Coauthor Irina Rotenko
Concerning the wolf Canis lupus breeding in Europe and wider in Eurasia there is a widely spread belief that the species is strictly monogamous with a certain way of breeding and pack formation in family pattern (e.g. Bibikov D.I., 1985 and references therein; Jędrzejewska & Jędrzejewski, 1998).
Those beliefs suggested the three following theses: first, during breeding season, a wolf pack has merely one litter or there are no pups; second, usually a wolf pack consists of parent wolves and their pups of the current and previous biological years as a normal maximum; additionally, such a pack may subordinate some non-relative wolves; and third, usually offspring disperse from their maternal pack around mating season, in the second year of their life, when they are 20-22 months old.
Actually, in our study in Belarus since the late 1990s we have found so many abnormalities, that we start thinking about what actually prevails in the wolf reproduction: the “rules” or “exceptions”. Such “abnormalities” occurring often is a very interesting phenomenon.
This post is entirely addressed this batch of questions on the wolf reproduction by basing on the example of one model wolf pack in Naliboki Forest, the central-western Belarus. This pack was leaded (still leading) by female, which is called by us as Torn Ear. This wolf female has been easily recognized by the tear in her right ear since February 2017, when we began to call her Torn Ear. Before that we recognized her by whitish stripes in the lower part of her head and black marks at the eyes.


In Naliboki Forest since February 2013 Torn Ear has been registered living and reproducing in the wolf group of three breeding females and one or two adult males altogether. The whole this quite long period of 8 years just Torn Ear leaded and still leading her pack. Her leading of the wolf pack was evidenced by numerous photos taken by camera-traps.
On 11th May 2013 we discovered three wolf litters in the dens within the small spot of spruce-black alder forest somewhere between the localities of Drazdy and Rabachova. The wolf dens that belonged to the three breeding female wolves were situated at the distance of about 430 meters, 220 meters and 380 meters between each other. The pups in one litter were about 40 days old, the second litter was about 15-18 days old, and the third one consisted of small blind pups. The first and second litters were mixed as: 2 pups of 40 days old and 2 pups of 15-18 days old; and as 2 and 3 such different age pups, respectively. It happened like that because the mothers stole pups from the dens of each other. None of the pups including the litter of Torn Ear survived by the winter 2013-2014.


During the breeding seasons of 2014 and 2015 we did not collect enough precise information about the reproduction in this wolf group. At least, we knew that they bred and had pups till the late summer or autumn, but during the coming winter there were again five adults only without any pups i.e. three adult females and two adult quite big males. Sometimes, one of the males walked separately, but within the same territory of the other wolves of the pack. Again, sometimes we registered that Torn Ear walked separately with one of the males, while other two females stayed with another male within entirely the same territory.
In April-May 2016 one or more females from the wolf breeding group that included Torn Ear gave birth in the Rabachova locality; there was registered minimum one litter. But all the pups disappeared by the end of May.
In February 2017 Torn Ear (when she was with a normal right ear) and other four members of the wolf pack were taken in a fladry enclosure in the localities of Kurhany and Symonava by the local hunters. That day, by driving there, we saw a number of vehicles and more than thirty hunters, who did the hunt-drive to kill the wolf pack. The hunter tried to do their bloody work during the whole day. Nevertheless, Torn Ear and other wolves escaped more or less safely from the fladry enclosure. One shot of hunter wounded the right ear of the female (since that time the female have got the torn ear). Also one male wolf from the pack was wounded to the right hind leg, but he survived.
In the late April 2017 all three females of the wolf pack gave birth definitely. The distances between their initial dens were 0.92, 0.72 and 0.95 km from each other. One male mostly looked after two breeding females including Torn Ear, whereas the other male took care about the third breeding female. Torn Ear was registered visiting not only own den, but also the den of another breeding female, to which the same male served. Interestingly that at one of the dens all these five wolves were registered altogether on 1st May. We documented that two litters were killed by adult male lynx that we called Jury. It happened in the end of April and the first days of May, whereas the third litter disappeared in the mid-June. In 10-15 days afterwards all the five wolves started walking widely within the previous winter home range and marking a lot.
After the clear documentation that the two wolf dens were extirpated by the adult lynx male we started thinking that all previous breeding failures of the wolf pack happened due to the same lynx male aggressivity to breeding in the wolves.





