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European Bison (Bison bonasus) or ‘Wisent’ bounced back from complete extinction in the wild. Every animal is descended from just 12 individuals – part of a successful captive breeding programme, which started in the mid 20th century.

These huge herbivores were wiped out in their original open habitat, and for many years, we wrongly believed that they were an animal of dense forest. However, they do need woodland, for it provides a shady refuge from summer heat and a source of leaves, bark and twigs, which are a supplement to their usual diet of grasses.

History of the European Bison

European Bison in the UK

There is no confirmed record of European Bison in the UK, although it lived in France as late as the 14th century, and fossils of its close relative, Steppe Bison, have been found just off the British coast. However, potential European Bison bone fragments which have been discovered are almost impossible to identify, as Bison DNA and bones are very similar to those of cattle.

We cannot confirm whether European Bison were present on the UK mainland from the fossil record. The similarity of bison bones and DNA to their cattle equivalents often make a true identification impossible (Rüther et al 2021). A lack of evidence doesn’t necessarily mean it wasn’t present, just that fossils haven’t yet been found or identified.

Collagen analysis shows that remains of animals in the Bos or Bison genus (containing both Bison and cattle) were being washed into a cave south of Sheffield over 12,000 years ago (Buckley et al 2017).

However, ‘Bison art’ found in the same cave system at Church Hole was dismissed by Wilson (2018) as likely to be a trick of the eye caused by an unusual rock formation.

In 2000, an investigation began into the seafloor between the UK and the Netherlands. This region, referred to as ‘Doggerland’, was once above sea level and formed a land bridge with Europe. This study found fossils of now-extinct Steppe Bison (Bison priscus) alongside other creatures typical of the Wooly Mammoth’s habitat (Mol et al 2006). However, European Bison was not found.

European Bison were recorded in North East France until the 7th century and in the Ardennes and Vogues regions until the 14th century, as reported by a IUCN review (Pucek et al 2004) – its historic presence in France is confirmed by an old article (Ahrens 1921). This places it within our geographical region, and the land bridge of Doggerland provides a past opportunity for population exchange.

European Bison Extinction

As humans moved into Europe in the ancient past, they pushed Bison from grassland into forests. These habitats became the last refuge of the species, where they were eventually wiped out in the early 20th century by hunting. The last wild European Bison went extinct in 1927 in the Soviet Union.

As humans moved into Europe, their activity increasingly pushed Bison towards woodland habitats to escape predation. This shows up in the isotopes of bone fragments from Bison killed at different points in the past – Early and pre-Neolithic Holocene European Bison ate vegetation characteristic of grassland, whereas Neolithic and Late Holocene animals had carbon and nitrogen isotope values more characteristic of forests according to a 2019 study (Hofman‐Kamińska et al 2019).

According to an old article (Ahrens 1921), despite having legal protection in Prussia as early as the 18th century, poaching, habitat loss and hunts continued to bring down the population of wild Bison. The Germans issued an order protecting the ‘Wisent’ in the Białowieza ˙ Primeval Forest at the start of WWI, when only 160 remained in this final stronghold. Numbers increased over the course of the war, but in December 1918, ‘retiring German soldiers [and] inhabitants’ shot ‘all  or nearly all’ of the remaining animals.

By the 1920s, the only remaining herd was in the Soviet Caucasus, and the final wild animal was killed in 1927, leaving only 54 animals remaining in zoos (Kuemmerle et al 2011).

European Bison Reintroductions

After the last wild European Bison went extinct in 1927, a captive breeding programme of European Bison from zoos saw numbers recover from a founder population of just 12 animals. This ‘bottleneck’ has led to some inbreeding issues.

Wilder Blean, the first UK reintroduction programme for European Bison will start in 2022 at Blean Woods in Kent.

A successful captive breeding programme saw the numbers of European bison recover, with reintroductions beginning in the 1950s (Rewilding Europe). By the 2010s, there were about 3000 European Bison, of which 1500 were living in the wild across about 30 herds in Central and Eastern Europe (Kuemmerle et al 2011).

However, the whole population went through a ‘genetic bottleneck’, being entirely descended from just 12 individuals. This makes them vulnerable to disease, with the lost genetic diversity resulting in ‘inbreeding depression’ in the Lowland-Caucasian line of bison (Tokarska et al 2011).

