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The unique case of marking behaviour in juvenile lynx

The unique case of marking behaviour in juvenile lynx Active scent marking with urine has been described in a number of mammal species, including felids. In Eurasian lynx (Lynx lynx), scent marking plays a role in intra-sexual competition and territory defence as well as in attracting sexual partners during the mating season. Marking is most frequent during the mating season and least frequent during the period when females give birth and lactate. Males generally mark more frequently than females and resident animals mark more frequently than dispersers. Juveniles have never been recorded actively marking. Here, however, we present a well-documented case of an actively marking juvenile lynx. Lynx females Koka and Baronka were born in 2019 to mother Nela, who disappeared in December 2019. Nela’s territory stayed vacant, with her juvenile daughters being the only females recorded there. On the th 30 March 2020, during the mating season, Koka was recorded on a camera trap video actively marking with urine, rubbing her face and again marking with urine on a rock. On the following day, she again marked with urine on the same marking site. After that, Koka left the area and the former territory of Nela was taken over by Baronka. In the BBA lynx population, juvenile lynx females were documented to successfully reproduce. This poses the question of whether their adult-like mark- ing behaviour should be explained in the context of early reproduction, or if it has other reasons. Neither Koka nor Baronka successfully reproduced with the local male during their first year of life, but that does not mean they did not try to attract him by marking. Also, juveniles’ competition over the territory left vacant by their mother is a feasible explanation. We sug- gest that high population turnover may result in unusual social situations, with juveniles consequently performing unusual behaviour. This does not have to be related to early reproduction. Keywords Eurasian lynx · Lynx lynx · Scent marking · Behaviour · Juvenile · Orphan · Bohemian-Bavarian-Austrian lynx population · BBA Article Mammals are complex social creatures, who are known to communicate visually, acoustically, and also by odours * T. Mináriková (Halpin 1986; Kean et al. 2017; Campbell-Palmer and Rosell tereza.minarikova@alkawildlife.eu 2011). These so-called “social odours “ may carry informa- ALKA Wildlife, Lidéřovice 62 , Dačice, Lidéřovice 380 01, tion about the animal's sex, reproductive status, individual Czech Republic identity, territory ownership, dominance, and health status Faculty of Environmental Sciences, Czech University (Johson 1973; Brennan and Kendrick  2006; Wyatt 2003; of Life Sciences Prague, Kamýcká 1176, Prague CZ-16521, Vogt et al. 2016). They consist of complex mixtures of mol- Czech Republic ecules sensed by chemosensory systems and have important Šumava National Park Administration, 1.máje 260, influences on a variety of behaviours that are vital for repro- Vimperk 385 01, Czech Republic ductive success, such as parent–offspring attachment, mate Faculty of Forestry and Wood Sciences, Czech University choice, and territorial marking (Johson 1973, Brennan and of Life Sciences Prague, Kamýcká 1176, Prague CZ-16521, Kendrick 2006; Wyatt 2003; Vogt et al. 2014). In many mam- Czech Republic mal species, the development of scent glands and the offset Hnutí DUHA Šelmy, Dolní náměstí 38, Olomouc 779 00, of marking behaviour corresponds with the sexual maturity Czech Republic of animals (Johnson 1973). In other cases, however, the trig- Agentura ochrany přírody a krajiny ČR, regionální gers of scent marking are still unclear (Ewer 1968; Allen pracoviště Jižní Čechy, Nám. Přemysla Otakara II. č. 34, et al. 2016). České Budějovice 370 01, Czech Republic Vol.:(0123456789) 1 3 acta ethologica Active scent marking with faeces and urine has been lynx juveniles have never been observed actively marking. described in several mammal species (reviewed in Gosling and Juvenile scent marking has not been previously recorded Roberts 2001a, b), including felids (Mellen 1993; Mohorović in the Bohemian-Bavarian-Austrian lynx population, either and Krofel 2020; Asa 1993; Vogt et al. 2014; Sæbø 2007; (own unpublished data). Stromková 2020). Despite intensive research on this topic in When it comes to the social status of the animal, both recent years, the basic information on scent marking is lacking residents and dispersing lynx were found to scent mark, for 23% of all felid species, and information on 21% of other however with different frequencies (Vogt et al. 2014; Sæbø felid species comes solely from a single study in captivity (Allen 2007; Stromková 2020). Resident