In the late April 2018 the wolf breeding group consisting of Torn Ear, two more breeding females and still two adult males began preparing denning in the same Rabachova locality again. They prepared minimum 5 dens before any parturition. Everywhere in the places they stayed there were registered lynxes in the close proximity. Then suddenly before any giving birth the wolf breeding group disappeared and they were not found in the whole Naliboki Forest during the summer. We hypothesized that they emigrated from Naliboki Forest to breed in the surrounding more rural area where there were not so many lynxes around because of the dramatic experience of the previous May, when Jury the lynx had killed all their pups.

After a long absence, Torn Ear with one of the males and one of the females was found to come back to their former territory in early March 2019. They were with another unknown third female, which could be a young adult (i.e. the one survived pup of the year). The area was already occupied by other three wolves. These wolves were gradually sent away by the former owners including Torn Ear.
In late April 2019 Torn Ear and at least one more female of this wolf group gave birth. Torn Ear’s den was placed in the Asovyja locality, while another female denned in the Rabachova locality. Their single male served for both mothers, which together with their pups were situated at the distance about 3 km apart. Then since the mid May both mothers with their litters were registered on stay in the Asovyja locality. Another female’s litter was situated in a badger outlier, where the pups were killed by a male lynx. We registered the tracks of the lynx entering the burrow-den and two pup remains about 40 and 120 meters away of the den. The litter of Torn Ear was saved and in July it consisted of 6 pups you may see in the photos below. Both females and their male took care about the pups.





Till December of 2019 only Torn Ear, another adult female, their adult male and 6 pups composed the pack. In the mid of December another bigger adult male joined the pack. Afterwards Torn Ear became more related to the first male, while another breeding female seemingly paired with the arrived adult male. Anyway these four adults (two breeding females and two males) walked mainly altogether till the mid-April 2020. Moreover, in the mid-January this pack subordinated three more wolves. In the late February the whole large pack already of 13 wolves were taken in fladry by local hunters. In the effect of that hunting 6 wolves were killed, while 7 of them including the mentioned two pairs of adults succeeded to escape. In the mid-April in the core area of the pack home range we found the two couples including Torn Ear with her male. Till the late May both couples stayed on the area about 30 square km maximally in the localities of Rabachova, Drazdy, Kazialiets and Patashnja.



The second breeding female, that walked together with Torn Ear, gave birth around 3rd May (on 13th May we found her den with 9 pups about 10 days old), and the new adult male served for her (see the photos below). This subfamily of the Torn Ear breeding group (hereafter the first subfamily) moved for about 1.5 km after we discovered their active den.




Moreover, on 18th of May we photographed one more heavily pregnant female (not Torn Ear) at the distance of about 800 meters from the discovered den (see the pregnant female in the photo below). That still pregnant female was plausibly one of the three wolves, which seemingly disappeared in the early April. Maybe they still stayed not faraway, but we did not know. This pregnant female plausibly gave birth on 19th or 20th of May, because on 21st May we found all the features of presence of one more den with pups there on the distance about 1-2 km from the first discovered active den. Another male served for this mother. That was the third founding male in the Torn Ear breeding group in May 2020.

Then in the last days of May this subfamily moved for distance of 2-2,5 km from the initial denning area and appeared at the distance about 1.8 km from the first mentioned subfamily actual denning area. In the photo below see one of the new dens of the second subfamily.

