1. Introduction
The primary purpose of modern zoological institutions is the conservation of wildlife. This is achieved through direct field engagement, environmental education, public awareness and advocacy, breeding programs, fundraising, as well as research collaborations and partnerships [
1]. While the conservation of wildlife is the core purpose of leading zoological institutions, maintaining high standards of animal welfare has become a central operational priority. Animal conservation and welfare have become closely intertwined, creating opportunities to combine these to enhance the species-specific knowledge required to secure the survival and the welfare of the wildlife [
1]. The importance of welfare assessments for zoological institutions lies in their role in ensuring the physical and psychological well-being of animals under human care, as stated in the current Animal Welfare Act (Animal Welfare Act, 2024). These assessments are not only critical for maintaining ethical standards but also support broader conservation goals. Zoological institutions have evolved to play a vital role in preserving and maintaining genetic diversity through coordinated breeding programs and by facilitating scientific research, supported by zoo-managed animal populations [
2].
Assessing animal welfare is often time-consuming due to the considerable variation in behavior across species. Lowland tapirs (
Tapirus terrestris L.) are generally considered to be predominantly nocturnal in situ, where they rest during the day and forage at night, particularly around midnight [
3,
4]. They are the largest native herbivores in the tropical forests of South America and play a key role in seed dispersal of various fruits and plants, helping to maintain their environment [
5]. Lowland tapirs are solitary by nature, often traveling alone and only associating during the mating season, which lasts 48 h and occurs every 50 to 80 days [
3]. In zoological institutions, lowland tapirs are long-lived mammals, which emphasized the importance of efficient and sustainable long-term welfare monitoring.
Differences between zoological enclosures and lowland tapirs’ natural environment complicate accurate welfare assessment. Identifying the optimal observation period is therefore essential for constructing reliable activity budgets. A common tool for behavioral studies is the use of ethograms, which categorize specific behaviors [
6]. When applied over a defined observation period, an ethogram can be used to construct both time and activity budgets of the focal animal [
7], allowing researchers to determine when and how behaviors are expressed.
The general aim of this study was to optimize welfare assessment of lowland tapirs in zoological institutions by identifying an efficient observation period and exploring the potential of automated behavioral assessment methods. Specifically, the objectives were to identify the most efficient time period for assessing the welfare of lowland tapirs. Furthermore, the aim was to explore whether machine learning (ML) could support behavioral assessment by providing rapid, automated estimates of activity and to evaluate whether automated tracking could serve as a proxy for manual ethogram-based observation, potentially reducing observer effect while maintaining meaningful welfare insight.
2. Materials and Methods
In this study, the activity pattern of six lowland tapirs was studied, with three individuals from Aalborg Zoo and three at Randers Regnskov.
The lowland tapirs in Aalborg Zoo consisted of a 21-year-old female (F1A), a 15-year-old male (M1A), and their 1-year-old female calf (F2A). F1A arrived from Newquay Zoo in England in 2004, and M1A arrived from Neuwied Zoo in Germany in 2019.
In Randers Regnskov, the lowland tapirs consisted of a 7-year-old female (F1R), a 7-year-old male named (M1R), and their 1-year-old female calf named (F2R). F1R arrived from Poznan Zoo in Poland in 2021, and M1R arrived from Zooparc de Beauval in France in 2019. F1A and F1R were pregnant at the time of this study. As the first-generation lowland tapir, M1R is vital for the lowland tapir breeding program, by lowering the risk of inbreeding with the potential introduction of new genes present in in situ individuals.
The enclosure at Aalborg Zoo comprises an inside area of 35 m
2 and an outside area of 780 m
2. The interior area contains sleeping pens along with a pool where the lowland tapirs can swim and dive. The temperature and humidity resembled rainforest conditions. The outside area contains an open area, where the lowland tapirs can walk unobstructed and the weather conditions were determined by local seasonal changes (
Supplementary Figure S1). During the test period, the lowland tapirs shared the outside enclosure with three female vicuñas (
Vicugna vicugna).
