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Research Article
The diet of Barn Owl (Tyto alba) in Georgia
expand article infoDenis Kitel, Zurab Javakhishvili§
‡ University of Bialystok, Bialystok, Poland
§ School of Natural Sciences and Medicine, Ilia State University, Tbilisi, Georgia
Open Access

Abstract

The Barn Owl (Tyto alba) has only recently colonized Georgia, yet has rapidly expanded throughout the country's lowland agricultural landscapes. Despite its increasing presence, no prior comprehensive study has examined its diet in Georgia. We analyzed 6627 prey items from Barn Owl pellets collected between 2023 and 2025 at 34 sites across five Georgian regions. After sterilization and dissection, prey remains were identified to the lowest possible taxonomic level and biomass was estimated. The results show that mammals dominate the diet, accounting for 98.6% of all prey items, with rodents (53% by number, 75% by biomass) and shrews (45% by number, 23% by biomass) being the most common prey groups. Other prey included birds, bats, amphibians, reptiles, and invertebrates. Notably, Mus macedonicus, Microtus socialis, and Crocidura spp. were the most frequent mammalian prey. Geographic variation in diet composition was observed, with some sites showing a predominance of shrews. This study provides novel insights into the Barn Owl’s feeding ecology in Georgia and offers the first regional data on several small mammal species, including updated records for Micromys minutus and Suncus etruscus. Given the owl’s preference for rodents, it may serve as a valuable component of sustainable rodent control strategies in Georgian agriculture.

Key words:

Barn Owl, diet, pellets, prey, rodents, shrews, pest control

Introduction

The Barn Owl (Tyto alba) has established itself in Georgia only in recent decades and, within approximately 25 years, has expanded across much of the country's lowland agricultural regions (Kitel et al. 2025). Its current distribution is closely tied to areas intensively utilized by local communities for farming (Belik 2024). Despite this recent and rapid spread, information on the Barn Owl in Georgia remains limited and fragmentary (Bukreev 2003; Galvez et al. 2005; Budagashvili and Javakhishvili 2024; Topaloudis et al. 2025). Although interest in the country’s biodiversity has grown in recent years (Mumladze et al. 2019), Barn Owl remains among the least studied bird species in Georgia.

As a specialized nocturnal predator, the Barn Owl plays an important role in regulating populations of small mammals, especially rodents and shrews. Analyses of owl pellets have proven to be an effective method for investigating predator diets and offer valuable insights into local small mammal fauna (Kryštufek et al. 2009; Riegert et al. 2011; Milana et al. 2018). In Georgia, dietary studies of raptors are rare and have so far focused only on diurnal species (Abuladze 2013). Moreover, recent comprehensive reviews of the country's small mammal fauna make no mention of data derived from owl pellet analysis (Bukhnikashvili et al. 2023; Kandaurov et al. 2023; Natradze et al. 2023), indicating a significant knowledge gap.

Globally, Barn Owls are increasingly recognized as effective agents of biological pest control, especially in agricultural ecosystems (Tores et al. 2005; Sebele et al. 2020; Bontzorlos et al. 2024). Although their diet is largely dominated by small mammals, they also opportunistically consume other taxa, including birds, bats, reptiles, amphibians, and invertebrates (Roulin and Dubey 2012, 2013; Vale-Gonçalves et al. 2015; Roulin 2016).

In Georgia’s fertile lowlands, where agricultural activity is intense, the Barn Owl may represent a promising natural solution for controlling rodent pests. Based on this context, the present study hypothesizes that rodents constitute the primary component of the Barn Owl’s diet in Georgia and that the species may play an ecologically and economically valuable role in supporting sustainable pest management practices.

Material and methods

Study area

Barn Owl pellets were collected across Georgia between 2023 and 2025 from 34 locations situated in five administrative regions: Adjara, Imereti, Kakheti, Kvemo Kartli, and Samegrelo-Zemo Svaneti (Fig. 1). Sampling sites were associated with known roosting and breeding areas of dispersing Barn Owls.