animals are found to mark et al. 2016). the most, probably in order to defend their territories (Vogt In tigers, females mark intensively prior to oestrus but not et al. 2014; Sæbø 2007; Stromková 2020). In Scandinavia, during it, and males mark most frequently when females are adult lynx placed scent marks more often at the borders than in oestrus (Smith et al. 1989). Marking is the most intensive in the centers of their home ranges (Sæbø 2007). In other when tigers are establishing territories (Smith et al. 1989). In regions, lynx females were observed to mark more often pumas, scent communication at community scrapes is used in core areas, where they spent most of their time (Gosling by females to select dominant resident males to mate with, and Roberts 2001b; Wyatt 2003; Hucht-Ciorga 1988). Lynx while males advertise themselves for mating and try to deter marking behaviour is however not only related to intra-sexual competitors (Allen et al. 2014; Allen et al. 2015). Dependent competition and territory defence, but also plays an important kittens do occasionally mark at community scrapes, but gen- role in attracting sexual partners during the mating season erally do not exhibit this behaviour, probably because they (Vogt et al. 2014). have no need to establish territories nor advertise availability The territorial behaviour of animals is generally consid- to mate (Allen et al. 2021). ered to be determined mainly by food availability and dis- Small cats (Felinae) exhibit strong uniformity in both tribution, which has a major impact on animal fitness, espe- scent marking and social behaviours (Mellen 1993). Repro- cially in females (Carpenter and MacMillen 1976; Maher ductively active animals mark more frequently than others, and Lott 2000; McLoughlin et al. 2000; Adams 2001; Borger especially during oestrus (Mellen 1993). In a female black- et  al. 2008). In Eurasian lynx, prey availability strongly footed cat, the marking frequency peaked before the mat- affects territory size (Mengüllüoğlu et al. 2021; Aronsson ing season and was absent during the period of rearing of et al. 2016) and reproductive success in females (Yom-Tov young, thus advertising female reproductive status (Molteno et al. 2010; Nilsen et al. 2010). Lynx territoriality is also et al. 1998). In the bobcat, marking is primarily exhibited strongly dependent on the animal's social status, with age, by adults (Bailey 1974), although juveniles were found to sex, reproductive status, population density, and spatial distri- mark from 5 months of age (Wassmer et al. 1988). In both bution affecting the size and even the very existence of lynx bobcat and Iberian lynx, scent marking seems to mainly play territories (Mengüllüoğlu et al. 2021; Aronsson et al. 2016). a role in territory defence, while its connection to mating In the Bohemian-Bavarian-Austrian lynx population, behaviour is not yet clear (Wassmer et al. 1988; Bailey 1974; the mean annual home range sizes are 438 km for adult Beltran et al. 1987; Barja et al. 2005). males and 278 km for adult females (own unpublished data, The Eurasian lynx (hereafter: lynx) is known to scent Belotti et al. 2012; Magg et al. 2016; Wölfl et al. 2001). mark with urine at visually conspicuous sites such as rocks, Generally, the home range of one male overlaps with those fallen tree trunks, roots, wood piles, wooden cabins, and of one to three females (Mináriková et al. 2020; Wölfl et al. along frequented lynx paths, where it is probable that con- 2020; Engleder et al. 2019, 2021). Home ranges of adult specifics will find the scent mark (Allen et al. 2017; Sæbø lynx of the same sex may partially, but never entirely, over- 2007; Vogt et al. 2014; Stromková 2020; Krofel et al. 2017). lap (Mináriková et al. 2020; Wölfl et al. 2020; Engleder et al. Faeces are also frequently found exposed at marking sites, 2019, 2021). although at resting sites or nearby kill lynx usually hide Here we present a unique, well-documented case of an them (Sæbø 2007; Stromková 2020, own unpublished data). actively marking juvenile lynx. Marking behaviour is not equally distributed in time but is It was recorded in the so-called Bohemian-Bavarian- most frequent during the mating season (January-April) and Austrian (BBA) lynx population, whose distribution area least frequent during the period when females give birth stretches across the borders between the Czech Republic, and lactate (May–July) (Vogt et  al. 2014). Males mark Bavaria, and Austria (Fig. 1). more frequently than females (Vogt et al. 2014; Sæbø 2007; Camera trapping is used as a major lynx monitoring Stromková 2020). Juveniles visit marking sites with their method in all three countries, complemented by snow track- mothers, but according to the three large-scale scent mark- ing and genetic