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Use the following of citation: North-western Journal of Zoology 2021: e212701

Paper Submitted to The North-Western Journal of Zoology

1 *Handling editor:

2 *Manuscript Domain:

3 *Manuscript code: nwjz_21_TH_04

4 *Submission date: 08_02_2021

5 *Revised: 22_05_2021

6 *Accepted / Rejected: 25_05_2021

7 *No. of words 1558 8

9

10 Title of the paper:

11 First records of golden and European non-antagonisticZoology interaction at the of 12 Lower Danube (Bulgaria) 13 Running head: proofing 14 First records of jackal-badger interaction Journaluntil 15 Authors (First LAST - without institution name!):

16 Yulian KONSTANTINOV, Nikolai SPASSOVpaper and Ilya ACOSTA-PANKOV 17

18 Key Words (at least five keywords):

19 jackal-badger interaction,North-western commensalism, non-antagonistic, Lower Danube, NW Bulgaria Accepted 20

21 No. of Tables: 1

22 No. of Figures: 2

23 No. of Files: 4

24 25 Use the following type of citation: North-western Journal of Zoology 2021: e212701

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26 {First records of and non-antagonistic interaction at the

27 Lower Danube (Bulgaria)}

28 {Yulian, KONSTANTINOV}1, {Nikolai, SPASSOV}2 and {Ilya, ACOSTA-PANKOV }2

29 1. Affiliation

30 Bulgarian Society for Regional Cultural Studies (BSRCS) [email protected]

31 2. Affiliation

32 National Museum of Natural History, Bulgarian Academy of Scienses, 1 Tsar Osvoboditel

33 Blvd, 1000 Sofia, Bulgaria, [email protected]; [email protected]

34 * Corresponding authors name and email address: Ilya Acosta-Pankov, Vertebrates

35 department, National Museum of Natural History, Bulgarian Academy of Scienses, 1 Tsar Zoology 36 Osvoboditel Blvd, 1000 Sofia, Bulgaria (E-mail: [email protected]). of 37 proofing 38 Abstract Journal 39 We present a case of non-antagonistic interaction ofuntil golden jackal (Canis aureus) with 40 European badger ( meles), recorded at the Bulgarian bank of the Lower Danube in 41 the summers of 2019 and 2020. Such interactionspaper among the mentioned species have not 42 been reported so far, however are reminiscent of similar mesopredators’ relationships 43 known among other canids and large mustelids. The relationship occurred in a jackal 44 family group researchNorth-western context, which has been studied during consecutive summers since 45 2017. The research was Acceptedbased on establishing a bait-station monitored by a camera trap 46 since 2018. In the field season of May-September 2019, and again in April–September 47 2020, as well as in April to May 6, 2021, the camera trap registered numerous cases of a 48 badger being present simultaneously with a member or members of the jackal group at the 49 bait-site. These findings made us wonder to what extent commensalism or mutualism 50 could exist in this intraspecific behaviour. This main question will have to be answered by 51 future research. The observed interaction revealed intraspecific tolerance and can be 52 approached from the perspective of the modern concept of the influence of facilitation on 53 the organization of natural communities. 54 Use the following type of citation: North-western Journal of Zoology 2021: e212701

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55 Key Words: jackal-badger interaction, commensalism, non-antagonistic, Lower Danube, NW

56 Bulgaria

57 Running title:

58 First records of jackal-badger interaction

59 Introduction

60 It is assumed that among ’ overt aggression between species occupying similar 61 ecological niches is a major factor influencing their ecology and evolution (Macdonald et al. 62 2004 and references herein). The coexistence of mesopredators competing for the same prey 63 is related to the complex ecosystem relationships. They have been evolutionary created for 64 spatial and temporal distribution of ecological niches, and mutual avoidance. These 65 relationships are dynamic and dependent on population density, prey selection, catch, and 66 seasonal variation (Barrull et al. 2014). In these complex interactions,Zoology not only aggressiveness 67 but also interspecific reciprocity is also often built up, rangingof from tolerance to various 68 degrees of symbiotic relationships. In central Bulgaria, (Tsunoda et al. 2018) have reported 69 non-simultaneous co-occurrence of golden (Canis proofingaureus) and European 70 (Meles meles) at seven camera trap stations in a field study with 17 stations. Journaluntil 71 Interspecific tolerance bordering on commensalism, or even mutualism in hunting and feeding 72 behavior, has been noted in rare cases between sympatric mesopredators as some canids and 73 large mustelids in North America, Africapaper ancd Europe. Such cases of nonaggressive 74 interactions or tolerance have been observed in foraging behavior between (Canis 75 latrans) and (Taxidea taxus) (Aughey 1884, Hawkins 1907, Cahalane 1950, 76 Lehner 1981, MintaNorth-western et al. 1992, Cook 2000) , as well as in some non-antagonistic forms of 77 -American badger relationsAccepted (Clark et al. 2015). Interactions between black-backed jackal 78 (Canis mesomelas) and the (Mellivora capensis) in relation to prey have been 79 described in Namibia (Gorta 2020) as either commensal interaction, or a kind of ‘producer- 80 scrounger’ kleptoparasitism. Relationships of marked tolerance between and European 81 badgers (Meles meles) have also been observed in artificial feeding sites in England, where 82 despite the dominance of the badger, no aggression was observed, and it was even assumed 83 that foxes sometimes seek the company of badgers (Macdonald et al. 2004). 84 In this short note we are reporting non-antagonistic interactions between the golden jackal 85 (Canis aureus) and the European badger (Meles meles), like the cases mentioned above, 86 especially as regards – golden jackal interactions. Golden jackal – badger interactions Use the following type of citation: North-western Journal of Zoology 2021: e212701

