# Golden jackal expansion in northernmost Europe: records in Finland

Ilpo Kojola, Heikki Henttonen, Samuli Heikkinen, Nathan Ranc

<figure><img src="image_1.png" /></figure>

## To cite this version:

Ilpo Kojola, Heikki Henttonen, Samuli Heikkinen, Nathan Ranc. Golden jackal expansion in northernmost Europe: records in Finland. Mammalian Biology: Zeitschrift für Säugetierkunde, 2023, 10.1007/s42991-023-00382-3. hal-04278630

HAL Id: hal-04278630

https://hal.inrae.fr/hal-04278630

Submitted on 10 Nov 2023

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_2.png" /></figure>
<figure><img src="image_3.png" /></figure>

<figure><img src="image_4.png" /></figure>

# Golden jackal expansion in northernmost Europe: records in Finland

Ilpo Kojola<fnref n="1" /> ID, Heikki Henttonen<fnref n="2" /> ID, Samuli Heikkinen<fnref n="3" /> ID, Nathan Ranc<fnref n="4" /> ID

Received: 20 January 2023 / Accepted: 2 October 2023

© The Author(s) 2023

## Abstract

The Eurasian golden jackal (*Canis aureus*) has been rapidly expanding its distribution range in Europe. Whether jackals will be able to adapt to new environmental conditions in northern Europe remains largely unresolved. Herein we provide additional evidence for the species' ability to colonize northern environments by presenting the new records of golden jackal occurrence in Finland. During 2018–2022, golden jackals were recorded at six localities of which one located in southern, four in central and one in northern Finland. We drafted potential main routes of dispersal movements by jackals in northern Europe, and discuss ecology and management of this newly colonizing species in Finland.

**Keyword** Mesocarnivore · Range expansion · Long-distance dispersal

The Eurasian golden jackal (*Canis aureus*) has been rapidly expanding its distribution range in Europe (Trouwborst et al. 2015; Krofel et al. 2017; Hatlauf et al. 2021; Ranc et al. 2022). Native to Europe but initially limited to small areas along the Mediterranean and Black Sea coasts of southeastern Europe, golden jackals have undergone a continental-scale range expansion during the twentieth century, possibly accelerating during the last 20 years (Spassov 1989; Krofel et al. 2017). The scale and spatiotemporal patterns of golden jackal expansion resembles that of two American canid mesocarnivores—the coyote (*Canis latrans*) (Hody and Kays 2018) and the crab-eating fox *Cerdocyon thous*. In the case of golden jackals (and of coyotes), strong evidence exists for the role of historical wolf decline in the species expansion (Krofel et al. 2017; Newsome et al. 2017), yet other mechanisms (e.g., changes in land use, in management practices

and global warming) may be playing important roles as well (Šálek et al. 2014; Rutkowski et al. 2015; Cunze and Klimpel 2022). The species now reproduces throughout Southeastern and Central European countries, with dispersers reaching far into western and northern Europe (Ranc et al. 2022). Up to date, golden jackals have been reported in 34 European countries, 11 of which have been newly colonized since 2010 (Hatlauf et al. 2021; Sørensen and Lindsø 2021).

The golden jackal is a highly adaptable generalist species, characterized by a large diet breadth, with small mammals and scavenging being consistently important food sources (Lanszki et al. 2022). Jackals use and breed in very diverse environmental conditions, with highest densities observed in hetoregenenous agricultural environments and wetlands (Šálek et al. 2014). The species also appears particularly successful in human-dominated environments (Ćirović et al. 2016; Lanszki et al. 2018; Fenton et al. 2021). Vagrants are capable of long-distance dispersal, including in highly anthropized landscapes (Rutkowski et al. 2015; Lanszki et al. 2018), suggesting that most of the continent could be colonized in the near future. The rapid range expansion and the regular occurrence of vagrants far from the known core distribution provides evidence of high dispersal capacity; however, the dispersal ecology of the species remains largely unknown (Lanszki et al. 2018).