analyses. To space the camera traps evenly ing studies of Alpine, Scandinavian, and Carpathian lynx across the region, we used a 10 km × 10 km grid (ETRS populations (Vogt et al. 2014; Sæbø 2007; Stromková 2020), LAEA 5210 10 km grid) and installed two to eight camera 1 3 acta ethologica Fig.1 Map of Bohemian- Bavarian-Austrian lynx popu- lation. The wider region of Vlachovo Březí forests, where the yearling female Koka was documented, is circled in red trapping sites per grid cell, depending on the area and habi- – “Independent lynx”—lynx older than 1 year, i.e., sub- tat type. Altogether, 131 grid cells were covered by camera adult or adult. trapping in the years 2017–2021 as part of the 3Lynx project – “Resident lynx” lynx staying for minimally 12 months in and following national activities (Mináriková et al. 2020; the same area after becoming independent. Wölfl et al. 2020; Engleder et al. 2021). The time unit for data analysis was lynx year, that is Lynx females B748 Koka and B749 Baronka were born st th the period starting on the 1 May and ending on the 30 in spring 2019 to B718 Nela, a resident female in the Vla- April of the following year, based on the lynx life cycle chovo Březí area (Czech Republic, South Bohemian region, (Mináriková et al. 2020). In the data analysis, the follow- Fig.  1). They were born in the territory of resident male ing categories were used for lynx according to their status: B716 Karlos. Karlos and Nela both disappeared during the lynx year – “Juvenile lynx”—lynx in the first year of life (also 2019 and have not been recorded since then, thus we assume th called “kitten”), recorded from birth till the 30 April they are dead. Karlos’s territory was taken over by the sub- of the following calendar year (0–1 year of age). adult young male B740 Piškot in November 2019. Nela was th – “Subadult lynx”—lynx in the second year of life, last recorded with both her kittens on the 13 December st recorded from the 1 May of the year following the birth 2019, but the exact date of her death remains unknown as th till the 30 April of the next year (1–2 years of age). she could have been missed by camera traps for some time. – “Adult lynx”—lynx older than 2 years. Her territory was not taken over by any subadult or adult 1 3 acta ethologica female. Therefore, both her juvenile daughters were left with the possibility of settling into the vacant territory they already inhabited as kittens. th On the 30 March 2020, during the mating season, Koka was recorded on a video actively marking with urine, rub- bing her face, and again marking with urine on a marking site (Fig. 2). On the following day, she again marked with th urine on the same spot (Fig. 3). On the 29 March this mark- nd ing site was visited by Piškot and on the 2 April it was visited by Baronka, but they were not recorded marking, they both merely inspected it. It was a rock, previously used for marking by resident female Nela, previous resident male Karlos, new resident male Piškot, and later Baronka. rd th Between the 3 and 26 of April 2020, after the end of the mating season, Koka left the area and dispersed 40 km far to the Boletice military training area and then to the Protected Landscape Area Blanský les, where she has lived since July 2020. The former territory of Nela was taken over by Bar- onka, who is till now (December 2022) the resident repro- ducing female lynx here, sharing the area with male Piškot. Lynx females are known to successfully reproduce from the age of 2 years (Breitenmoser-Würsten et al. 2007; Nilsson 2013; Nilsen et al. 2010, 2012; Kvam 1991). Juvenile lynx ovulation was recorded in Norway, but no juvenile pregnancy nor parturition was found (Kvam 1991). The BBA lynx popu- lation is, to the best of our knowledge, the only European lynx population where juvenile lynx females were documented to successfully give birth and raise kittens to independence (Engleder et al. 2019; Palmero et al. 2021, own unpublished data). This poses the question of whether the adult-like mark- ing behaviour of a juvenile female lynx should be explained in the context of early reproduction or if it has other reasons. While the marking behaviour in lynx generally serves both to attract conspecifics of the opposite sex and to mark the territory (Vogt et al. 2014), we cannot assess what was the main purpose of Koka’s behaviour. Neither Koka nor Baronka successfully reproduced with the resident male Piškot during their first year of life but that does not mean they did not try to attract him (or any other male) by mark- ing. Also, juveniles’ competition over the territory left vacant by their mother is a possible explanation for the marking behaviour, with Baronka winning over and displac- ing Koka, who then dispersed to a new area. The BBA lynx