Fresh pellets were characterized by a glossy, lacquered surface with a blackish tint due to salivary coating, consistent with descriptions by Mikusek (2005) and Poprach (2010).

To ensure biosafety and facilitate dissection, all pellets were sterilized by oven-baking at 160°C for 40 minutes, following the protocol of Orosz-Coghlan et al. (2022), and then soaked in warm water. A total of 6627 prey items were extracted and identified to the lowest possible taxonomic level (Table 1).

Figure 1. 

A map of Georgia with 34 sites where Barn Owl (Tyto alba) pellets have been collected in 2023–2025.

Table 1.

The number of preys from Barn Owl (Tyto alba) pellets by regions in Georgia.

Region Number of sites Number of preys
Adjara 2 302
Imereti 1 37
Kakheti 16 3804
Kvemo Kartli 7 1827
Samegrelo-Zemo Svaneti 8 657
Total 34 6627

Species identification

Prey remains extracted from Barn Owl pellets included mammals, birds, amphibians, reptiles, and invertebrates. Mammals (Rodentia, Eulipotyphla, Chiroptera, Carnivora) were identified using standard taxonomic keys and monographs (Pucek 1981; Kryštufek and Vohralík 2001, 2005, 2009; Aulagnier et al. 2009; Zaitsev et al. 2014; Tembotova 2015; Kryštufek and Shenbrot 2022, 2025; Russo 2023). Species nomenclature followed recent national checklists (Bukhnikashvili et al. 2023; Kandaurov et al. 2023), with the only exception being the use of Crocidura gueldenstaedtii instead of Crocidura suaveolens, as used in earlier and modern classifications (Bukhnikashvili 2004; Gazzard and Kryštufek 2024).

Due to local morphological variation in Caucasian rodents, standard identification methods were sometimes insufficient, and additional references were consulted. For example, distinguishing Microtus socialis required intact skulls and ratios of rostrum height and interorbital constriction to condylobasal length, along with proportionally large auditory bullae (Vinogradov and Gromov 1952; Kryštufek and Kefelioğlu 2001). Although Microtus rossiaemeridionalis has recently been recorded in Georgia (Maglakelidze et al. 2024), its morphological similarity to M. obscurus necessitated treating all specimens as M. obscurus in the absence of DNA data.

Cranial features were used to separate Rattus rattus and R. norvegicus, mainly based on molar shape (Pucek 1981; Yiğit et al. 1998; Kryštufek and Vohralík 2009).

Identification of Pine Voles (Microtus majori) relied on specific M1 patterns of the lower mandible, while M. subterraneus was excluded based on its known distributions; in addition, M. daghestanicus was excluded due to its preference for high-altitude habitats, whereas all our data were collected in lowland areas (Çolak et al. 1998; Kryštufek and Vohralík 2004; Aulagnier et al. 2009).

In many cases, Mus and Apodemus specimens were too fragmented for species-level identification. For Mus, separation of M. musculus and M. macedonicus used zygomatic index (Zagorodniuk 2002; Çolak et al. 2006; Kryštufek and Vohralík 2009), though this was not always conclusive due to damaged remains. Third molar ratios provided limited accuracy. Apodemus species were identified to genus level unless diagnostic features (e.g., bullae or tooth row length) were intact, as overlapping morphotypes (e.g., A. ponticus, A. witherbyi) complicate identification (Darvish et al. 2014; Mohammadi et al. 2014; Okulova et al. 2018; Maglakelidze et al. 2024).

Bird remains were identified mainly from skulls (Brown et al. 2021; Drewitt 2024), using an osteological reference collection. Ulna shape was of limited diagnostic value (Panteleev 2005). As no published data exist on bird biomass in Georgia, we used mean body mass data from the Aras Bird Ringing Station in Türkiye, based on measurements of 2511 individuals collected in 2023–2024.

