1. Introduction
The Eurasian Scops Owl (
Otus scops) is a predominantly nocturnal owl belonging to the western Palaearctic faunal zone [
1]. As one of Europe’s smallest raptors, the species is ecologically characterized by its high degree of specialization on insectivorous prey [
2]. This specialization dictates both its habitat selection and its migratory life history. The species is largely an Afro-Palaearctic long-distance migrant, with the majority of individuals moving to sub-Saharan Africa during the winter months [
1]. For the Scops Owl specifically, quantitative data on density and breeding performance remains scarce relative to other European owl species [
3,
4]. Despite considered as Least Concern [
5], reports of declines in countries at the edge of its distribution [
6,
7] and within traditional strongholds (e.g., Italy, Greece, Spain) [
2,
8,
9], yielded an overall population decline for the European and Mediterranean populations (IUCN). The owl’s dependence on invertebrates is profound, with diet analysis in some breeding season studies demonstrating that insect material can comprise up to 90% of the prey items consumed, numerically [
2,
7]. This exceptional degree of specialization, particularly on Orthoptera, makes
O. scops highly vulnerable to the systemic decline in insect populations [
10,
11] and to changes in insect communities [
12] observed across Europe. This perspective firmly establishes the Eurasian Scops Owl as an indicator of the ecological health and invertebrate biomass availability in European agro-ecosystems [
7,
13].
The analysis of pellets and nest remains has long demonstrated that O. scops relies primarily on insects while breeding [
2,
7,
8,
14]. Unfortunately, pellet analysis inherently underestimates the intake of soft-bodied prey due to thorough digestion, a methodological limitation [
15]. Direct observation and nestbox cameras effectively overcome bias by allowing researchers to accurately record every prey item delivered to the nest, regardless of digestibility. This technique has been used in combination with pellets [
2], demonstrating that traditional pellet analysis systematically under-represents the total prey items consumed. Several studies on small owls, such as Aegolius funereus and Athene noctua, confirmed the precise evaluation of representation of individual prey when using cameras, without biasing the proportion of soft-bodied prey such as earthworms [
16,
17]. But while its diet is well-documented as primarily insectivorous, focusing on Orthoptera and Lepidoptera, the fine-scale mechanisms of prey selection remain poorly understood.
The primary objective of this study is to describe, with high temporal and taxonomic resolution, the trophic ecology and provisioning behavior of a breeding pair of Otus scops during the critical nestling phase. Specifically, we aim to:
i) Quantify the actual diet through continuous video monitoring to avoid bias detection; ii) Analyze energetic profitability (biomass vs. abundance), contrasting the numerical frequency of prey items with their actual biomass contribution to determine if there is active selection towards prey of higher energetic value; iii) Describe circadian provisioning rhythms: Determine nocturnal activity patterns and verify the existence of strategic feeding peaks designed to maximize chick satiation. iv) Document prey handling behaviors: Record direct evidence of food preparation (e.g., removal of ovipositors or legs) and “surplus provisioning” decisions (such as the rejection or caching of vertebrate prey).
4. Discussion
Our study provides a high-resolution insight into the trophic ecology of
Otus scops, revealing that behind its apparent generalist insectivorous diet lies a highly sophisticated and selective provisioning strategy. By employing continuous video monitoring, we were able to detect prey items and behavioral nuances — such as specific prey processing and sex-biased selection — that are typically obscured in traditional studies based on indirect sampling techniques, such as pellet analysis [
15]. However, we acknowledge that these findings stem from the intensive monitoring of a single breeding pair over a ten-day period; therefore, while this case study offers a detailed mechanistic understanding of provisioning decisions, broader research is required to confirm the generalizability of these selective strategies at the population level.
Scops owl foraging activity was strictly nocturnal, commencing on average at 21:58 (earliest record 21:15) and ceasing at 05:40 (latest record 05:55), coinciding with the astronomical twilight. This pattern aligns with the described circadian rhythm of the species [
1] and matches the phenology of their main prey [
26]. Most Orthoptera identified species, including
Uromenus rugosicollis,
Thyreonotus corsicus,
Barbitistes fischeri, and
Meconema thalassinum, are strictly nocturnal or evening-active, while
Tettigonia viridissima exhibits crepuscular habits, in contrast to the diurnal
Locusta migratoria and
Decticus albifrons which are associated with hot, sunny weather and are unlikely to be available during the owl’s foraging window [
26]. Provisioning rhythms were not uniform, with a peak of activity recorded during the first hour of darkness (22:00 h Local Time)[
27], reaching rates of ~ 14.5 items/hour, a decline between 23:00 -and 4:00, and a secondary peak before dawn (05:00 h). This pattern of prey delivery was different from the one found in Hungary, where the first period of the night showed higher provisioning rates and then declined until dawn [
27].