population has a high turnover of adult Fig. 2 Juvenile female Koka actively marking with urine, rubbing her th and subadult lynx, similarly to its neighbours – the West- face and again marking with urine on a marking site (the 30 March 2020) ern Carpathian and the Alpine lynx populations (Wölfl et  al. 2020; Duľa et  al. 2021; Breitenmoser-Würsten et al. 2007). Specifically, 23% of adults and 44% of sub- adults vanish every year, especially in the outskirt part lynx presence (Mináriková et al. 2020; Wölfl et al. 2020; Engleder et  al. 2021). This instability may result in of the BBA population’s distribution (Wölfl et al. 2020). This inevitably causes social instability, with territories unusual social situations, like the one presented in this paper, with juveniles consequently performing unusual changing their owners frequently and some areas being temporary without lynx reproduction or entirely without behaviours. 1 3 acta ethologica Funding Open access publishing supported by the National Technical Library in Prague. This study was supported by the CE1001 3Lynx Project, funded by Interreg Central Europe. Data availability The datasets analysed during the current study are available from the corresponding author on reasonable request. Declarations Conflict of interest The authors declare that they have no conflict of interest. Open Access This article is licensed under a Creative Commons Attri- bution 4.0 International License, which permits use, sharing, adapta- tion, 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 Fig. 3 Koka again marking with urine on the same marking site (the otherwise in a credit line to the material. If material is not included in st 31 March 2020) 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://cr eativ ecommons. or g/licen ses/ b y/4.0/ . 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Acta Theriol 46(2):181–194. https:// doi. org/ 10. 4098/ at. arch. 01- 20 Publisher's Note Springer Nature remains neutral with regard to Wyatt TD (2003) Pheromones and animal behaviour (Vol. 626). Cam- jurisdictional claims in published maps and institutional affiliations. bridge University Press, Cambridge. ht tps:/ / doi . or g/ 10. 10 17/ cbo97 80511 615061 1 3 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png acta ethologica Springer Journals

The unique case of marking behaviour in juvenile lynx

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Abstract

Active scent marking with urine has been described in a number of mammal species, including felids. In Eurasian lynx (Lynx lynx), scent marking plays a role in intra-sexual competition and territory defence as well as in attracting sexual partners during the mating season. Marking is most frequent during the mating season and least frequent during the period when females give birth and lactate. Males generally mark more frequently than females and resident animals mark more frequently than dispersers. Juveniles have never been recorded actively marking. Here, however, we present a well-documented case of an actively marking juvenile lynx. Lynx females Koka and Baronka were born in 2019 to mother Nela, who disappeared in December 2019. Nela’s territory stayed vacant, with her juvenile daughters being the only females recorded there. On the th 30 March 2020, during the mating season, Koka was recorded on a camera trap video actively marking with urine, rubbing her face and again marking with urine on a rock. On the following day, she again marked with urine on the same marking site. After that, Koka left the area and the former territory of Nela was taken over by Baronka. In the BBA lynx population, juvenile lynx females were documented to successfully reproduce. This poses the question of whether their adult-like mark- ing behaviour should be explained in the context of early reproduction, or if it has other reasons. Neither Koka nor Baronka successfully reproduced with the local male during their first year of life, but that does not mean they did not try to attract him by marking. Also, juveniles’ competition over the territory left vacant by their mother is a feasible explanation. We sug- gest that high population turnover may result in unusual social situations, with juveniles consequently performing unusual behaviour. This does not have to be related to early reproduction. Keywords Eurasian lynx · Lynx lynx · Scent marking · Behaviour · Juvenile · Orphan · Bohemian-Bavarian-Austrian lynx population · BBA Article Mammals are complex social creatures, who are known to communicate visually, acoustically, and also by odours * T. Mináriková (Halpin 1986; Kean et al. 2017; Campbell-Palmer and Rosell tereza.minarikova@alkawildlife.eu 2011). These so-called “social odours “ may carry informa- ALKA Wildlife, Lidéřovice 62 , Dačice, Lidéřovice 380 01, tion about the animal's sex, reproductive status, individual Czech Republic identity, territory ownership, dominance, and health status Faculty of Environmental Sciences, Czech University (Johson 1973; Brennan and Kendrick  2006; Wyatt 2003; of Life Sciences Prague, Kamýcká 1176, Prague CZ-16521, Vogt et al. 2016). They consist of complex mixtures of mol- Czech Republic ecules sensed by chemosensory systems and have important Šumava National Park Administration, 1.máje 260, influences on a variety of behaviours that are vital for repro- Vimperk 385 01, Czech Republic ductive success, such as parent–offspring attachment, mate Faculty of Forestry and Wood Sciences, Czech University choice, and territorial marking (Johson 1973, Brennan and of Life Sciences Prague, Kamýcká 1176, Prague CZ-16521, Kendrick 2006; Wyatt 2003; Vogt et al. 2014). In many mam- Czech Republic mal species, the development of scent glands and the offset Hnutí DUHA Šelmy, Dolní náměstí 38, Olomouc 779 00, of marking behaviour corresponds with the sexual maturity Czech Republic of animals (Johnson 1973). In other cases, however, the trig- Agentura ochrany přírody a krajiny ČR, regionální gers of scent marking are still unclear (Ewer 1968; Allen pracoviště Jižní Čechy, Nám. Přemysla Otakara II. č. 34, et al. 2016). České Budějovice 370 01, Czech Republic Vol.:(0123456789) 1 3 acta ethologica Active scent marking with faeces and urine has been lynx juveniles have never been observed actively marking. described in several mammal species (reviewed in Gosling and Juvenile scent marking has not been previously recorded Roberts 2001a, b), including felids (Mellen 1993; Mohorović in the Bohemian-Bavarian-Austrian lynx population, either and Krofel 2020; Asa 1993; Vogt et al. 2014; Sæbø 2007; (own unpublished data). Stromková 2020). Despite intensive research on this topic in When it comes to the social status of the animal, both recent years, the basic information on scent marking is lacking residents and dispersing lynx were found to scent mark, for 23% of all felid species, and information on 21% of other however with different frequencies (Vogt et al. 2014; Sæbø felid species comes solely from a single study in captivity (Allen 2007; Stromková 2020). Resident animals are found to mark et al. 2016). the most, probably in order to defend their territories (Vogt In tigers, females mark intensively prior to oestrus but not et al. 2014; Sæbø 2007; Stromková 2020). In Scandinavia, during it, and males mark most frequently when females are adult lynx placed scent marks more often at the borders than in oestrus (Smith et al. 1989). Marking is the most intensive in the centers of their home ranges (Sæbø 2007). In other when tigers are establishing territories (Smith et al. 1989). In regions, lynx females were observed to mark more often pumas, scent communication at community scrapes is used in core areas, where they spent most of their time (Gosling by females to select dominant resident males to mate with, and Roberts 2001b; Wyatt 2003; Hucht-Ciorga 1988). Lynx while males advertise themselves for mating and try to deter marking behaviour is however not only related to intra-sexual competitors (Allen et al. 2014; Allen et al. 2015). Dependent competition and territory defence, but also plays an important kittens do occasionally mark at community scrapes, but gen- role in attracting sexual partners during the mating season erally do not exhibit this behaviour, probably because they (Vogt et al. 2014). have no need to establish territories nor advertise availability The territorial behaviour of animals is generally consid- to mate (Allen et al. 2021). ered to be determined mainly by food availability and dis- Small cats (Felinae) exhibit strong uniformity in both tribution, which has a major impact on animal fitness, espe- scent marking and social behaviours (Mellen 1993). Repro- cially in females (Carpenter and MacMillen 1976; Maher ductively active animals mark more frequently than others, and Lott 2000; McLoughlin et al. 2000; Adams 2001; Borger especially during oestrus (Mellen 1993). In a female black- et  al. 2008). In Eurasian lynx, prey availability strongly footed cat, the marking frequency peaked before the mat- affects territory size (Mengüllüoğlu et al. 2021; Aronsson ing season and was absent during the period of rearing of et al. 2016) and reproductive success in females (Yom-Tov young, thus advertising female reproductive status (Molteno et al. 2010; Nilsen et al. 2010). Lynx territoriality is also et al. 1998). In the bobcat, marking is primarily exhibited strongly dependent on the animal's social status, with age, by adults (Bailey 1974), although juveniles were found to sex, reproductive status, population density, and spatial