The enclosure in Randers Regnskov consists of an inside area of 114 m
2, with rainforest-like climate. This enclosure was divided between an area with concrete flooring and an imitation of a forest floor with dirt and wood chips. It also contains a pool which allows them to swim and dive. The lowland tapirs share the enclosure with various animals, such as birds and tamarins, moving freely in the entire South American biosphere section (
Supplementary Figure S2).
Before the experiment, an ethogram was constructed based on previous studies of lowland tapirs according to Montenegro [
8]. This newly constructed ethogram was later adapted to cover the behavior of zoo lowland tapirs, based on in-person observations (
Table 1).
A definition of what constituted a behavior was provided, ensuring a coherent understanding of the behavioral categories, thereby supporting reliable data analysis and reducing observer bias. Swimming was recorded as a separate behavioral category due to the availability of water pools and the species-specific use of aquatic environments for thermoregulation, resting, defecation, and enrichment, and was therefore distinguished from other locomotor behaviors. The behaviors observed in the recordings were cataloged, including the start time of the observation period. This allowed a general understanding of when the animals were active and which behaviors were common during different times of the day. On top of behaviors, an “out of sight” category was also made for when an individual was absent or left during a recording.
Data were collected with the use of wildlife cameras (3 × Boly BG584 (Boly Inc., Santa Clara, CA, USA) and 1 × Spromise S378 (Spromise, Alexandria, Australia)). Before the experiment, the cameras were placed in the lowland tapir enclosure to test their recording reliability over 24 h. This also served to acclimate the cameras to the indoor enclosures, especially towards the increased humidity. These data were not used in the results. Thereafter, an interobserver reliability test was made with the use of camera reliability data [
9]. This test was needed for a comparison between observers to establish the accuracy of all observers [
9], thereby facilitating valid conclusions and supporting meaningful scientific analysis.
The test was made by observing nineteen video recordings individually among five observers. The observations were then compared with one another, and the percentage difference among the observers was calculated. For the observations to be considered excellent, a minimum of 75% similarity is needed [
9]. If any individual failed to meet the requirement, the definitions of behaviors were further specified, and a new set of nineteen videos was observed. Afterwards, the interobserver reliability test results were collected, and the reliability coefficient was calculated by using Fleiss Kappa test, in RStudio (2023.12.1 Build 402, Posit Software, PBC, Boston, MA, USA) [
10]. Fleiss Kappa test is used to measure the reliability between multiple independent raters’ results in determined classified categories, such as walking, standing, etc. (
Supplementary Table S1).
In the case of both indoor and outdoor areas, both areas were monitored with wildlife cameras. The cameras were placed to minimize the area of blind spots in the lowland tapir enclosures. In Aalborg Zoo, two cameras were installed indoors and two outdoors. In Randers Regnskov, three cameras were installed indoors. The cameras were configured to trigger on the detection of movement and would record for 30 s, meaning each video recorded counted as an observation period. The animals were monitored over a three-day (72 h) period. The cameras’ batteries and memory cards were replaced daily, every 24 h. The recording period in Aalborg Zoo began on 1 November 2024 and ended on 4 November 2024. In Randers Regnskov, the recording period began on 18 November 2024 and ended on 21 November 2024. After the 72 h period, the recordings were sorted based on which camera they came from. If a lowland tapir was observed in the recording, all behavioral tendencies were noted in Google Sheets (Google LLC, web-based application, Mountain View, CA, USA) (along with the date and start time of the recording. These recordings were used, with the standardized ethogram, to observe individuals’ activity budgets from the two different enclosures and construct a time budget for the period [
6]. These time budgets were used to identify the most efficient period for assessing the welfare of lowland tapirs.