Whether golden jackals will be able to adapt to new environmental conditions, especially in the northern countries, is unresolved. However, the recent establishment of a growing population in Estonia (although limited to coastal areas), suggests that the species could indeed thrive in Boreal

Handling editor:Carlos Rouco.

<figure><img src="image_5.png" /></figure>

Ilpo Kojola
ilpo.kojola@luke.fi

<footnote n="1">¹ Natural Resources Institute Finland (Luke), Ounasjoentie 6, 96200 Rovaniemi, Finland</footnote>

<footnote n="2">² Natural Resources Institute Finland (Luke), Latokartanonkaari 9, 00790 Helsinki, Finland</footnote>

<footnote n="3">³ Natural Resources Institute Finland (Luke), Paavo Havaksentie 3, 90570 Oulu, Finland</footnote>

<footnote n="4">⁴ Université de Toulouse, INRAE, CEFS, Castanet-Tolosan, France</footnote>
ecosystems (Männil and Ranc 2022) despite the ecological conditions there significantly differ from those in the source populations of Southeastern and Central Europe, and of the Caucasus. In addition, several dispersing individuals have very recently been noticed as far as the European Subarctic and Arctic regions, in Norway and in Russia (Sørensen and Lindsø 2021; Rykov et al. 2022). Herein we provide additional evidence for golden jackal's ability to colonize northern environments by presenting the records of jackal presence in Finland and discuss potential dispersal routes into Finland by taking into account the geographic location of occurrences. Furthermore, we discuss the future of golden jackal and its management in Finland.

The monitoring of large carnivore populations in Finland is extensively based on opportunistic point observations (sightings, tracks, roadkills) by hunters, foresters and naturalists; then verified and recorded by approximately 2000 volunteer large carnivore contact persons distributed all over the country (Kojola et al. 2018). Persons for this network are nominated in local game management districts (n=280) and educated by Finnish Wildlife Agency and Natural Research Institute Finland. In addition, thousands of private trail cameras used for local wildlife monitoring are maintained by hunters, providing an additional, important source of opportunistic data. Suspected trail camera images of golden jackals are sent to wildlife experts working for Natural Resources Institute Finland for assessment. Hunters, naturalists and large carnivore contact persons are encouraged to send images that could be jackal images. The images sent are mostly images of fox or wolf. After this first filtering during which the authenticity is evaluated by Finnish specialists, potential golden jackal records are evaluated by several European golden jackal experts. Our data here comprise records from six different locations, originating from different data sources—trail camera images (three), picture from photographer (one), recovery of dead animal (one) and multiple data from the presence of one possibly resident individual (snow tracks, vocal records; see Table 1 and Fig. 1 for record summary and location). In this article, we present a photographic document (Fig. 1), the date, the type

of the document and surrounding environment (Table 1), for each record.

We detail below the records in their chronological order. The first verified record of a golden jackal in Finland dates of August 5th 2018, when an individual was photographed by a trail camera within a house yard near Kajaani, in east-central Finland. Second, a few months later, on January 16th 2019, a single golden jackal was photographed by another trail camera at feeding site in a coniferous forest near Reisjärvi, in central Finland. Third, on July 23rd 2019, a jackal moving along the road side was photographed from car in Rautavaara (a municipality neighboring Kajaani; i.e. 90 km from the first record). Fourth, the presence of one individual was repeatedly reported, between November 22nd 2020 and April 16th 2022, by a trail camera located at a roe deer feeding station Lumijoki-Siikajoki, west-central Finland. The same individual, probably a male (cf. Table 1), may have also been observed nearby by an ornithologist on September 1st 2022. Fifth, in early March 2022, a jackal was inadvertently trapped (and killed) by a fox-snare in Lapland 120 km north from the Arctic Circle (northernmost record in Finland). The individual was a male that weighed 10 kg and proved out to be in good physical condition based on laboratory investigation by Finnish Food Authority (personal communication). The species identification was confirmed in DNA analysis (J. Aspi et al., unpublished data). The potential relatedness of this animal to Estonian jackals is not known. Sixth and finally, snow tracks and spontaneous vocalizations by a single jackal have been reported in late March 2022, in southeastern Finland, near the Russian border (Fig. 1). Despite the accumulating data (records #4 and #6) and in the absence of additional information on these two areas of repeated occurrences, the most plausible scenario is that an individual has settled in each area. Further efforts should be conducted to ascertain the identity of the individuals and evaluate the unlikely, but possible, occurrence of territorial (reproductive) groups.