distri- mark from 5 months of age (Wassmer et al. 1988). In both bution affecting the size and even the very existence of lynx bobcat and Iberian lynx, scent marking seems to mainly play territories (Mengüllüoğlu et al. 2021; Aronsson et al. 2016). a role in territory defence, while its connection to mating In the Bohemian-Bavarian-Austrian lynx population, behaviour is not yet clear (Wassmer et al. 1988; Bailey 1974; the mean annual home range sizes are 438 km for adult Beltran et al. 1987; Barja et al. 2005). males and 278 km for adult females (own unpublished data, The Eurasian lynx (hereafter: lynx) is known to scent Belotti et al. 2012; Magg et al. 2016; Wölfl et al. 2001). mark with urine at visually conspicuous sites such as rocks, Generally, the home range of one male overlaps with those fallen tree trunks, roots, wood piles, wooden cabins, and of one to three females (Mináriková et al. 2020; Wölfl et al. along frequented lynx paths, where it is probable that con- 2020; Engleder et al. 2019, 2021). Home ranges of adult specifics will find the scent mark (Allen et al. 2017; Sæbø lynx of the same sex may partially, but never entirely, over- 2007; Vogt et al. 2014; Stromková 2020; Krofel et al. 2017). lap (Mináriková et al. 2020; Wölfl et al. 2020; Engleder et al. Faeces are also frequently found exposed at marking sites, 2019, 2021). although at resting sites or nearby kill lynx usually hide Here we present a unique, well-documented case of an them (Sæbø 2007; Stromková 2020, own unpublished data). actively marking juvenile lynx. Marking behaviour is not equally distributed in time but is It was recorded in the so-called Bohemian-Bavarian- most frequent during the mating season (January-April) and Austrian (BBA) lynx population, whose distribution area least frequent during the period when females give birth stretches across the borders between the Czech Republic, and lactate (May–July) (Vogt et  al. 2014). Males mark Bavaria, and Austria (Fig. 1). more frequently than females (Vogt et al. 2014; Sæbø 2007; Camera trapping is used as a major lynx monitoring Stromková 2020). Juveniles visit marking sites with their method in all three countries, complemented by snow track- mothers, but according to the three large-scale scent mark- ing and genetic analyses. To space the camera traps evenly ing studies of Alpine, Scandinavian, and Carpathian lynx across the region, we used a 10 km × 10 km grid (ETRS populations (Vogt et al. 2014; Sæbø 2007; Stromková 2020), LAEA 5210 10 km grid) and installed two to eight camera 1 3 acta ethologica Fig.1 Map of Bohemian- Bavarian-Austrian lynx popu- lation. The wider region of Vlachovo Březí forests, where the yearling female Koka was documented, is circled in red trapping sites per grid cell, depending on the area and habi- – “Independent lynx”—lynx older than 1 year, i.e., sub- tat type. Altogether, 131 grid cells were covered by camera adult or adult. trapping in the years 2017–2021 as part of the 3Lynx project – “Resident lynx” lynx staying for minimally 12 months in and following national activities (Mináriková et al. 2020; the same area after becoming independent. Wölfl et al. 2020; Engleder et al. 2021). The time unit for data analysis was lynx year, that is Lynx females B748 Koka and B749 Baronka were born st th the period starting on the 1 May and ending on the 30 in spring 2019 to B718 Nela, a resident female in the Vla- April of the following year, based on the lynx life cycle chovo Březí area (Czech Republic, South Bohemian region, (Mináriková et al. 2020). In the data analysis, the follow- Fig.  1). They were born in the territory of resident male ing categories were used for lynx according to their status: B716 Karlos. Karlos and Nela both disappeared during the lynx year – “Juvenile lynx”—lynx in the first year of life (also 2019 and have not been recorded since then, thus we assume th called “kitten”), recorded from birth till the 30 April they are dead. Karlos’s territory was taken over by the sub- of the following calendar year (0–1 year of age). adult young male B740 Piškot in November 2019. Nela was th – “Subadult lynx”—lynx in the second year of life, last recorded with both her kittens on the 13 December st recorded from the 1 May of the year following the birth 2019, but the exact date of her death remains unknown as th till the 30 April of the next year (1–2 years of age). she could have been missed by camera traps for some time. – “Adult lynx”—lynx older than 2 years. Her territory was not taken over by any subadult or adult 1 3 acta ethologica female. Therefore, both her juvenile daughters were left with the possibility of settling into the vacant territory they already inhabited as kittens. th On the 30 March 2020, during the mating season, Koka was recorded on a video