To explore automated alternatives to manual behavioral scoring, an ML approach was implemented, focusing on activity level detection. ML refers to computational methods that identify patterns in data and was here used to support behavioral assessment by providing automated estimates of activity. Using the video recordings, lowland tapir movement was analyzed with SLEAP (Version 1.4.1a2, Princeton University, Princeton, NJ, USA), an open-source deep learning-based framework for multi-animal pose tracking. Rather than classifying detailed behaviors, the analysis focused on estimating general activity levels (e.g., periods of movement versus rest), which is more robust to issues such as partial occlusion and limited nighttime visibility.
SLEAP (Version 1.4.1a2) was used to track the head, shoulder, and hip of each lowland tapir across time. To quantify movement, a body centroid was calculated for each frame by averaging the x- and y-coordinates of the shoulder and hip points. The Euclidean distance between centroids in consecutive frames was then calculated to estimate moment-to-moment movement. Because this raw displacement data is subject to noise from pose estimation jitter and small positional fluctuations, a smoothing procedure was applied. A centered rolling average over thirty frames (equivalent to one second at 30 frames per second) was used to reduce short-term noise and better reveal longer-term activity trends.
The resulting smoothed activity profiles allowed visualization of changes in movement intensity over time, facilitating identification of patterns such as bursts of locomotion or extended periods of rest. This approach provides a scalable and objective method for assessing activity levels across large video datasets without the need for manual annotation.
Statistical visualizations of data were made using Google Sheets (Google LLC, web-based application). This included (1) tables of activity budgets for every 24 h and all 72 h combined for each lowland tapir (
Table 2;
Supplementary Tables S2 and S3). (2) In addition, diagrams illustrating the hourly distribution of instances of behaviors across all 72 h were combined for each lowland tapir (
Figure 1A–F). (3) A table showing the percentage of how many times “out of sight” was denoted for each lowland tapir with the total observation periods (
Supplementary Table S4). “Out of sight” was also denoted every time a lowland tapir was on camera and walked out of the picture.
Statistical tests were made using the program RStudio (2023.12.1 Build 402) [
10], which uses the coding language R. The level of significance was set to
α = 0.05. Chi-square test was used to determine and analyze differences in behavior between observation days for each lowland tapir, between total behaviors across the 72 h observation period among individuals from their respective zoological institutions and between total behaviors across zoological institutions based on their family role comparing dam-dam, sir-sir, and calf-calf (
Supplementary Tables S5–S9).
Furthermore, Mann–Whitney
U-test was used to test for differences in medians for two independent groups, with time intervals in hours, which in summary aggregate to a 24 h period (
Supplementary Tables S10–S14). This helped to conclude in which period to observe possible differences in activity for zoologically based lowland tapirs.
4. Discussion
The present study is based on a limited sample size comprising six lowland tapirs from two zoological institutions, including parent–offspring pairs. Consequently, the findings should be interpreted within an exploratory and method-oriented framework rather than as population-level inferences. Furthermore, environmental conditions and enclosure characteristics were not quantified or analyzed as independent variables but instead formed part of the overall context in which activity patterns were assessed. The discussion therefore focuses on institution-specific patterns shaped by multiple interacting factors.
In addition, welfare assessment in this study is based on activity patterns, which provide an indirect but a commonly applied indicator of animal welfare. The integration of physiological stress measures would provide valuable complementary insight and represent an important direction for future research beyond the scope of the present study.
The results show a clear difference in behaviors, particularly “resting” and “swimming”, between Aalborg Zoo and Randers Regnskov, despite the individuals sharing the same genders and familial roles. This can be attributed to a difference in enclosures, both in area size and content, as seen in
Supplementary Figure S1 and S2 [
11]. Within the two institutions, the individuals show a greater degree of similarity, generally following the same behavioral patterns, as seen on the histograms in
Figure 1A–F. This indicates that the gender and age of the lowland tapirs play a lesser role in determining their behavior, while the enclosure type plays a greater role. This is supported by their natural behavior, where there is little to no difference between the genders in terms of behavior [
12].