Amphibians and reptiles were not identified to species level; representative species (Hyla orientalis, Lacerta agilis) were used for biomass estimation (Shlyakhtin et al. 2019; Burraco et al. 2021). Invertebrate remains were identified by specialists (A. Seropian, A. Schröter, T. Stalling), and biomass was estimated using published mean weights, rounded to 1 g where necessary (Cornwell 1976; Meresman and Ribak 2020; Hendrix and Verdonschot 2021).

Results and discussion

A Diet composition in pellets

A total of 6627 prey items were extracted from Barn Owl pellets and identified to the lowest possible taxonomic level. Of these, 996 individuals originated from western regions (Adjara, Imereti, Samegrelo-Zemo Svaneti), and 5631 from eastern Georgia (Kakheti and Kvemo Kartli). By number and mass Rodentia prevail among other preys with 3528 individuals or 87726.7 g. The second largest group of mammals as preys is Eulipotiphla with 2985 individuals or 28037.9 g. The rest of animal groups estimate 114 individuals or 1749.1 g (Table 2).

Small mammals overwhelmingly dominated the diet: rodents accounted for 53% of all prey by number and 75% by total biomass, while shrews (Soricidae) comprised 45% by number and 23% by biomass. Together, these two groups made up 98% of total biomass (Fig. 2). Other vertebrates—bats, birds, amphibians, reptiles—and invertebrates were rare (Table 2).

Among 1099 unbroken pellets, length ranged from 21.6 to 81.0 mm (mean = 39.8 mm), and width from 14.6 to 45.9 mm (mean = 26.0 mm) (Fig. 3). Each pellet contained between 1 and 13 individual prey items (mean = 3.53) (Fig. 4). Only 6 pellets contained more than 10 prey items, with those dominated by small Soricidae. Altogether, 3883 prey items (58.6% of total) were recovered from intact pellets.

The pellet with the highest prey count (13 individuals) came from Poti (Samegrelo-Zemo Svaneti) and contained 11 Sorex volnuchini, 1 Neomys teres, and 1 Crocidura sp. Another highly diverse pellet from Shiraki (Kakheti) held 11 Crocidura sp.

Taxonomically, the Barn Owl diet in Georgia included representatives of five vertebrate classes and invertebrates (Fig. 5). Mammals dominated with 98.6% of prey by number, followed by birds (1.0%), and a combined 0.4% from amphibians, reptiles, and invertebrates.

Table 2.

Total prey composition of Barn Owl (Tyto alba) pellets in Georgia by taxonomic group and biomass contribution.