The first UK reintroduction programme will take place at the West Blean woods nature reserve in Kent, in Spring 2022. This exciting project, run by the Kent Wildlife Trust, will measure the impact of the European Bison with an extensive ecological surveying programme to detect changes in biodiversity and habitat structure.

Ecology of the European Bison

Territory Size

European Bison don’t defend a territory, but they range across an area of over 100km² – larger in the absence of human activity.

Our knowledge of the range of European Bison comes from modern day herds, which are closely monitored. According to the IUCN (Pucek et al 2004), their ‘territories’ overlap and are not defended.

Kuemmerle et al (2011) found a median herd ranged over a territory of 106km² or just over 10,500 hectares – an area about the size of Sheffield. This number was backed up by a study in Lithuania, which found a territory size of 100-120km² (Balčiauskas 1999).

However, modern territories are limited due to the small size of wilderness areas, and there was a large variation in the 2011 study, with some territories much larger than the median.

Diet

In their natural habitat, Bison mainly eat grass (65-85% of diet), with leaves from Spring to Autumn (up to 20%). They also eat bark and twigs in Spring and Winter (up to 20%).

An adult European Bison eats up to 60kg of fresh food every day, with calves eating 9kg daily and young adults from 20-29kg. It can take up to 14 days for food to pass through an adult’s gut.

There is now strong evidence that by the early 20th century, bison no longer lived in the habitats for which their bodies evolved. For example, isotopic analysis of ancient bones showed that their diet was typical of open habitats, not dense forest (Bocherens et al 2015). Forest-dwelling herds in Poland have traditionally been fed hay in winter to reduce the likelihood of starvation (Pucek et al 2004).

European Bison now largely inhabit forested areas (though this has recently begun to change), as this is where the last herds survived extinction at the beginning of the 20th century. This means that the diet recommended by the IUCN conservation action plan (Pucek et al 2004) was based on a misunderstanding of the species’ preferred habitat. However, a 1972 study recognised that 67% of their summer diet was made up of grasses, sedges and herbs (Knez 1972).

A well-designed recent study assessed the actual diet preferences of bison when housed in the diverse habitat of the Kraansvlak Bison Project, where a wide variety of food types are available (Cromsigt et al 2018). While the proportion varied over the year, over 60%, and up to 85%, of the diet was grass, with the remaining share made up largely of the leaves, bark and twigs of woody plants. Of these, leaves were the largest proportion particularly in Summer and Autumn (about 20% of total diet), except in Winter, with bark and twig consumption peaking in Spring and Winter (up to 15% of total diet combined). Herbs made up a very small remainder (<10%) of the diet, mostly in autumn.

The 2004 conservation action plan suggested that adult Bison typically eat 15-22g of food per kg of body mass per day. The exact figure depended on the age of the animal and the type of food. The literature reviewed in this action plan contained a variety of figures, from 23kg to 60kg per day for an adult European Bison. Young adults could eat 20kg to 29kg per day, while calves were expected to eat 8.5kg of ‘fresh food’ daily (Pucek et al 2004).

The Bison’s digestive system appears to work very slowly, with a study from the 1960s finding that dye added to food taking 8-14 days to finish showing up in dung (Gill 1967).

Habitat

Bison naturally inhabit an open mosaic of grassland, shrubland and deciduous woodland. Their bodies appear to have largely evolved for a life in scrubby grassland, but over time human activity caused them to seek refuge exclusively in dense woodland. This led to Bison being wrongly classed by the 20th century as purely a ‘forest species’. 

As stated in Diet, analysis of isotopes in ancient European Bison bones has shown that these creatures are native to open habitats, rather than forest (Bocherens et al 2015). However, as the last wild bison to survive extinction were found in dense woodland, scientists, even into the 21st century, recommended that ‘the most suitable habitats [are] deciduous forest types’ (Pucek et al 2004).

A study in a French Bison enclosure examined movement patterns, where feeding racks were located between conifer woodland and wet meadow habitats. The animals spent an overwhelming majority of the day in the more open area, with less than 20% of time in the forest (Ramos et al 2016).