Both groups of lowland tapirs show a higher activity during the daytime, which contrasts with their documented natural behavior in situ [
3] ). A more natural enclosure, like Randers Regnskov, would be expected to facilitate more natural behavior. Other factors, like feeding time and visitor numbers, could have attributed to this, as eating behavior seems to be focused during the afternoon hours. Other studies have shown a similar pattern, though with a species of elephants (
Elephas maximus), where feeding was more prevalent during the day than night despite their natural behavior [
13]. Another effect of feeding times is the emergence of anticipatory behaviors. These often manifest as an increase in activities like walking in the form of pacing right before being fed [
14]. A study has shown that an unpredictable feeding schedule helped facilitate more natural behavior in a group of chimpanzees (
Pan troglodytes) [
15]. Implementing an unpredictable feeding schedule should be tested on lowland tapirs kept in zoos to observe potential changes in their behavior. Visitor numbers are another source of behavioral disturbances for most ex situ animals [
16]. The detrimental effect of visitor numbers is discussed in a study from Arignar Anna Zoological Park, in India, on four individuals of ex situ Indian gaur (
Bos gaurus) [
17]. The study indicates that zoo visitors can be a potential source of disturbance and stress for ex situ animals. Another study showed that an increase in visitors caused a group of Malayan tapirs (
Tapirus indicus) to become more anxious and alarmed, becoming less active, thereby lowering their welfare [
11]. Conversely, the Malayan tapirs became more active during low visitor hours, where walking and swimming became more prevalent [
11]. A different study conducted on lowland tapirs in situ in Brazil showed a similar result [
18]. The study found that the presence of humans in or near their habitat can lead to altered behavior. The study suggests that lowland tapirs exhibit behavioral adaptations to optimize resource access while minimizing disturbances in environments with dynamic or uncertain conditions [
18]. The idea that lowland tapirs modify their behavior in response to perceived environmental changes, no matter how slight, is supported by these studies. This suggests that the behavioral patterns observed in Aalborg Zoo and Randers Regnskov may also be influenced by a natural adaptive mechanism in response to human visitors.
The distribution of individual activity budgets, as shown in
Figure 1A–F, indicates that lowland tapirs at Aalborg Zoo and Randers Regnskov generally follow similar behavioral patterns within each institution. This consistency suggests that the lowland tapirs exhibit some level of synchronization. These findings support studies which challenge the long-held belief that lowland tapirs are strictly solitary animals [
12]. Instead, it has been observed that lowland tapirs can live together in family groups, typically consisting of a monogamous pair and their offspring [
12].
This study revealed significant behavioral differences between days in some cases, observed in both individuals at Aalborg Zoo and Randers Regnskov. Behaviors such as “foraging”, “resting”, and “swimming” varied daily for almost every individual.
Similarly, a study conducted on lowland tapirs in the Atlantic Forest found that daily activity patterns were influenced by factors such as temperature, lunar phases, and increased rainfall [
3].These factors affected energy utilization and encouraged foraging under less stressful conditions. During months with high precipitation and low temperatures, there was an increase in diurnal activity compared to days with higher temperatures [
3]. Additionally, periods of increased lunar light led to reduced activity and foraging, as lowland tapirs sought areas with more cover or dense vegetation to avoid potential threats from predators or humans [
3]. These factors could be considered crucial for explaining the significant behavioral differences observed between the individuals in each institution in this study. The lowland tapirs at Aalborg Zoo had access to an outdoor area of approximately 780 m
2, with minimal vegetation cover, during the study period. This suggests that lunar light might have had a pronounced impact on nocturnal activities at Aalborg Zoo.
The indoor facility at Aalborg Zoo spanned 35 m
2 and maintained high humidity (80–100% RH) and temperature. Compared to the conditions described in a previous study [
3], access to an open outdoor area during November, combined with the humid indoor environment, is likely to influence the lowland tapirs’ ability to regulate their body temperature. Research indicates that the optimal temperature range for lowland tapirs to thrive is between 21 °C and 27 °C and they struggle to adapt to significantly lower temperatures [
19]. This could be an influencing factor in their time spent outdoors in colder months, while the higher humidity and temperature indoors could impact activity level, such as a decrease in foraging or an increase in resting and swimming indoors. The size of the facility could also play a significant role in shaping these behaviors [
11].