Species Numb. of Ind. West Numb. of Ind. East Total Numb. of Ind. Mean Ind. Mass (g) Total Biomass (g)
Sorex satunini 0 15 15 5.9 88.5
Sorex volnuchini 110 0 110 4.3 473
Sorex raddei /satunini 0 4 4 5.9 23.6
Neomys teres 11 0 11 18 198
Crocidura leucodon 11 206 217 12.8 2777.6
Crocidura gueldenstaedtii 22 270 292 9.3 2715.6
Crocidura sp. 318 1968 2286 9.3 21259.8
Suncus etruscus 0 41 41 1.9 77.9
Talpa levantis 6 0 6 47.1 282.6
Talpa sp. 3 0 3 47.1 141.3
Dryomys nitedula 2 3 5 24.1 120.5
Glis glis 0 2 2 89.6 179.2
Apodemus uralensis 4 0 4 20.2 80.8
Apodemus uralensis /witherbyi 9 4 13 20.2 262.6
Apodemus sp. 76 312 388 20.2 7837.6
Micromys minutus 85 4 89 9 801
Rattus norvegicus 31 1 32 259 8288
Rattus rattus 5 0 5 188.4 942
Rattus sp. 17 1 18 188.4 3391.2
Mus musculus 44 32 76 22.7 1725.2
Mus macedonicus 0 513 513 16.1 8259.3
Mus sp. 36 1013 1049 16.1 16888.9
Mesocricetus brandti 0 1 1 108.7 108.7
Arvicola amphibius 1 4 5 170 850
Microtus majori 64 1 65 23.8 1547
Microtus obscurus 88 54 142 36.4 5168.8
Microtus socialis 0 454 454 27.9 12666.6
Microtus obscurus /socialis 0 667 667 27.9 18609.3
Mustela erminea 1 0 1 150 150
Myotis blythii 0 1 1 15 15
Myotis sp. 0 1 1 3.8 3.8
Pipistrellus sp. 10 6 16 3 48
Hirundo rustica 4 0 4 20.2 80.8
Hirundinidae sp. 13 1 14 20.2 282.8
Galerida cristata 0 1 1 38.2 38.2
Acrocephalus sp. 0 2 2 11.6 23.2
Turdus philomelos 1 0 1 70 70
Turdus sp. 1 0 1 70 70
Passer domesticus 0 4 4 26.9 107.6
Fringilla coelebs 3 7 10 21.5 215
Carduelis cannabina 0 2 2 17.8 35.6
Carduelis carduelis 0 2 2 16.3 32.6
Emberiza sp. 0 1 1 26.1 26.1
Passeriformes sp. 9 13 22 22.3 490.6
Anura sp. 4 1 5 5.4 27
Squamata sp. 0 1 1 6.9 6.9
Dytiscus sp. 1 0 1 1.9 1.9
Protaetia sp. 0 1 1 1 1
Scarabaeidae sp. 3 3 6 1 6
Blatta orientalis 1 1 2 1 2
Bolivaria brachyptera 0 1 1 1 1
Calliptamus barbarus 0 1 1 1 1
Decticus albifrons 0 2 2 1 2
Decticus sp. 0 3 3 1 3
Gryllus sp. 1 5 6 1 6
Gryllotalpa gryllotalpa 1 0 1 1 1
Xeropicta derbentina 0 1 1 1 1
Rodentia 462 3066 3528 87726.7
Eulipotyphla 481 2504 2985 28037.9
Carnivora , Mustelidae 1 0 1 150
Chiroptera 10 8 18 66.8
Aves 31 33 64 1472.5
Amphibia , Anura 4 1 5 27
Reptilia , Squamata 0 1 1 6.9
Invertebrata 7 18 25 25.9
Total 996 5631 6627 117513.7
Figure 2. 

Prey composition in the diet of Barn Owl (Tyto alba) in Georgia by: A: number of individuals, B: biomass (%).

Figure 3. 

Distribution of Barn Owl (Tyto alba) pellet dimensions (n = 1099).

Figure 4. 

Number of prey items per unbroken Barn Owl (Tyto alba) pellet (n = 1099).

Figure 5. 

Selected prey remains from Barn Owl (Tyto alba) pellets: A: Mus musculus; B: Decticus sp. (Orthoptera); C: Pipistrellus sp. (Chiroptera); D: Dryomys nitedula; E: Rodentia: Apodemus sp.; F: Crocidura gueldenstaedtii; G: Sorex volnuchini; H: Hirundo rustica; I: Micromys minutus; J: Talpa levanti.

Variation across sites

Although rodents dominated the overall diet of Barn Owls in Georgia, notable geographic variation was observed. At sites with over 200 prey items (n=10) —indicating long-term owl presence—shrews (Soricidae) occasionally outnumbered rodents by number or biomass (e.g. Nazarlo, Chailuri, Poti) (Table 3). This pattern may reflect temporary declines in rodent populations, potentially driven by population cycles or pesticide use in agricultural areas.

Table 3.

The share of Rodents and Shrews in Barn Owl (Tyto alba) diet at different sites.