Owing to the high dispersal capacity of golden jackals and the opportunistic nature of our data, it is not currently possible to estimate the number of distinct individuals in

<table><caption>**Table 1** Verified records of golden jackal presence in Finland. Locations in Fig. 1</caption><thead><tr><td>Date</td><td>Document</td><td>Number in Fig. 1</td><td>Surrounding environment</td></tr></thead><tbody><tr><td>Aug. 5, 2018</td><td>Trial camera picture</td><td>1</td><td>House-yard</td></tr><tr><td>Jan. 16, 2019</td><td>Trial camera picture</td><td>2</td><td>Forest</td></tr><tr><td>July 23, 2019</td><td>Photo taken by camera</td><td>3</td><td>Roadside</td></tr><tr><td>From Nov. 22 2020 to Sept. 1, 2022</td><td>Trail camera pictures (Nov. 22 2020, Jan 8 2021, April 16 2022), probably a male based on its posture during urination</td><td>4</td><td>Forest, feeding station</td></tr><tr><td>March 1, 2022</td><td>Adult male found dead, captured in a fox-snare (Fig. 1), vocal records earlier in winter</td><td>5</td><td>Forest</td></tr><tr><td>March 27, 2022</td><td>Photos of tracks, vocal records in March 2022</td><td>6</td><td>Forest</td></tr></tbody></table>

<figure><img src="image_6.png" /></figure>
<figure><img src="image_7.png" /><figcaption>**Fig. 1** Verified records of golden jackal presence in Finland (circles), Norway (squares) and northwestern Russia (hexagons) during 2018–2022 and potential movement routes (arrows) from Estonian jackal populations (ovaloid shapes) to Finland and North-Western Russia. Numbers associated to Finnish records refer to numbers in Table 1. Observations outside Finland were obtained from Sørensen</figcaption></figure>

and Lindsø 2021 (Norway) and Rykov et al. 2022 (Russia). Photos: 1 by Tanja Nuotio, 2 by Hanna-Leena Paananen, 3 by Jarmo Korhonen, 4 by Tommi Lainas, 5 by Seppo Muttilainen and 6 by Tiia Kimmo. Shaded areas show wolf territories in 2018–2022 (cf. Mäntyniemi et al. 2022)

Finland. There are thousands of trail cameras maintained in the country, primarily deployed by moose and deer hunters, but also many at baits used for shooting foxes (*Vulpes vulpes*) and raccoon dogs (*Nyctereutes procyanoides*); because golden jackals tend to use anthropogenic food source extensively (Lanzski et al. 2022), and probably even more so during the severe winter climate of Finland, it is likely that the detection probability is relatively high. In addition, we estimate that private owners of trail cameras are likely to report potential records of golden jackals in Finland since there is a general interest to monitor the species both for ecological reasons and potential conflict with human interests, and hunting and nature magazines have extensively published on the species colonization. Owing to the very large effort, albeit opportunistic in nature, we conclude that jackal occurrence is still rare in the country but that the trend is increasing, with even an indication of individuals settling (in at least two separate locations).