The breeding story of the old female Torn Ear in May 2020 was unclear, at first. Nowadays we got more or less sufficient information to reconstruct precisely enough her breeding story in 2020. Torn Ear is an old female wolf and in this year she got estrus quite late in the mid-March. Respectively, Torn Ear gave birth approximately on 20th of May at the distance about 0.7 km from the initial den of the first mentioned subfamily. However, that place was situated on the opposite site of the Biarezina medium-sized river.


In the conclusion, we would like to say that the presented multiannual example of out-rule packing and breeding in wolves suggests that the given story is not something abnormal, exceptional or extraordinary. The phenomenon of pack multi-breeding and presence of several (2 or even 3) big males in one breeding group of wolves, which are playing a founding role in the pack breeding, was repeated across this story from year to year with changing composition of the pack. Therefore it is quite opposite, the revealed phenomenon looks like something usual, common and routine.
Also, here it is worthwhile mentioning one our hypothesis on multi-breeding in a wolf pack relating presence of either merely one or several founding males in the pack. Our (still insufficient) data suggest that in case a leading individual in a wolf pack is a female, there can be not only one but two or even three founding males if double or triple breeding happens in the pack. Two founding males look like a norm in such a case, and there can be even three such males. When a male leads a pack, a founding male is mainly one even in a case of multi-breeding.

I would like to know more about what is meant by “founding males”。Are they made up of wolves of different bloodlines? Did they have different territories before forming large packs? Also, what do you think about the so-called restriction of mating activities of wolves of the same sex during the breeding season to form what is often called “wolf monogamy”? Tank you.
A founding male means father of pups. It looks like in the Torn Ear pack there were several founding males. It is very plausible. Anyway, in the denning period of 2020 in a small locality there were 3 wolf couples with pups each; their dens were situated in a proximity, and they met each other every day peacefully. The three adult males (everyone was big) hunted together in the early denning period. Then in the mid-summer they stayed altogether with the mothers and their pups. In May 2021 in the Torn Ear breeding group there were again three breeding females and two big males; one male served for two females with pups. We do not know for sure what is a mating system in such a breeding group of wolves, however, it looks like it may be that a given female in heat could mate with two or even three males. These breeders (6 in 2020 and 5 in 2021) stayed together on the same territory during the winters before denning in May. I assume that a ‘wolf monogamy’ happens less often than the species polygamy, at least, in Belarus. In Belarus, a founding male wolf mates with a founding female and his daughters on the second year of their life not infrequently.
Thank you Mr. Sidorovich for your answer, there are some more things I would like to know. Regarding the naliboki region of Belarus, is wolf mortality due to conflicts between different wolf packs common? What is the proportion of deaths caused by intra-species struggle in natural mortality? What is the attitude shown by local wolves towards non-packs members? I once read an article about wolves in the Yellowstone National Park area and it mentioned that wolves there have a 67% mortality rate due to intraspecific factors. I know that the wolf population density there is high, so I would like to know the following mortality events and mortality rates caused by intra-species struggle of wolves in places like the naliboki area where wolves do not have a great advantage over other carnivores.
In Naliboki Forest as well as in the whole Belarus killing of a wolf by another one (or by a pack) is rare. Conflicts between wolves are common, but they do not kill each other. I know this question good enough. For my part, American wolves are like a different species taking into account all the features (Sidorovich & Rotenko, 2019). Therefore, the pattern from Yellowstone National Park is not a good argument.
Yeah, we really can’t generalize about wolves around the world. Yellowstone wolves do have a very high population density, even much higher than in many parts of North America, and I feel that the intra-species struggle of wolves in that part of the world has more to do with that. The composition of wolf populations, inter-population relationships, ecological niches, etc. may also be very different in different parts of the world and cannot be generalized. For example, wolves can exclude pumas in some parts of North America, but are bullied by Eurasian lynx in naliboki. I’ve been lucky enough to see animal lovers on some forums sharing your and other scholars’ research on wolves, and I think it’s amazing how diverse those wolves are in all aspects of their behaviour in different habitats.
Thanks, I agree with that.