Site Name Nr of preys Rodents. % Shrews. %
Nr Biomass Nr Biomass
Koghoto, Kakheti 1586 68 82 31 17
Nazarlo, Kvemo Kartli 931 33 52 67 48
Gremi, Kakheti 866 70 84 30 16
Chailuri, Kakheti 424 22 38 77 62
Akhali Samgori, Kvemo Kartli 382 56 74 43 26
Avgia, Adjara 300 62 90 28 5
Poti, Samegrelo-Zemo Svaneti 286 22 51 69 39
Tkviri, Samegrelo-Zemo Svaneti 247 45 72 54 26
Kondoli, Kakheti 224 43 61 57 39
Kumisi, Kvemo Kartli 211 73 86 27 14

These local deviations align with previous observations that Barn Owls exhibit selective opportunism, primarily preying on rodents when available (Tores et al. 2005). In other countries, high consumption of shrews by Barn Owls has also been reported, including Germany (Gurney 1876), Italy (Bose and Guidali 2001), Greece (Bontzorlos et al. 2005), and Turkey (Hoppe 1986). However, due to the absence of similar pellet studies in the Caucasus region, it remains unclear whether the patterns observed here are typical for the region or unique to Georgia.

Certain species account

Owl pellet content is representative for real proportions of the small mammal species in their communities at the open areas (Andrare et al. 2015). Pellet analysis also provided new insights into the distribution of several small mammal species in Georgia.

Pygmy White-toothed Shrew (Suncus etruscus). Previously known only from eastern Georgia (Buknikashvili 2004), this species was detected exclusively in Kvemo Kartli and Kakheti. The presence of 41 individuals suggests a more common and widespread status than previously assumed, corroborating recent regional observations (Kandaurov et al. 2023). Similar records of S. etruscus in owl pellets have been reported from southeastern Turkey (Coşkun and Kaya 2013).

Harvest Mouse (Micromys minutus). Historically reported from only seven western Georgian sites, mainly from older records (Bukhnikashvili et al. 2023), M. minutus was recorded here from Adjara and, for the first time, from eastern regions (Kvemo Kartli, Kakheti), with 89 individuals identified. These findings significantly extend its known range.

Transcaucasian Water Shrew (Neomys teres) and Caucasian Pygmy Shrew (Sorex volnuchini). Despite being widely distributed in Georgia (Buknikashvili 2004), both species were found only in pellets from the Poti region. This site—formerly a marshland and now a canal system with rich vegetation and no agriculture—likely supports suitable habitats. Their absence elsewhere likely reflects a sampling bias toward agricultural landscapes, which are less favorable for these species.

Brandt’s Hamster (Mesocricetus brandti). Only a single individual was recorded at Shiraki (Kakheti), a steppe habitat considered atypical for Barn Owl hunting. However, with the owl’s ongoing range expansion into suboptimal habitats, occasional predation on large desert rodents may become more frequent.

Conclusion

This study represents the first comprehensive analysis of the Barn Owl’s diet in Georgia, covering a wide geographic range and providing a dataset of over 6600 prey items identified from owl pellets. The overwhelming dominance of small mammals, particularly rodents and shrews, confirms that the Barn Owl functions primarily as a specialist predator on terrestrial micromammals, although its capacity to exploit other taxa—including birds, bats, amphibians, reptiles, and invertebrates—reflects opportunistic tendencies when primary prey availability fluctuates.

The high representation of rodents (53% by number and 75% by biomass) supports the hypothesis that Barn Owls may play a significant ecological role in regulating rodent populations in Georgia’s agricultural landscapes. This aligns with findings from other regions where Barn Owls are already integrated into pest management strategies. At the same time, the significant presence of shrews in certain areas and years suggests sensitivity to local prey dynamics, possibly influenced by natural population cycles or anthropogenic factors such as pesticide use.

From a faunistic standpoint, the study contributes valuable records for small mammal distribution in Georgia, including the easternmost confirmations of Suncus etruscus and previously undocumented occurrences of Micromys minutus in eastern regions. These results underline the utility of owl pellet analysis as a complementary method for biodiversity assessments, especially in understudied or inaccessible habitats.

The research also identifies several methodological and interpretative challenges, including species-level identification difficulties due to cranial damage or cryptic morphology, and the need for more detailed ecological data on the small mammal communities being preyed upon.

Going forward, a multi-seasonal and multi-year monitoring program, complemented by live-trapping and habitat assessments, would help clarify temporal prey fluctuations, reveal effects of agricultural practices, and further test the feasibility of using Barn Owls as a nature-based solution for rodent control. Such efforts could also enhance our understanding of the ecology of prey species, many of which remain poorly documented in Georgia.