Our results confirmed the strong preference for Orthoptera in the study area, which were considered the staple prey during the breeding period, both in frequency and biomass, as in other European countries [
2,
7,
8,
14,
27,
28]. Quantitative analysis showed that males delivered smaller prey (2.57 ± 0.93 cm) compared to the joint (male + female) hunting period (2.90 ± 1.03 cm). Due to allometric scaling [
19], this moderate 12.8% length increase translates into a disproportionate 40.2% biomass gain (0.89 g vs. 1.25 g), illustrating an ‘allometric amplification’ of profitability. While this size divergence suggests niche partitioning (intersexual-competition hypothesis), the fact that the male alone matched the pair’s daily biomass delivery (75.0 vs. 67.2 g) strongly supports the small-male hypothesis [
29], linking smaller size to superior foraging agility and efficiency. These differences can arise because females forage in the immediate vicinity of the nest selecting accessible prey, and males exploit larger territories to capture high-profitability vertebrates, assuming higher search costs [
16].
An interesting finding is the significant predation bias towards female orthopterans, particularly
Meconema thalassinum. This small species is ecologically rare and cryptic; its unique drumming call is virtually inaudible to the human ear [
18,
26]. The unexpectedly high prevalence of elusive species such as
M. thalassinum and
Tettigonia viridissima [
14,
30] is likely explained by their strong positive phototaxis, which drives them to congregate around artificial light sources [
26].
O. scops appear to exploit this ‘streetlight trap’ mechanism, foraging in areas where prey are unnaturally concentrated. This behavior is facilitated by the owl’s synanthropic habit of nesting in villages [
2], where they can directly benefit from artificial illumination, and light pollution is known to disrupt insect behavior [
31], triggering positive phototaxis in many nocturnal species [
32]. This phenomenon can create ‘ecological traps’ or, conversely, ‘foraging hotspots’ [
33,
34] where phototactic prey congregate, potentially altering the predator’s natural diet composition and foraging efficiency. The pervasive impact of artificial light at night is well-documented, affecting critical aspects of survival and behavior across taxa, ranging from foraging and colonization to migration and seasonal reproduction [
35]. Consequently, the diet composition observed here can be biased by urban lighting and may not fully represent the species’ foraging ecology in pristine, unlit environments.
Orthoptera females dominated the diet numerically (2:1 ratio) and, crucially, contributed significantly more biomass than males. These results contradict the ‘passive acoustic detection’ hypothesis, which posits that owls should capture more males due to the conspicuousness of their mating calls [
25]. Instead, our findings align with the ‘mobility hypothesis’ proposed by [
25], which suggests that females are rendered vulnerable not by acoustic signaling, but by their increased mobility during phonotaxis or oviposition. Furthermore, we propose that
O. scops engage in active selection based on profitability. As confirmed by our biometric data, female-biased Sexual Size Dimorphism (SSD) acts as a powerful multiplier of energetic value. This is driven by the specific allometric relationship between length and biomass in Ensifera [
19]. Due to this power law, even moderate differences in body length translate into disproportionate gains in biomass. For instance, while
T. viridissima females in our study were only 27% longer than males (SSDL = 1.27), this allometric amplification results in them providing nearly double the biomass (SSDW = 1.92). Therefore, by actively selecting the larger sex, the predator capitalizes on this non-linear profitability, prioritizing the exponentially larger ‘energy package’ of the female over the acoustic detectability of the smaller singing male. However, it must be noted that without concurrent data on prey availability in the foraging range, strict positive selection cannot be statistically confirmed, despite the balanced sex-ratio [
25].
An interesting observation is the systematic removal of the sclerotized ovipositor from female
T. viridissima. To our knowledge, this is the first documented observation of systematic ovipositor removal in
Otus scops. This behavior optimizes energy intake (as predicted by Central Place Foraging theory; [
36]), by discarding this indigestible and potentially dangerous appendage, parents mitigate physical injury and maximize energy density for nestlings with limited gut capacity [
37]. But also reveals a critical bias in traditional diet analyses: the removal of the primary diagnostic feature of female katydids [
25] likely leads to a significant underestimation of female predation in pellet-based studies.