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87 have not been previously recorded in Bulgaria, nor in the existing golden jackal-related 88 literature as regards jackal presence elsewhere. The lack of such observations is probably due 89 not only to the poor study of such interactions, but also to the relatively recent active dispersal 90 and long-term entry of jackals into European ecosystems (Arnold et al. 2011, Spassov & 91 Acosta-Pankov 2019). 92 93 Material and Methods

94 Between 2017 and May 2021, a field study of jackal family group behaviour was conducted on 95 the Bulgarian bank of the Lower Danube, in North-West Bulgaria, near the village of Stanevo 96 (Lom Municipality). The habitat of this area is suitable for jackals, whose expansion has been 97 well noted during the last 20-30 , while previous to this period they were unknown in this 98 part of Bulgaria (Spassov 1989, Stoyanov 2013, Spassov & Acosta-Pankov 2019). The 99 observations were made with camera-traps since 2018, using bait as attractant. Food was placed Zoology 100 regularly at the bait-site close to the field-research tent camp. The distance between bait-site of 101 and camp depended on changing river water levels. It was 22 m in 2018, 6 m at very high-water 102 level in May–June 2019, 22 m in July–September the sameproofing , and 20 m in April–September 103 2020, as well in April 2021. This relatively short distance allowed a resident jackal family Journal 104 group of 5–7 members to get used to all visual, auditory,until and olfactory signals of anthropogenic 105 origin coming from the camp. Continuous observations made it possible to establish the 106 approximate age and gender of family members,paper as well as their hierarchical position. All 107 details were entered into field-diaries and visual recording protocols. 108 Results

109 The camera trap coveredNorth-western nights recording jackal group visits added up to 257 nights for 2018– 110 2020, within a total of 307Accepted observed nights since 2017. In the currently ongoing field-study of 111 2021, 14 nights have been monitored so far. The family group ‘appropriated’ the bait-site in the 112 course of time, almost completely barring access to other competing scavenging species (foxes, 113 feral and ). This was done by regular nightly presence, bait-site use, and olfactory 114 marking. 115 The visits of other species, who happened to be registered at the site, were very rare. There 116 were only six such visits: two with foxes, three with stray cats, and one with a stray , none 117 of them was interactive. In the current 2021 field study, a feral appeared briefly in the first 118 two nights and then disappeared, again without interacting in any way. Use the following type of citation: North-western Journal of Zoology 2021: e212701

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119 In this context, the appearance of a badger and its subsequent interaction with jackals stood 120 out. Some of the camera trapped jackal – badger interactions during these periods are 121 described below. 122 The first appearance of a badger was recorded on the night of 24–25 May (2019), and the first 123 record of a jackal and a badger together was on the night of 28–29 May (2019). During the 124 last two observation periods (2019–2020, and April to May 6, 2021) a total of 93 jackal- 125 badger interactions had been recorded, including some in which a jackal and a badger were 126 eating side by side or nose-to-nose (Fig. 1a). In the specific registered cases, badger visits 127 occurred in ‘mixed’ nights, i.e., nights in which badger and jackals were present together at 128 the bait-site (‘interactive visits’), or the visits of both species succeeded each other at intervals 129 of varying length (Table 1). 130 The behaviour during the registered total of 88 interactive visits can be categorized as non- 131 antagonistic behaviour of interspecific tolerance as no case of a scuffle or fighting between 132 the two species had been noted. Only some behavioural responsesZoology of subadult jackals were 133 registered showing some tension in the presence of the badger,of while, in general, the badgers 134 appeared to be tolerant to the presence of jackals (Fig. 2). proofingAnother pattern of behaviour that 135 was observed during this period (2019–2020) we called ‘jackal sentry response’. Behaviour of 136 a jackal or jackals standing sideways in a ‘sentry’Journal poseuntil and looking in all directions were 137 recorded, while the badger was eating. Several similar interactions were registered (Fig. 1). 138 There thus appears to be a similarity between this behaviour assumed by jackals, and what paper 139 Minta et al. (1992) call ‘coyote sentinel’ in the context of coyote – American badger 140 association for hunting ground-squirrels. 141 Aside from occasionalNorth-western scenes in which lower mutual tolerance was shown, the predominant 142 case has been of jackal(s)Accepted and badger feeding together (Fig. 1a, 2), after an initial period of 143 tension had passed. 144 Discussion