The most likely source of jackals observed in Finland is Estonia, where the species bred for the first time in 2012 and has gradually colonized the westernmost coastal areas of the country (Männil and Ranc 2022). The northern locations of Finnish observations might suggest that most jackals have moved into Finland via the isthmus located between the lakes Ladoga and Onega (Fig. 1), while the Megapolis of St. Petersburg may block the route through the isthmus between the Finnish Gulf and Lake Ladoga. If so, it could explain why most of the jackal observations are relatively north in Finland. The Russian observations from Arkhangelsk and Vologda regions (Rykov et al. 2022) tend to corroborate this scenario. However, given the species' high ecological flexibility, their arrival through the St. Petersburg Isthmus cannot entirely be ruled out. Although Estonia is the most likely origin of jackals recorded in Finland, Norway and Northwestern Russia, some long-distance dispersers could also potentially originate in Central Europe, or even in the Caucasus (Rykov et al. 2022). In Russia the golden jackal is present and breeding in southernmost segments of the country and
during the 2000s, four individuals have been killed in North-
western Russia. In addition to jackals reported by Rykov
et al. 2022 (Leningrad region 2007, Moscow region 2016,
Archangelsk region 2021) and a newspaper article reports a
jackal kill with an excellent photo at Belozersk in Vologda
region, 400 km east of St. Petersburg in 2021 (personal com-
munication with Alexei V. Abramov, Zoological Museum
of St. Petersburg). However, our understanding of the status
of golden jackals in Russia remains partial. In this context,
obtaining data on the distribution of golden jackals in Rus-
sia is critical to understand and forecast future colonization
of Finland.

Very little is known of golden jackal ecology in northern
environments (Männil and Ranc 2022), and even less so
in Finland. In such environments, it is likely that winter
represents the most significant constrain for jackal survival.
Observations in Finland indicate that one jackal has survived
over two consecutive winters (record 4), and two others (5
and 6) possibly one winter. Information provided by local
hunters, concerning the two northernmost individuals
(records 4 and 5) indicates that these individuals have been
highly dependent on anthropogenic food—regularly visiting
feeding stations where oat was provided for roe deer in
winter (record 4), and a location where local professional
fishermen clean fish (record 5). Southern Finland is
probably the most suitable habitat for golden jackals as the
landscape is more heterogeneous, with a higher proportion
of agricultural lands, and more diverse and abundant food
sources—high small mammal density, abundant prey such
as European hare (Lepus europaeus), roe deer (Capreolus
capreolus) and white-tailed deer (Odocoileus virginianus),
as well as a high density of mammalian and avian road kills.
Cunze and Klimpel (2022) predicted that the changes in
climate would create suitable environments for jackal west
and north from its present distribution range but Finland
was not regarded as a potential new area in their projections.
Given observed spatiotemporal patterns in Central Europe
(e.g., Slovenia), it can take several decades before sporadic
occurrences lead to an actual population growth similar to
what is observed in the species' core areas of southeastern
Europe (Krofel et al. 2023).

In multiple countries, golden jackal presence seems
affected by the occurrence of larger, dominant grey wolves
(Canis lupus) (Krofel et al. 2017). In particular, the Balkan
region experienced a rapid increase of golden jackal
population following the decrease and fragmentation of
historic wolf populations (Krofel et al. 2017). In Estonia,
golden jackals seem to avoid regions which are inhabited
by resident wolf packs (Männil and Ranc 2022). In Finland,
only one record (record 4) was made in the periphery of
a wolf breeding territory, while the five others are located
outside wolf territory boundaries. So far no records of
golden jackals have been reported from Southwestern

Finland, where food sources is certainly highest and snow
cover duration shortest but which harbors the highest density
of permanent wolf packs and territory-marking pairs of
wolves in the country (Mäntyniemi et al. 2022). The spatial
pattern of future occurrence of golden jackals may allow us
to evaluate the influence of wolf on the species' colonization
pattern in Finland. Future studies should evaluate how
golden jackal will influence animal communities in Boreal
environments. For example, jackals may locally influence
the reproductive success of ground-nesting birds but also
increase mortality of red fox and raccoon dog through
intraguild competition and predation.