An analysis of Bison’s dental characteristics relative to a very large collection of other mammals placed them unambiguously in the group of grazers typical of grasslands, at the opposite end of the scale to forest-dwelling browsers (Mendoza 2008).

However, Massilani et al (2016) point out that the complementary fluctuations in past populations of European Bison and Steppe Bison during periods where woodland cover varied suggests that the former species is more of a woodland specialist than the latter. The lower head position of the Steppe Bison also makes grazing easier, while browsing of shrubs is easier for European Bison (Massilani et al 2016).

Newborn calf and maternal bison behaviour is typical of a species that inhabited open habitats, rather than closed woodland. The calf spent its first week of life close to its mother, with no hiding phase that would be characteristic of forest-dwelling mammals (Daleszczyk 2004).

The preference for grasses revealed in ‘Diet’ shows that grassland/meadow is required for European Bison to thrive. Indeed, Bison which were not fed hay over winter in dense forest habitats were more likely to die in Poland (Pucek et al 2004). However, they also seek out leaves, bark and twigs in the colder months, even when grasses are abundant, indicating that they are not a purely grassland species and instead rely on a mosaic/mixed habitat to survive (Cromsigt et al 2018).

A review of the Bison’s incorrect classification as a ‘forest species’ concluded that their use of forest habitats would have increased in response to predation by humans (Kerley et al 2012). Given that we drove this species to extinction in the wild by 1927, perhaps it isn’t surprising, then, that the final remnants of the species were exclusively found in forest habitats.

Bisons’ strong preference for the shade of deciduous woodland in hot conditions revealed in unpublished data (Pape 2015) means that this habitat forms an important part of their environment, allowing them to stay cool in temperatures over 27.5°C.

European Bison are well-adapted to snowy terrain, able to graze even in deep drifts, pushing aside 30-40cm of snow with their heads to get to the vegetation beneath (Baskin & Danell 2003).

Rewilding the European Bison

Habitat Engineering

Bison usually feed on grasses and herbs, but their unfussy habit of grazing tree saplings, and occasional browsing and debarking of shrubs and trees can prevent meadows turning back into woodland. 

There is a common belief that Bison break up woodland like Elephants, by knocking over and uprooting trees, but the evidence does not support this. Instead, they likely move in to areas opened up by natural and human-caused disturbances like storm damage and forest fires, then keep these areas open with their browsing and physical activity. 

Bison may also open up woodland over very long periods by eating young saplings before they grow up into the canopy. Other large European herbivores have been shown to do this in experimental trials.

A study of Bison and other large herbivores in the Białowieza Primeval Forest found that the number of tree saplings in meadows was ‘substantially reduced’ by Bison grazing. The same effect was not found for other ungulates like Red Deer, and the authors suggested that this was likely due to the lack of ‘pickiness’ in the Bison’s feeding strategy. The high density of European Bison in this area was noted and it is not known whether this effect would be as marked in lower density populations. When Bison visited more often, they also significantly reduced the density of shrubby vegetation by browsing leaves and branches, stopping the edges of meadows from turning back into mature woodland (Kowalczyk et al 2021).

A well-written but critically flawed MSc thesis (Oquinena Valluerca 2011) examined satellite maps and GPS tracking of the Bison enclosure at Kraansvlak between 2003 and 2009, and found that areas of high Bison activity were associated with a reduction in woody vegetation, an increase in short grasses and a reduction in tall grasses. Low activity was associated with reduced bare sand cover. However, the area was only inhabited by bison from 2007, with Konik Horses and Scottish Highland Cattle living on the land for the previous 4 years – this study was not peer-reviewed and the conclusions, lacking proper experimental design, are not scientifically valid. 

However, a well-designed follow-up study at Kraansvlak examined habitat change over 8 years in 3 neighbouring areas. This found no impact of Bison on woody vegetation cover. The study compared the decline in woody plant vitality, grass height and cover across areas grazed by Bison, Cattle or wild Deer. Bison did not appear to significantly influence habitat structure relative to these controls, contrary to expectations. In fact, more Spindle Tree debarking from deer was found in the exclusion zone than in the Bison zone, although Sycamore was only debarked in the Bison area (Valdés-Correcher 2018).