145 As shown above, a case of interaction is at hand, which can be generally considered as non- 146 antagonistic. If any antagonism is manifested, then the aggression is weak and does not 147 exceed that which is sometimes manifested between the members of the jackal family group. 148 The two species use the resource of the bait-site in a spirit of mutual tolerance with occasional 149 tense behavior exhibited by the badger. Such tension is only very rarely exhibited by jackals, 150 except for the alpha male. Overwhelmingly, the jackals show a submissive role in respect of 151 the badger, barring an occasional evasive movement on the badger’s part in the presence of Use the following type of citation: North-western Journal of Zoology 2021: e212701

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152 the alpha male. This type of behaviour suggests a badger role consistent with the hierarchy of 153 the jackal group. In this sense, the badger behaves as a meta-beta member, subservient only to 154 the alpha-jackal. Another feature of the interaction is its seasonal dynamic. In late spring/ 155 early summer, when the resource base is still scanty, cases of higher tension tend to be more 156 numerous than later in the summer when food sources are more abundant. As said at the 157 beginning, given the recent appearance of jackals at the study area (in the late 1990s) the 158 interactions registered are of very recent origin. These newly developed behaviour of 159 tolerance support the claim that interacting species should evolve an adaptive response to 160 conflict, which minimizes costs to their own fitness (Huntingford & Turner 1987, Macdonald 161 et al. 2004). 162 Observed interspecific relationships exhibiting interspecific tolerance can be interpreted from 163 the perspective of the modern concept of the influence of facilitation (i.e. positive species 164 interaction) on the organization of natural communities and the importance of a wide variety 165 of facilitative interactions in these communities (Bruno et al.Zoology 2003). Such relationships of 166 mutual facilitation between different species have been describedof for some 167 communities in Białowieża primeval forest, which includeproofing invasive newcomers (Kowalczyk 168 et al. 2008). The case we have described also partially resembles the one in Clark et al. (2015) 169 related to the authors’ description and analysisJournal of Sanuntil Joaquin ( macrotis 170 mutica) interacting with American badger (Taxidea taxus). In our case, there has been 171 evidence of non-antagonistic jackal-badger interaction in a context of human-created paper 172 additional conditions. What each one of the species ‘gets’ from the relationship, remains 173 another open question. At the moment, a plausible explanation seems to be what has been 174 stated by Clark et al.North-western (2015), who suggests that this kind of tolerance may be preferred as a 175 least-cost pathway. The Acceptedjackals’ behavior pattern called by us ‘sentry response’ may suggest, 176 however, an interpretation extending these limits. Is it possible that the badger benefits from 177 eager jackals ‘sentries’ around him/her, particularly at hibernating periods? Such possible 178 interpretations, connected with give-and-take mutualism, based on denning, remain, however, 179 questions to be answered by future research. 180 181 Acknowledgement

182 We are grateful to Familia Nikola Ltd. for supplying us free of charge with oat-flake products 183 for bait since 2017. 184 Use the following type of citation: North-western Journal of Zoology 2021: e212701

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185 References

186 Arnold, J., Humer, A., Heltai, M., Murariu, D., Spassov, N., Hackländer, K. (2011): Current

187 status and distribution of golden jackals Canis aureus in Europe. Review 42 (1):

188 1-11. https://doi.org/10.1111/j.1365-2907.2011.00185.x.

189 Aughey, S. (1884): Curious companionship of the coyote and the badger. American Naturalist

190 18: 644–645.

191 Barrull, J., Mate, I., Ruiz-Olmo, J., Casanovas, J.G., Gosàlbez, J., Salicrú, M. (2014): Factors

192 and mechanisms that explain coexistence in a Mediterranean carnivore assemblage: an

193 integrated study based on camera trapping and diet. Mammalian Biology 79 (2): 123-131.

194 Bruno, J.F., Stachowicz, J.J., Bertness, M.D. (2003): Inclusion of facilitation into ecological Zoology 195 theory. Trends in ecology & evolution 18 (3): 119-125. of 196 Cahalane, V.H. (1950): Badger-Coyote Partnerships. Journal of Mammalogy 31 (3): 354-355. proofing 197 Clark, H.O., Powers, R.M., Uschyk, K.L., Burton, R.K. (2015): Observations of antagonistic Journal 198 and nonantagonistic interactions between the Sanuntil Joaquin kit fox (Vulpes macrotis

199 mutica) and the American badger (Taxidea taxus). The Southwestern Naturalist 60 (1): paper 200 106-110.