In contrast, the facility in Randers Regnskov was located within a dome, characterized by high humidity (80–100% RH, stable temperatures, and coverage from various trees and bushes). Here, natural lunar light was reduced, and the indoor temperature remained consistent. While the individuals in Randers Regnskov did not experience the same fluctuations in environmental conditions or outdoor access as those in Aalborg Zoo, significant behavioral differences between days among individuals in Randers Regnskov still emerged, even when these factors remained constant.
All above mentioned studies highlight the importance of accounting for both macro-and micro-environmental factors, as well as abiotic and biotic elements, when conducting behavioral studies on lowland tapirs [
18]. By considering this broader context, researchers can gain a more comprehensive understanding of the factors shaping lowland tapir behavior.
Besides welfare assessment, this study demonstrates the feasibility and utility of applying pose estimation techniques, specifically SLEAP, to quantify basic activity levels in lowland tapirs using video data. By tracking key body parts and calculating centroid displacement over time, we were able to derive meaningful movement profiles that distinguish periods of rest from bouts of locomotion without relying on detailed behavioral annotations. This approach offers a promising automated alternative to manual scoring, which is often time-consuming and subject to observer bias.
The results highlight how smoothing displacement data can effectively mitigate the influence of pose estimation noise and small positional jitter inherent in ML-based tracking. The centered rolling average over one second (30 frames) proved sufficient to reveal underlying activity trends, enabling the identification of natural behavioral rhythms such as resting phases and short bursts of movement. This method’s robustness to partial occlusion and challenging lighting conditions, common in wildlife videos, is a notable advantage for studying elusive or nocturnal species like lowland tapirs.
While this approach does not classify complex behaviors or social interactions, focusing on coarse activity levels is particularly useful in ecological and behavioral monitoring contexts where large datasets demand scalable analysis. Future work could extend these findings by integrating additional body points or combining pose data with environmental or physiological measurements to better interpret the functional significance of observed activity patterns.
Given these technical constraints and the challenges associated with nocturnal video data, the ML analysis using SLEAP was intentionally limited to coarse activity level estimation (movement versus rest) to ensure robustness under conditions of partial occlusion, low nighttime visibility, and limited training data. While this approach does not capture complex behaviors or social interactions, it provides a scalable and objective proxy for general activity patterns. Extending pose estimation to multi-class behavioral classification represents an important direction for future work.
Limitations include the reliance on video quality and the potential for missed or mis-tracked points during occlusions or rapid movements, which may reduce accuracy. Additionally, while centroid displacement provides a convenient proxy for movement, it may not capture subtle postural changes or non-locomotor activities such as grooming or feeding. Addressing these limitations may require more sophisticated multi-point tracking models or complementary sensor data.
Overall, the application of SLEAP-based pose estimation for lowland tapir activity monitoring represents a valuable step toward automated behavioral analysis in wildlife research, offering an objective, reproducible, and efficient tool to quantify activity levels across extensive video datasets.
Technical difficulties with the cameras could have influenced the results, particularly when considering the total observation periods. The number of observation periods was significantly higher for Randers Regnskov, likely due to the cameras nearly covering the entire facility. Additionally, the facility housed various other species, which could have triggered the cameras to start recording. This does not influence the results when looking at individuals within institutions but can complicate the comparison between Aalborg Zoo and Randers Regnskov (
Supplementary Table S1). Furthermore, the use of percentages in the Chi-square analysis stems from the fact that the total observation counts varied between individuals and zoological institutions. While this facilitated relative comparisons across individuals and sites, it does not conform to standard Chi-square procedures, which are conventionally based on raw counts [
20,
21]. The Chi-square results should therefore be interpreted with caution and were cross-checked against the underlying raw data presented in
Table 2. Future analyses should apply Chi-square tests directly to raw frequency data to avoid potential biases.