Altogether. the Barn Owl emerges not only as a key predator with potential agricultural utility, but also as a valuable bioindicator and sampling tool for small mammal communities in the region. As Georgia continues to integrate biodiversity research into conservation and land-use planning, expanding the scope of such integrative studies is essential for promoting sustainable coexistence between wildlife and human activities.

Acknowledgements

Our main thanks go to Dr. P. Mirski from University of Bialystok (Poland) for his help in preparing this publication. The authors are grateful to the following people who participated in the work on collecting Barn Owl pellets from different sites over Georgia: P.Afonin, O.Bystritskaya, S.Bystritsky, A.Evseeva, V.Fenchuk, O.Ilyina, E.Kliuzheva, M.Kliuzhev, V.Kochetkov, A.Kuzmenkova, N.Paposhvili. Special thanks to L.Mumladze for opportunity to work with zoological collection in Ilya State University, to Ç.Şekercioğlu and Aras Bird Ringing Station in Türkiye for providing the data on certain birds mean mass and to K.Deoniziak for the access to the bird skull collection in the University of Bialystok.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The preparation of this paper was carried out within the framework of the EU4Belarus SALT II program and funded by the European Union.

Author contributions

DK collected and desiccated Barn Owl pellets, conceived an idea of the paper and wrote the manuscript; ZJ was responsible for the collaboration between Ilia State University (Georgia) and University of Bialystok (Poland) to implement the project.