A key finding is the marked discrepancy between numerical frequency (>98% insects) and biomass contribution. While the diet appears numerically insectivorous during the breeding period [
7,
8,
14], the energetic foundation relies heavily on vertebrates and large orthopterans. This suggests the Scops Owl operates under a mixed foraging strategy consistent with the principles of Central Place Foraging [
36]: the owl maintains a high-frequency inflow of low-cost prey (maintenance) while relying on stochastic, high-value captures (investment) for bulk calories. However, we believe that biomass delivered by mice could be well overestimated, since in almost half of the videos we were unable to determine which individual ate the prey (nestling, female, or none). This mice influx coincided with an exceptional demographic peak recorded by the SEMICE small mammals monitoring program in 2025 (
www.semice.org). Such superabundance reduced search costs to negligible levels, triggering an opportunistic surplus provisioning response [
38] where delivered biomass exceeded the brood’s consumption capacity. This phenomenon underscores the critical distinction between delivered biomass and effective consumption. Furthermore, this finding challenges the reliability of biomass estimates derived solely from nest remains, as the presence of uneaten prey suggests that delivery does not always equate to intake, potentially leading to overestimations of the nestlings’ actual energy consumption [
39]. Indeed, our findings mirror observations in other generalist raptors in the study area like
Buteo buteo, where large prey often go uneaten while smaller staples drive actual intake [
40]. In our study, rodents likely functioned as a redundant insurance resource, while the easier-to-process female orthopterans acted as the true energetic driver. This underscores that assessing diet solely based on deliveries — without verifying consumption — can lead to biased conclusions regarding the functional value of prey [
17].
Figure 1.
Schematic diagram of the video monitoring setup installed at the nest site. The wooden nest box was modified with a 20 cm cylindrical perch to facilitate adult access and prey delivery observations, and a transparent methacrylate side panel to allow recording of internal nesting activity. An external metal arm mounted on the tree trunk holds a camouflage-patterned video camera, powered by an integrated solar panel, oriented to cover both the entrance hole and the transparent panel.
Figure 1.
Schematic diagram of the video monitoring setup installed at the nest site. The wooden nest box was modified with a 20 cm cylindrical perch to facilitate adult access and prey delivery observations, and a transparent methacrylate side panel to allow recording of internal nesting activity. An external metal arm mounted on the tree trunk holds a camouflage-patterned video camera, powered by an integrated solar panel, oriented to cover both the entrance hole and the transparent panel.
Figure 2.
Videoframes extracted after nest prey provisioning records in the Scops Owl nest. (a) Uromenus rugosicollis, (b) Tettigonia viridissima, (c) Bacillus rossius, (d) Mus spretus.
Figure 2.
Videoframes extracted after nest prey provisioning records in the Scops Owl nest. (a) Uromenus rugosicollis, (b) Tettigonia viridissima, (c) Bacillus rossius, (d) Mus spretus.
Figure 3.
Composition of the diet delivered to Otus scops nestlings. The left chart illustrates the provisioning composition based on numerical frequency (% N), showing a massive dominance of Orthoptera (64.5%). The right chart displays the composition based on estimated fresh biomass (% g), highlighting the significant energetic contribution of vertebrates (Rodentia, 19.9%) to the chicks’ intake, despite their low numerical abundance. Percentages are displayed only for categories contributing more than 2% to the total. “Undetermined” items were assigned zero biomass. Colors correspond to the prey categories listed in the legend.
Figure 3.
Composition of the diet delivered to Otus scops nestlings. The left chart illustrates the provisioning composition based on numerical frequency (% N), showing a massive dominance of Orthoptera (64.5%). The right chart displays the composition based on estimated fresh biomass (% g), highlighting the significant energetic contribution of vertebrates (Rodentia, 19.9%) to the chicks’ intake, despite their low numerical abundance. Percentages are displayed only for categories contributing more than 2% to the total. “Undetermined” items were assigned zero biomass. Colors correspond to the prey categories listed in the legend.
Figure 4.
Biomass contribution of Orthopteran species to the Scops Owl’s diet. Percentages represent the proportion of the total biomass (weight) provided by each identified species. The thickness of the red arrows corresponds to importance of each prey item, highlighting Uromenus rugosicollis and Tettigonia viridissima as the primary energy sources.
Figure 4.
Biomass contribution of Orthopteran species to the Scops Owl’s diet. Percentages represent the proportion of the total biomass (weight) provided by each identified species. The thickness of the red arrows corresponds to importance of each prey item, highlighting Uromenus rugosicollis and Tettigonia viridissima as the primary energy sources.
Figure 5.