A  study of sites in Zubryatnik and Polushkino where Bison have been reintroduced for 15 years claimed to ‘quantify [Bisons’] influence… on the structure of the vegetation’. But the experimental design was not longitudinal, with all data gathered in one month, and no exclosures or enclosures were used. Without any effective controls, the conclusion that the ‘forest vegetation has been significantly transformed by bison’ is not valid, as it fails to account for the strong likelihood that Bison selectively inhabit areas with this vegetation structure (Ivanova et al 2018).

However, there is some evidence that debarking might cause loss of mature trees due to fungal infection, e.g. 80% of beeches are infected with white rot from debarking on ‘Wisent Island’ in Germany. This may be a possible mechanism for opening up woodland, but a habitat change was not documented and deer were jointly responsible for the damage (Kelterborn et al 2009).

There is a common belief that Bison are able to transform woodland habitat by the physical action of knocking down or uprooting trees – and they have been observed ‘breaking through vegetation… and opening up scrubland’ (Vlasakker 2014). However, a key forest conservation textbook asserts that this is not the case; ‘the role of large ungulates on forest structure during the early Holocene was negligible’; instead, large ungulates simply keep gaps open, whether these gaps are human-caused or natural. Disturbances like forest fires, storm damage and waterlogging all contribute to the creation of these natural gaps (Honnay et al 2004).

However, Vera (2000) suggests that large herbivores slowly reduced forest cover over time by browsing out new saplings in woodland areas, gradually turning woodland into ‘parkland’. This recruitment method would take place over decades, or hundreds of years, reducing ‘recruitment’ of new trees and turning the landscape into a mosaic of scrubland and open woodland patches. 

A well-designed experiment (Smit et al 2015) in the Netherlands on land grazed by Konik ponies, Heck cattle and Red Deer confirmed that no experimental saplings survived the study period on this land, while some saplings survived in fenced-off control areas (exclosures). These data suggest that, while Bison are not necessary to prevent recruitment of new trees, as a large herbivore, they may play a role in reducing new canopy formation. However, the authors point out that the grazing density of herbivores in this area was high, with no natural predators (probably unnnaturally-so).

Biodiversity

Bison are very effective at spreading plants across a landscape, especially deciduous woodland and open grassland species. Their dung can contain over 100 species of plant seeds and their hooves can also carry over 30 types of seed, driving them deep into the soil in a protective ‘mud package’.

Bison may also have an impact on insect biodiversity, especially where they intensively graze meadow habitats.

A study of Bison dung found that these deposits contained an abundance of deciduous woodland seedlings, left in a coniferous habitat. While the paper didn’t examine how successful these seedlings were over a longer period, it was expected that in the long term, the dung could cause a change in habitat. Patches of deciduous vegetation were seen throughout the conifer wood, which was consistent with this idea (Jaroszewicz et al 2008).

Bison not only disperse seeds in their dung, but also in the material attached to their hooves (and, to a lesser extent their fur). A study of these seeds found that they were overwhelmingly different species to those transported in the digestive system (Schulze et al 2014). Seeds mainly originated from open habitats, and were transported into closed woodland. The bison’s hooves not only trampled vegetation, embedding the seeds into deeper soil layers and creating new exposed areas of ground, but also surrounded seeds in a ‘package of mud’ which was thought to improve their chances of germination. Dung further enriched the soil, which could again increase a seedling’s odds of survival.

The dung of forest-dwelling mammals, including Bison, was examined by Jaroszewicz et al (2013). While the most plants were found in Red Deer dung (137 species), this is likely to have been due to a lack of European Bison dung samples, which was a close runner-up. Bison deposited 109 different taxa of plant seeds in their dung, compared to just 24 in Wild Boar.

A recent study, designed to find any impact of Bison grazing on Carabid Beetle populations found no effect on total numbers. However, the increase in grazing caused a slight increase in the number of species of beetle (Schwerk et al 2021).

Tree Damage

Bison are known for ‘debarking’ trees; however, less than 2% of their tree damage is to the trunk bark, and they mostly eat branches, leaves and twigs. In a tiny proportion of cases (0.2%) they will uproot a tree. Debarking may cause tree loss when fungal infection occurs.