Author ORCIDs

Denis Kitel https://orcid.org/0000-0002-5298-2611

Zurab Javakhishvili https://orcid.org/0000-0001-8909-1081

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

References

  • Abuladze A (2013) Birds of Prey of Georgia. Materials towards a Fauna of Georgia. Issue VI. Tbilisi, 218 pp.
  • Aulagnier S, Haffner P, Mitchell-Jones AJ, Moutou F, Zima J (2009) Mammals of Europe, North Africa and the Middle East. London, Bloomsbury Wildlife, 272 pp.
  • Belik VP (2024) Current expansion of the Barn Owl (Tyto alba) (Tytonidae, Aves) in Northern Eurasia. Biological Bulletin of the Russian Academy of Sciences 51: 3154–3168. https://doi.org/10.1134/S1062359024701930
  • Bontzorlos VA, Peris SJ, Vlachos CG, Bakaloudis DE (2005) The diet of Barn Owl in the agricultural landscapes of central Greece. Folia Zoologica 54(1–2): 99–110.
  • Bontzorlos V, Cain S, Leshem Y, Spiegel O, Motro Y, Bloch I, Cherkaoui SI, Aviel S, Apostolidou M, Christou A et al. (2024) Barn Owls as a nature-based solution for pest control: A multinational initiative around the Mediterranean and other regions. Conservation 4: 627–656. https://doi.org/10.3390/conservation4040039
  • Bose M, Guidali F (2001) Seasonal and geographic differences in the diet of Barn Owl in an agro-ecosystem in northern Italy. Journal of Raptor Research 35(3): 240–246.
  • Brown R, Lees D, Ferguson J, Lawrence M (2021) Tracks and Signs of the Birds of Britain and Europe. 3rd ed. Bloomsbury Publishing, London, 416 pp.
  • Budagashvili N, Javakhishvili Z (2024) The birds of Georgia: an updated checklist using citizen science platforms. Sandgrouse 46: 2–29. [Accessed: 20 September 2025]
  • Bukhnikashvili A (2004) On cadastre of small mammals (Insectivora, Chiroptera, Lagomorpha, Rodentia) of Georgia. Tbilisi, 136 pp. [In Russian]
  • Bukhnikashvili A, Kandaurov A, Sheklashvili G, Natradze I (2023) All records of rodents (Mammalia, Rodentia) and hares (Mammalia, Lagomorpha) in Georgia from 1855 through to 2022. Biodiversity Data Journal 11: e108740. https://doi.org/10.3897/BDJ.11.e108740
  • Bukreev SA (2003) The data on the breeding of Barn Owl on the Caucasus. Strepet 2: 80–81. [In Russian]
  • Coşkun Y, Kaya A (2013) Additional records of Suncus etruscus (Savi, 1822) (Mammalia: Soricidae) from Southeastern Anatolia, Turkey. IUFS Journal of Biology 72(2): 31–35.
  • Çolak E, Sözen M, Özkurt Ş (1998) A study on taxonomic status of Microtus subterraneus and Microtus majori in Turkey. Turkish Journal of Zoology 22: 119–129.
  • Çolak E, Yiğit N, Sözen M, Çolak R, Özkurt Ş, Kankiliç T (2006) The morphological analysis of Mus domesticus and Mus macedonicus in Turkey. Turkish Journal of Zoology 30(3): 309–317.
  • Darvish J, Mohammadi Z, Ghorbani F, Mostafavi E (2014) Morphometric characterisation of the Eastern Broad-toothed Field Mouse Apodemus mystacinus from Zagros Mountains, Iran. Acta Zoologica Bulgarica 66(4): 461–468.
  • Galvez RA, Gavashelishvili L, Javakhishvili Z (2005) Raptors and Owls of Georgia. GCCW & Buneba Print Publishing, Tbilisi, 128 pp.
  • Gurney JH (1876) Barn Owl and Shrew. Zoologist 2nd Series 11: 4871.
  • Hendrix P, Verdonschot P (2021) Observations on the growth of Dytiscus dimidiatus larvae reared on single or multiple prey species. Entomologische Berichten 81(4): 153–159.
  • Kandaurov A, Bukhnikashvili A, Sheklashvili G, Natradze I (2023) The occurrence of insectivores (Mammalia, Eulipotyphla) in Georgia from 1864 through to 2022. Biodiversity Data Journal 11: e106256. https://doi.org/10.3897/BDJ.11.e106256
  • Kitel D, Songulashvili Z, Javakhishvili Z (2025) Expansion of the Barn Owl Tyto alba in Georgia. Sandgrouse 47: 209–218. [Accessed: 20 September 2025]
  • Kryštufek B, Vohralík V (2001) Mammals of Turkey and Cyprus. Volume 1: Insectivora. University of Primorska, Koper, 140 pp.
  • Kryštufek B, Vohralík V (2005) Mammals of Turkey and Cyprus. Volume 2: Rodentia I: Sciuridae, Dipodidae, Gliridae, Arvicolinae. University of Primorska, Koper, 292 pp.
  • Kryštufek B, Vohralík V (2009) Mammals of Turkey and Cyprus. Volume 3: Rodentia II: Cricetinae, Muridae, Spalacidae, Calomyscidae, Capromyidae, Hystricidae, Castoridae. University of Primorska, Koper, 372 pp.