In Finland, the golden jackal is currently a fully protected
species by the law of environmental protection. Whenever
current expansion will lead to an increasing breeding
population, the status of jackals might change into a game
animal, which enables managers to approve licenses more
readily for removal of damage-causing individuals. The
examples from Estonia (Männil & Ranc 2022) and other
European countries (Trouwborst et al. 2015; Hatlauf et al.
2021) indicate that jackal populations can sustain very high
pressure. The benefits of such management in terms of jackal
population regulation and human-jackal conflicts remain
to be studied. Whenever models for potential harvest are
based on reliable data on the size and demographics of the
population, golden jackal populations can be successfully
conserved when they have a legal status of a game animal.

Golden jackal dispersers are increasingly being recorded
in Finland, especially in the central part of the country.
In southern Fennoscandia, where the potential for future
expansion and establishment of reproductive units is greatest,
habitat fragmentation and geography (Baltic Sea, big lakes)
may constrain dispersal routes. Thus far, all recorded
jackals appear isolated and many are closely associated with
anthropogenic food during winters. The ability for jackals to
establish reproductive groups will condition the long-term
potential of the species in Fennoscandia.

**Acknowledgements** We are grateful to Marja Isomursu, Tiia Kimmo, Jarmo Korhonen, Tommi Lainas, Seppo Muttilainen and Tanja Nuotio for pictures and background information about the Finnish records of golden jackals.

**Author contributions** Study concept and design: IK, HH, SH and NR; data collection: IK and HH; original draft preparation: IK and HH; writing—review and editing: IK, HH, SH and NR.

**Funding** Open access funding provided by Natural Resources Institute Finland.

**Data availability** Data are available from authors.

**Declarations**

**Conflict of interest** The authors declare no competing interests.
**Open Access** This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

## References

Ćirović D, Penezić A, Krofel M (2016) Jackals as cleaners: Ecosystem services provided by a mesocarnivore in human-dominated landscapes. *Biol Conserv* 199:51–55. https://doi.org/10.1016/j.biocon.2016.04.027

Cunze S, Klimpel S (2022) From the Balkan towards Western Europe: Range expansion of the golden jackal (*Canis aureus*) A climatic niche modeling approach. *Ecol Evol* 12:e9141. https://doi.org/10.1002/ece3.9141

Fenton S, Moorcroft PR, Ćirović D, Lanszki J, Heltai M, Cagnacci F, Breck S, Bogdanović N, Pantelić I, Ács K, Ranc N (2021) Movement, space-use and resource preferences of European golden jackals in human-dominated landscapes: insights from a telemetry study. *Mamm Biol* 101:619–630. https://doi.org/10.1007/s42991-021-00109-2

Hatlauf J, Bayer K, Trouwborst A, Hackländer K (2021) New rules or old concepts? The golden jackal (*Canis aureus*) and its legal status in Central Europe. Eur J Wildl Res 67:25. https://doi.org/10.1007/s10344-020-01454-2

Hody JW, Kays R (2018) Mapping the expansion of coyotes (*Canis latrans*) across North and Central America. *ZooKeys* 759:81–97

Kojola I, Heikkinen S, Holmala K (2018) Balancing costs and confidence: volunteer-provided point observations, GPS telemetry and the genetic monitoring of Finland's wolves. Mammal Res 63:415–423. https://doi.org/10.1007/s13364-018-0371

Krofel M, Giannatos G, Ćirović D, Stoyanov S, Newsome TM (2017) Golden jackal expansion in Europe: a case of mesopredator release triggered by continent-wide wolf persecution? *Hystrix, Ital J Mamm* 28:1–15. https://doi.org/10.4404/hystrix-28.1-11819

Krofel M, Berce M, Berce B, Kryštufek S, Lamut S, Tarman J, Fležar U (2023) New mesocarnivore at the doorstep of Central Europe: historic development of golden jackal (*Canis aureus*) population in Slovenia. Mammal Res 68:329–339. https://doi.org/10.1007/s13364-023-00686-2