actively marking with urine, rub- bing her face, and again marking with urine on a marking site (Fig. 2). On the following day, she again marked with th urine on the same spot (Fig. 3). On the 29 March this mark- nd ing site was visited by Piškot and on the 2 April it was visited by Baronka, but they were not recorded marking, they both merely inspected it. It was a rock, previously used for marking by resident female Nela, previous resident male Karlos, new resident male Piškot, and later Baronka. rd th Between the 3 and 26 of April 2020, after the end of the mating season, Koka left the area and dispersed 40 km far to the Boletice military training area and then to the Protected Landscape Area Blanský les, where she has lived since July 2020. The former territory of Nela was taken over by Bar- onka, who is till now (December 2022) the resident repro- ducing female lynx here, sharing the area with male Piškot. Lynx females are known to successfully reproduce from the age of 2 years (Breitenmoser-Würsten et al. 2007; Nilsson 2013; Nilsen et al. 2010, 2012; Kvam 1991). Juvenile lynx ovulation was recorded in Norway, but no juvenile pregnancy nor parturition was found (Kvam 1991). The BBA lynx popu- lation is, to the best of our knowledge, the only European lynx population where juvenile lynx females were documented to successfully give birth and raise kittens to independence (Engleder et al. 2019; Palmero et al. 2021, own unpublished data). This poses the question of whether the adult-like mark- ing behaviour of a juvenile female lynx should be explained in the context of early reproduction or if it has other reasons. While the marking behaviour in lynx generally serves both to attract conspecifics of the opposite sex and to mark the territory (Vogt et al. 2014), we cannot assess what was the main purpose of Koka’s behaviour. Neither Koka nor Baronka successfully reproduced with the resident male Piškot during their first year of life but that does not mean they did not try to attract him (or any other male) by mark- ing. Also, juveniles’ competition over the territory left vacant by their mother is a possible explanation for the marking behaviour, with Baronka winning over and displac- ing Koka, who then dispersed to a new area. The BBA lynx population has a high turnover of adult Fig. 2 Juvenile female Koka actively marking with urine, rubbing her th and subadult lynx, similarly to its neighbours – the West- face and again marking with urine on a marking site (the 30 March 2020) ern Carpathian and the Alpine lynx populations (Wölfl et  al. 2020; Duľa et  al. 2021; Breitenmoser-Würsten et al. 2007). Specifically, 23% of adults and 44% of sub- adults vanish every year, especially in the outskirt part lynx presence (Mináriková et al. 2020; Wölfl et al. 2020; Engleder et  al. 2021). This instability may result in of the BBA population’s distribution (Wölfl et al. 2020). This inevitably causes social instability, with territories unusual social situations, like the one presented in this paper, with juveniles consequently performing unusual changing their owners frequently and some areas being temporary without lynx reproduction or entirely without behaviours. 1 3 acta ethologica Funding Open access publishing supported by the National Technical Library in Prague. This study was supported by the CE1001 3Lynx Project, funded by Interreg Central Europe. Data availability The datasets analysed during the current study are available from the corresponding author on reasonable request. Declarations Conflict of interest The authors declare that they have no conflict of interest. Open Access This article is licensed under a Creative Commons Attri- bution 4.0 International License, which permits use, sharing, adapta- tion, 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 Fig. 3 Koka again marking with urine on the same marking site (the otherwise in a credit line to the material. If material is not included in st 31 March 2020) 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://cr eativ ecommons. or g/licen ses/ b y/4.0/ . 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Acta Theriol 46(2):181–194. https:// doi. org/ 10. 4098/ at. arch. 01- 20 Publisher's Note Springer Nature remains neutral with regard to Wyatt TD (2003) Pheromones and animal behaviour (Vol. 626). Cam- jurisdictional claims in published maps and institutional affiliations. bridge University Press, Cambridge. ht tps:/ / doi . or g/ 10. 10 17/ cbo97 80511 615061 1 3

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acta ethologicaSpringer Journals

Published: Feb 1, 2023

Keywords: Eurasian lynx; Lynx lynx; Scent marking; Behaviour; Juvenile; Orphan; Bohemian-Bavarian-Austrian lynx population; BBA

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