a decrease. An explanation might be that we excluded all behaviors shorter than 5 seconds, resulting in a large enough difference in mean and variance of the features for those behaviors for successful classification. Furthermore, the large number of observations in the minority classes (rarer behaviors) across individuals might have contributed positively during the training of the model, resulting in high model performance.</p>
<p>Finally, we predicted that adding more behaviors would affect model performance negatively and expected that the addition of behaviors would lead to loss of model performance (H2) as it has been shown in previous studies (<xref ref-type="bibr" rid="B68">V&#xe1;zquez Diosdado et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B37">Lush et&#xa0;al., 2018</xref>). Contrary to our prediction, we found that models performed well with an increase of behaviors and could accurately predict less frequent behaviors such as head butting, throwing head and shaking. Less frequent behaviors were observed for fewer individuals than more common behaviors, which may lead to a stronger impact of individual-specific accelerometer signatures in the random forest models. We addressed this through class weighting, and individual-based cross validation. Even though there was variation in prediction success across individuals and behaviors, the overall model performed similarly well as with the 5-fold (i.e. with random 80/20 data split) cross-validation model.</p>
<p>This study&#x2019;s sample size of individuals is larger than most accelerometry classification studies of free-ranging cattle (<xref ref-type="bibr" rid="B13">Chapa et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B33">Kour et&#xa0;al., 2021</xref>). <xref ref-type="bibr" rid="B51">Riaboff et&#xa0;al. (2022)</xref> recommended using a minimum of 10 animals and emphasized more robust analysis with at least 25 animals and a variety of breeds and farms. While we exceeded this recommendation (n = 38 individuals distributed on four breeds and four farms), we did not specifically account for breed, farm or individual characteristics such as body weight or age, as this would require an even larger sample.</p>
<p>Interestingly, we were able to differentiate between behaviors that we expected to have similar accelerometry signatures, such as laying ruminating and standing ruminating. When looking at posture, cattle laying causes the angle of the accelerometer to vary slightly compared to when they are standing, as the electronic housing often leans against the individual&#x2019;s chest. Similarly, we could identify vigilance behavior, which is likely due to our decision rule defining individuals as being vigilant only when no other behavior happened (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure&#xa0;1</bold>
</xref>).</p>
<p>Often, scientific studies develop models and tools that are appropriate for experimental settings, but too expensive or impractical to be used for commercial settings. Our results based on the accelerometers contained in the commercial Nofence collars open up for a range of end-user applications. For example, these could be Nofence tools for easy handling by the customers, such as a built-in algorithm in the collar converting accelerometer data directly to behavioral states or to time budget summaries, which could be continuously transmitted to the farmer. This could allow for an easy and fine-scale supervision and monitoring of free-ranging cattle in remote areas or dense habitats. Furthermore, the success in classification of ruminating, vigilance and social behaviors could contribute to the study of free-ranging cattle welfare, stress and productivity, through the quantification of precise nutrient intake and energy expenditure.</p>
<p>In conclusion, our study succeeded in categorizing high resolution accelerometer data into behaviors for free-ranging cattle in rugged terrain of the boreal forest. Not only were collar deployment errors confirmed to not significantly impact model performance, but our models also showed success in detecting more behaviors than previously published, including less frequent behaviors other than resting, grazing, ruminating and walking. Calibrating such data with an array of different behaviors makes a valuable contribution to livestock precision farming in extensive rangeland systems. It may allow farmers to monitor the welfare of their animals continuously and to detect non-normal behaviors caused by e.g. diseases or carnivore attacks.</p>
</sec>
<sec id="s5" sec-type="data-availability">
<title>Data availability statement</title>
<p>The data and R scripts to replicate the models of this study are openly available in Dataverse NO at <uri xlink:href="https://doi.org/10.18710/ND4CLL">https://doi.org/10.18710/ND4CLL</uri>.</p>
</sec>
<sec id="s6" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>Ethical review and approval was not required for the animal study because this study used commercially available GPS/virtual fence collars in Norway and is approved by the Norwegian authorities for use on cattle. Written informed consent was obtained from the owners for the participation of their animals in this study.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>EV, LJN, MS, BZ, and ALE conceived the study. BZ and ALE secured the funding. LJN conducted the filming and video analysis. EV and MS conducted the accelerometry analysis, with additional support of OD. EV and LJN drafted the manuscript. MS, BZ, AH, MT, OD, ALE reviewed and commented on the initial drafts. All authors contributed to the ideas and edits to the manuscript and approved the submitted version.</p>
</sec>
</body>
<back>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>This research is part of the CarniForeGraze project which is funded by the Norwegian Research Council (project number 302674).</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We would like to thank the farmers Anne Dieset, Lasse Holter, &#xd8;ystein Lageraaen, and Jens Gunnar Voldmo, for their cooperation and permission of approaching and filming their cattle, and for sharing of their data. Furthermore, we would also like to thank Nofence AS for their support in obtaining the accelerometry data. Additionally, we would like to thank Pierre Lissillour for his help during the filming, Malena Diaz G&#xf3;mez, Josh Hauer and Irene Garcia Cuesta for their help with the video analysis and Saskia H. Wulff for her illustration.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s11" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fanim.2023.1083272/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fanim.2023.1083272/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Presentation_1.pdf" id="SM1" mimetype="application/pdf"/>
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