Biometric characteristics of the main Orthopteran prey species delivered by adult Scops Owls to the nest. (a) Mean body length (cm) of the identified species. (b) Mean biomass (fresh weight in g) per individual. In both panels, bars represent the mean value, error bars indicate the standard deviation (SD), and N denotes the sample size for each species. Species are ranked by size/mass to illustrate the range of prey dimensions, from smaller species like Meconema thalassinum to large Tettigoniids like Tettigonia viridissima.
Figure 5.
Biometric characteristics of the main Orthopteran prey species delivered by adult Scops Owls to the nest. (a) Mean body length (cm) of the identified species. (b) Mean biomass (fresh weight in g) per individual. In both panels, bars represent the mean value, error bars indicate the standard deviation (SD), and N denotes the sample size for each species. Species are ranked by size/mass to illustrate the range of prey dimensions, from smaller species like Meconema thalassinum to large Tettigoniids like Tettigonia viridissima.
Figure 6.
Temporal patterns of Otus scops provisioning activity, comparing Orthoptera (red) with other prey types (blue). Kernel density estimates illustrate the daily distribution of (a) capture frequency (number of items) and (b) biomass delivery (fresh weight). Dashed vertical lines indicate the mean time for each category. Statistical comparisons (Mann-Whitney U test) show no significant difference in the temporal distribution of captures (p = 0.806), while the biomass per prey item differs significantly between groups (p < 0.001), driving the distinct patterns observed in biomass delivery.
Figure 6.
Temporal patterns of Otus scops provisioning activity, comparing Orthoptera (red) with other prey types (blue). Kernel density estimates illustrate the daily distribution of (a) capture frequency (number of items) and (b) biomass delivery (fresh weight). Dashed vertical lines indicate the mean time for each category. Statistical comparisons (Mann-Whitney U test) show no significant difference in the temporal distribution of captures (p = 0.806), while the biomass per prey item differs significantly between groups (p < 0.001), driving the distinct patterns observed in biomass delivery.
Figure 7.
Kernel Density Estimation (KDE) of parental provisioning return times categorized by prey type (N = 724). The graph illustrates the probability distribution of the time elapsed since the previous visit for Invertebrate prey (green) versus Vertebrate prey (orange). While invertebrate captures are associated with short return intervals (mean ~ 6.7 min), vertebrate predation involves significantly longer and more variable foraging bouts (mean ~ 17.8 min), reflecting the higher search costs associated with high-profitability prey. The difference between groups is statistically significant (Mann-Whitney U test, U = 5108.0, p < 0.001).
Figure 7.
Kernel Density Estimation (KDE) of parental provisioning return times categorized by prey type (N = 724). The graph illustrates the probability distribution of the time elapsed since the previous visit for Invertebrate prey (green) versus Vertebrate prey (orange). While invertebrate captures are associated with short return intervals (mean ~ 6.7 min), vertebrate predation involves significantly longer and more variable foraging bouts (mean ~ 17.8 min), reflecting the higher search costs associated with high-profitability prey. The difference between groups is statistically significant (Mann-Whitney U test, U = 5108.0, p < 0.001).
Figure 8.
Density distributions of Orthopteran prey size and biomass according to parental provisioning strategy. Panels show Kernel Density Estimates (KDE) for (a) Body length (mm) and (b) Fresh biomass (g). Blue curves represent deliveries during the “Male-Only Provisioning” phase (female at nest), while orange curves represent the “Mixed/Female Active Provisioning” phase (biparental foraging). Statistical analysis (Mann-Whitney U test) confirms that prey items delivered during the mixed foraging phase were significantly larger and heavier than those delivered by the male alone. Vertical lines indicate the mean for each phase and for the total (black dashed line).
Figure 8.
Density distributions of Orthopteran prey size and biomass according to parental provisioning strategy. Panels show Kernel Density Estimates (KDE) for (a) Body length (mm) and (b) Fresh biomass (g). Blue curves represent deliveries during the “Male-Only Provisioning” phase (female at nest), while orange curves represent the “Mixed/Female Active Provisioning” phase (biparental foraging). Statistical analysis (Mann-Whitney U test) confirms that prey items delivered during the mixed foraging phase were significantly larger and heavier than those delivered by the male alone. Vertical lines indicate the mean for each phase and for the total (black dashed line).
Figure 9.
(a) Sex ratio of the main Orthopteran prey species. Stacked bars display the relative frequency (%) of females (red) and males (blue) for each species identified in the diet of Otus scops. The vertical dashed line indicates a balanced 1:1 sex ratio (50%); (b) Comparison of the daily contribution of female and male orthopterans to the diet of Otus scops in terms of numerical abundance (left) and total biomass (right). Boxplots show the median (line), interquartile range (box), and full range (whiskers), with individual data points overlaid (N = 11 nights). Statistical differences were tested using the Mann-Whitney U test.