  • Kryštufek B, Vohralík V, Obuch J (2009) Endemism, vulnerability and conservation issues for small terrestrial mammals from the Balkans and Anatolia. Folia Zoologica 58(3): 291–302.
  • Maglakelidze S, Bukhnikashvili A, Sheklashvili G, Natradze I, Kandaurov A, Mumladze L (2024) Exploring small mammal diversity in Georgia (Sakartvelo) through DNA barcoding. Biologia 79: 2789–2803. https://doi.org/10.1007/s11756-024-01742-1
  • Meresman Y, Ribak G (2020) Elastic wing deformations mitigate flapping asymmetry during manoeuvres in rose chafers (Protaetia cuprea). Journal of Experimental Biology 223(24): jeb225599. https://doi.org/10.1242/jeb.225599
  • Mikusek R (Ed.) (2005) Methods of investigation and conservation of owls. FWIE, Kraków. [In Polish]
  • Milana G, Luiselli L, Amori G (2018) Forty years of dietary studies on Barn Owl (Tyto alba) reveal long-term trends in diversity metrics of small mammal prey. Animal Biology 68(2): 129–146. https://doi.org/10.1163/15707563-17000095
  • Mohammadi Z, Darvish J, Ghorbani F, Mostafavi E (2014) First record of the Caucasus field mouse Apodemus ponticus from Iran. Biodiversity Journal 5(4): 475–480.
  • Mumladze L, Japoshvili B, Anderson EP (2019) Faunal biodiversity research in the Republic of Georgia: a short review of trends, gaps, and needs in the Caucasus biodiversity hotspot. Biologia 75(1): 1385–1397. https://doi.org/10.2478/s11756-019-00398-6
  • Natradze I, Bukhnikashvili A, Sheklashvili G, Mumladze L (2023) Bats of Georgia – an occurrence dataset from 1835 through 2022. Biodiversity Data Journal 11: e103181. https://doi.org/10.3897/BDJ.11.e103181
  • Okulova NM, Bogdanov AS, Baskevich MI et al. (2018) [Skull sizes and proportions in west-Palearctic wood mice (Sylvaemus) from eastern Europe. 1. Interspecific variability]. Zoological Journal 94(11): 1418–1433. https://doi.org/10.1134/S0044513418110065 [In Russian]
  • Panteleev AV (2005) Identification of families of passerine birds by the distal part of the ulna. Russian Ornithological Journal 14 (express issue 304): 1033–1039. [In Russian]
  • Poprach K (2010) The Barn Owl. TYTO, Nenakonice, Czech Republic.
  • Pucek Z (1981) Keys to Vertebrates of Poland: Mammals. Warsaw: PWN – Polish Scientific Publishers.
  • Riegert J, Šindelář J, Zárybnická M, Horáček I (2011) Large-scale spatial patterns of small mammal communities in the Mediterranean region revealed by Barn Owl diet. Scientific Reports 11: 4985. https://doi.org/10.1038/s41598-021-84683-y
  • Sebele L, Mundy P, Fritz H, Guerbois C (2022) Perceptions on Barn Owls and their use in rodent control: a case study of Hwange District. Tropical Conservation Science 15: 1–12. https://doi.org/10.1177/19400829221120175
  • Shlyakhtin GV, Tabachishin VG, Yermokhin MV (2019) Seasonal diet variations of the Sand Lizard (Lacerta agilis) in the Northern Lower-Volga Region. Povolzhskiy Journal of Ecology 3: 396–401. https://doi.org/10.35885/1684-7318-2019-3-396-401 [In Russian]
  • Tembotova FA (2015) [Mammals of Caucasus and the Seas Washing it: A Guide]. Scientific Publishing Partnership KMK, Moscow, 352 pp. [In Russian]
  • Topaloudis A, Cumer T, Lavanchy E, Ducrest AL, Simon C, Machado AP, Paposhvili N, Roulin A, Goudet J (2025) The recombination landscape of the Barn Owl, from families to populations. Genetics 229(1): iyae190. https://doi.org/10.1093/genetics/iyae190
  • Vale-Gonçalves HM, Barros P, Braz L, Cabral JA (2015) The contribution of Barn Owl feeding ecology to confirm bat species occurrence in north Portugal. Barbastella 8(1): 22–26. https://doi.org/10.14709/BarbJ.8.1.2015.05
  • Vinogradov BS, Gromov IM (1952) Rodents of the Fauna of USSR. Academy of Sciences of the USSR, Moscow–Leningrad. [In Russian]
  • Yiğit N, Çolak E, Sözen M, Özkurt Ş (1998) The taxonomy and karyology of Rattus norvegicus and Rattus rattus in Turkey. Turkish Journal of Zoology 22: 203–212.
  • Zaitsev MV, Voyta LL, Sheftel BI (2014) [The mammals of Russia and adjacent territories. Lipotiphlans]. Nauka, St.Petersburg, 390 pp. [In Russian]
  • Zagorodniuk IV (2002) Taxonomic revision and diagnostics of the rodent genus Mus from Eastern Europe. News of Biosphere Reserve “Askania Nova” 4: 130–140. [In Russian]
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