Lanszki J, Schally G, Heltai M, Ranc N (2018) Golden jackal expansion in Europe: first telemetry evidence of a natal dispersal. *Mamm Biol* 88:81–84. https://doi.org/10.1016/j.mambio.2017.11.011

Lanszki J, Hayward MW, Ranc N, Zalewski A (2022) Dietary flexibility promotes range expansion: The case of golden jackals in Eurasia. J Biogeogr 49:993–1005. https://doi.org/10.1111/jbi.14372

Männil, Ranc N (2022) Golden jackal (*Canis aureus*) in Estonia: development of a thriving population in the boreal ecoregion. Mammal Res 67:245–250. https://doi.org/10.1007/s13364-021-00615-1

Mäntyniemi S, Helle I, Kojola (2022) Eur J Wildl Res 68:70. https://doi.org/10.1007/s10344-022-01615-5

Newsome TM, Greenville AC, Ćirović D, Dickman CR, Johnson CN, Krofel M, Letnic M, Ripple WJ, Ritchie EG, Stoyanov A, Wirsing AJ (2017) Top predators constrain mesopredator distributions. Nat Commun 8:15469

Ranc N, Acosta-Pankov I, Balys V, Bučko J, Cirovic D, Fabijanić N, Filacorda S, Giannatos G, Guimaraes N, Hatlauf J, Heltai M, Ionescu O, Ivanof G, Jansman H, Kowalchyk R, Krofel M, Kutal M, Lanszki J, Lapini L, Männil P, Melovski D, Migli D, Molinari P, Olsen K, Ozolinš J, Pavanello M, Šálek M, Selanec I, Stojanov A, Stoyanov S, Sunde P, Szabó L, Reinhardt I, Trace A, Trbojevic I, von Arx M, Yakovlev Y, Zimmermann F (2022) Distribution of large carnivores in Europe 2012–2016: Distribution map for Golden Jackal (*Canis aureus*). Zenodo. https://doi.org/10.5281/zenodo.6382216

Rutkowski R, Krofel M, Giannatos G, Ćirović D, Männil P, Volokh AM, Lanszki J, Heltai M, Szabó L, Banea OC, Yavruyan E,HRarapetyan V, Kopiliani N, Miliou A, Tryfonopoulos GA, Lymberakis P, Penezić A, Pakeltytė G, Schecka E, Bogdanowics W (2015) A European concern? genetic structure and expansion of golden jackals (*Canis aureus*) in Europe and the Caucasus. PLoS ONE 10:1–22. https://doi.org/10.1371/journal.pone.0141236

Rykov AM, Kuznetsova AS, Tirronen KF (2022) The first record of the golden jackal (*Canis aureus* Linnaeus, 1758) in the Russian Subarctic. Polar Biol 45:965–970. https://doi.org/10.1007/s00300-022-03037-0

Šálek M, Červinka J, Banea OC, Krofel M, Ćirović D, Selanec I, Penzeciz A, Grill S, Riegert J (2014) Population densities and habitat use of the golden jackal (*Canis aureus*) in farmlands across the Balkan Peninsula. Eur J Wildl Res 60:193–200. https://doi.org/10.1007/s10344-013-0765-0

Sørensen OJ, Lindsø LK (2021) The golden jackal *Canis aureus* detected in Norway – management challenges with naturally dispersed species new to the country (Original title: Gullsjakal påvist i Norge – forvaltningsutfordringer ved nye arter i landet). Fauna 74:74–87

Spassov N (1989) The position of jackals in the Canis genus and life-history of the golden jackal (*Canis aureus* L.) in Bulgaria and on the Balkans. Hist Nat Bulg 1:44–56

Trouwborst A, Krofel M, Linnell JDC (2015) Legal implications of range expansions in a terrestrial carnivore: the case of the golden jackal (*Canis aureus*) in Europe. Biodivers Conserv 24:2593–2610. https://doi.org/10.1007/s10531-015-0948-y

**Publisher's Note** Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.