Figure 9.
(a) Sex ratio of the main Orthopteran prey species. Stacked bars display the relative frequency (%) of females (red) and males (blue) for each species identified in the diet of Otus scops. The vertical dashed line indicates a balanced 1:1 sex ratio (50%); (b) Comparison of the daily contribution of female and male orthopterans to the diet of Otus scops in terms of numerical abundance (left) and total biomass (right). Boxplots show the median (line), interquartile range (box), and full range (whiskers), with individual data points overlaid (N = 11 nights). Statistical differences were tested using the Mann-Whitney U test.
Figure 10.
Bivariate plot of Sexual Size Dimorphism in weight (SSDW) against dimorphism in length (SSDL) for Orthoptera prey species. The distribution of species relative to the isometric line (black dotted, y = x) and the allometric regression curve (red dashed, y = x2.797) highlights the exponential increase in female biomass relative to their linear size increase. Point size scales with total number of measured specimens per species.
Figure 10.
Bivariate plot of Sexual Size Dimorphism in weight (SSDW) against dimorphism in length (SSDL) for Orthoptera prey species. The distribution of species relative to the isometric line (black dotted, y = x) and the allometric regression curve (red dashed, y = x2.797) highlights the exponential increase in female biomass relative to their linear size increase. Point size scales with total number of measured specimens per species.
Table 1.
Composition of the diet delivered to Otus scops nestlings during the study period. The table displays the taxonomic classification of prey items, their absolute abundance (N), numerical frequency (%N), total estimated biomass (g), and relative biomass contribution (% Biomass). Note that biomass could not be estimated for items in the general “Undetermined” category.
Table 1.
Composition of the diet delivered to Otus scops nestlings during the study period. The table displays the taxonomic classification of prey items, their absolute abundance (N), numerical frequency (%N), total estimated biomass (g), and relative biomass contribution (% Biomass). Note that biomass could not be estimated for items in the general “Undetermined” category.
| Order |
Species |
N |
% N |
Biomass (g) |
% Biomass |
| Araneae |
|
2 |
0.28% |
0.97 |
0.13% |
| |
Araneae undetermined |
2 |
0.28% |
0.97 |
0.13% |
| Homoptera |
|
1 |
0.14% |
1.95 |
0.26% |
| |
Lyristes plebejus |
1 |
0.14% |
1.95 |
0.26% |
| Lepidoptera |
|
14 |
1.93% |
5.63 |
0.74% |
| |
Brintesa circe |
1 |
0.14% |
0.25 |
0.03% |
| |
Caterpillar |
3 |
0.41% |
1.31 |
0.17% |
| |
Heterocera undetermined |
10 |
1.38% |
4.07 |
0.54% |
| Mantodea |
|
2 |
0.28% |
4.24 |
0.56% |
| |
Mantis religiosa |
2 |
0.28% |
4.24 |
0.56% |
| Orthoptera |
|
468 |
64.64% |
559.96 |
73.81% |
| |
Uromenus rugosicollis |
111 |
15.33% |
182.92 |
24.11% |
| |
Tettigonia viridissima |
50 |
6.91% |
125.78 |
16.58% |
| |
Meconema thalassinum |
35 |
4.83% |
11.84 |
1.56% |
| |
Barbitistes fischeri |
12 |
1.66% |
9.30 |
1.23% |
| |
Decticus albifrons |
6 |
0.83% |
14.86 |
1.96% |
| |
Thyreonotus corsicus |
6 |
0.83% |
6.78 |
0.89% |
| |
Locusta migratoria |
1 |
0.14% |
2.13 |
0.28% |
| |
Orthoptera undetermined |
247 |
34.12% |
206.35 |
27.20% |
| Phasmida |
|
12 |
1.66% |
27.89 |
3.68% |
| |
Bacillus rossius |
12 |
1.66% |
27.89 |
3.68% |
| Rodentia |
|
12 |
1.66% |
154.00 |
20.30% |
| |
Mus spretus |
11 |
1.52% |
136.00 |
17.93% |
| |
Apodemus sylvaticus |
1 |
0.14% |
18.00 |
2.37% |
| Squamata |
|
1 |
0.14% |
4.00 |
0.53% |
| |
Tarentola mauritanica |
1 |
0.14% |
4.00 |
0.53% |
| Undertemined |
|
212 |
29.28% |
|
|
| |
Undetermined |
212 |
29.28% |
|
|
| TOTAL |
TOTAL |
724 |
100% |
758.63 |
100% |