201 Cook, L. (2000): The coyote and the badger. Blue Jay 58 (4): 185-186.

202 Gorta, S.B.Z. (2020).North-western What goes around comes around: complex competitive interactions Accepted 203 between two widespread southern African mesopredators. Canid Biology & Conservation

204 22(2): 8-10.

205 Hawkins, A. (1907): Coyote and badger. Ottawa Naturalist 21: 37.

206 Huntingford, F.A., Turner, A.K. (1987). conflict. London. Chapman & Hall.

207 Kowalczyk, R., Jędrzejewska, B., Zalewski, A., Jędrzejewski, W. (2008): Facilitative

208 interactions between the Eurasian badger (Meles meles), the red fox (Vulpes vulpes), and Use the following type of citation: North-western Journal of Zoology 2021: e212701

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209 the invasive dog ( procyonoides) in Białowieża Primeval Forest,

210 Poland. Canadian Journal of Zoology 86 (12): 1389-1396.

211 Lehner, P. (1981): Coyote-badger associations. Great Basin Naturalist 41 (3): 347-48.

212 Macdonald, D.W., Buesching, C.D., Stopka, P., Henderson, J., Ellwood, S.A., Baker, S.E.

213 (2004): Encounters between two sympatric carnivores: red foxes (Vulpes vulpes) and

214 European badgers (Meles meles). J. of Zoology, Lond. 263: 385–392

215 Minta, S.C., Minta, K.A., Lott, D.F. (1992): Hunting associations between badgers (Taxidea

216 taxus) and (Canis latrans). Journal of Mammalogy 73 (4): 814-820.

217 Spassov, N. (1989): The position of jackals in the Canis and life history of the golden

218 jackal (Canis aureus L.) in Bulgaria and on the Balkans. Historia Naturalis Bulgarica 1: Zoology 219 44-56. of 220 Spassov, N., Acosta-Pankov, I. (2019): Dispersal history of the golden jackal (Canis aureus proofing 221 moreoticus Geoffroy, 1835) in Europe and possible causes of its recent population Journal 222 explosion. Biodiversity Data Journal, 7: e34825;until doi: 10.3897/BDJ.7.e34825.

223 Stoyanov, S. (2013): Population ecology studies on the jackal (Canis aureus Linnaeus, 1758) paper 224 in Bulgaria. PhD Thesis. University of forestry, Faculty of Forestry, Sofia. 148 pp. [In

225 Bulgarian].

226 Tsunoda, H., Ito, K.,North-western Peeva, S., Raichev, E., Kaneko, Y. (2018): Spatial and temporal Accepted 227 separation between the golden jackal and three sympatric carnivores in a human-modified

228 landscape in central Bulgaria. Zoology and Ecology 28 (3): 172-179.

229 https://doi.org/10.1080/21658005.2018.1504406.

230 Table captions (if applicable)

231 Table 1. Number of interactive visits during mixed nights in relation to all monitored nights in 232 the research periods of 2019 and 2020. 233 Monitored season 2019 Monitored season 2020 Total Use the following type of citation: North-western Journal of Zoology 2021: e212701

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Monitored nights 108 98 206 ‘Mixed’ nights 56 65 121 ‘Interactive visits’ 48 40 88 234 235 Figure captions (if applicable)

Zoology of proofing 236 237 Figure 1. An adult male jackal eating nose-toJournal-noseuntil with badger at the eating site, two sub- 238 adults in ‘sentry’ pose behind (a). Record of ‘sentry response’ in May 2021 (b). Record of 239 ‘sentry response’ in September 2019 (c). Badger eating, jackals at ‘sentry’ position on the paper 240 side, waiting for their turn at the eating site (d). (Photos: Y. Konstantinov). 241 North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e212701

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242 243 Figure 2. Recording of an adult male jackal turning back to lookZoology at an incoming badger (a). 244 Recording of a sub-adult jackal and badger eating side byof side. The sub-adult jackal was tense 245 while the badger was indifferent (b). Adult jackal eating nose-to-nose with a badger, while a proofing 246 sub-adult is waiting behind (c). Nose-to-nose eating with badger (d). (Photos: Y. 247 Konstantinov). Journaluntil 248

249 paper

North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e212701

Table 1 - Number of interactive visits during mixed nights in relation to all monitored nights in the research periods of 2019 and 2020.

Monitored season 2019 Monitored season 2020 Total

Monitored nights 108 98 206

‘Mixed’ nights 56 65 121

‘Interactive visits’ 48 40 88

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North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e212701

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North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e212701

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North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e212701

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North-western Accepted Use the following type of citation: North-western Journal of Zoology 2021: e212701

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