European Bison can cause damage to commercial tree crops with their ‘debarking’ behaviour, where they strip the bark from trees for its nutrition value. However, it is likely that this behaviour is simply a survival strategy in the absence of their appropriate diet, as providing hay fodder to other herbivores has been shown to reduce debarking, although tree damage is typically higher in the vicinity of feeding sites (Schröder et al 2019).

An examination of tree damage across a 180x100m transect within a larger Bison enclosure found that, of 4,701 trees damaged, just 10 were uprooted (Cătănoiu & Răzvan 2008). Over 98% of damage was the eating of branches, leaves and twigs, reducing tree vigour rather than removing individual trees. Debarking was relatively infrequent, occurring in just 1.4% of cases.

However, a long term study of tree damage in a German Bison reserve found that 80% of beeches had succumbed to white rot after being bark-stripped by Bison and other large forest-dwelling herbivores (Kelterborn et al 2009).

Body of the European Bison

Weight

Calves are born at 24-28kg, growing to adult size by age 6. An average adult male weighs 660kg, while adult females weigh 430kg.

The most recent data on European Bison body size comes from two Polish studies, both of which recorded data from forest-dwelling animals. However, this may not reflect the true potential size of Bison, as, for example, African forest elephants are smaller than their savannah-dwelling relatives (World Wildlife).

With this caveat, the IUCN report (Pucek et al 2004) states that the gender differences in body mass seen in adults are barely insignificant at birth, with calves averaging 24 – 28kg at birth. Calves grow most rapidly for the first year, but growth drops off significantly at age 3, and the animals reach full adult size by age 6.

Average adult male body mass ranged from 630kg to 750kg across two studies, with adult female body mass ranging from 420kg to 460kg. Averaging data across these two studies, the females’ mean mass was 431kg, while males’ mean mass was 661kg.

Size

Adult males grow up to 300cm long, and 188cm at the shoulders, with females reaching 270cm long and 167cm at the shoulders.

Adult male European Bison grow up to 300cm in length, with a girth of 280cm and a height at the withers (shoulders) of 188cm.

Adult female European Bison grow up to 270cm in length, with a girth of 246cm and a height at the withers (shoulders) of 167cm.

These figures are for bison both free-ranging in the Białowieza ˙ Primeval Forest and associated reserves.

(Pucek et al 2004)

Age

European Bison may live up to 24 years in the wild and 28 in captivity. Typically, females live longer than males.

Data from Poland in the 1970s showed that captive bred bulls lived up to 20 years, while their wild counterparts last 14-16 years. Captive bred cows lived up to 28 years, while the longest-lived wild cow on record was 24 years old (Krasiński 1978).

Reproduction

Bison typically have one calf at a time, giving birth after 264 days, from May to July. Adults reach maturity at 4 years old and, while European Bison (Bison bonasus) are a distinct species, they can breed with American Bison (Bison bison bison) and make fertile offspring.

Cows reach reproductive maturity in their third year, giving birth for the first time in their fourth year. They may continue giving birth until the end of life, but typically the upper limit is 18-20 years (Pucek et al 2004).

Bulls can reproduce from age 4-6, but are usually prevented from doing so in the wild by other males until around age 6. They continue reproducing until the age of 12. The rutting season occurs fro August to October (Pucek et al 2004).

Various studies of gestation have found different average durations, from 264 days to 267.4, as noted in the IUCN report. Twins are uncommon, with most mothers carrying one calf at a time. Cows lie down to calve and the newborn suckles within the first hour of life. Typically, calving happens from May to July, although it has been observed as late as October (Pucek et al 2004).

European and American Bison can interbreed and produce fertile offspring, revealing their close genetic similarity (Massilani et al 2016).

Temperature Range

Ideal range from 5°C to 27°C, but European Bison adapt to extreme heat by seeking out shade and resting, or warm up by travelling long distances across dry, sunny grasslands.

With their natural range across continental Europe, Bison are accustomed to a very broad range of temperatures, including severe heat and cold.

Unpublished data from a PhD thesis (Pape 2015) suggests that, when dealing with extreme temperatures, Bison showed behaviours which allowed them to adapt by varying the heat they created from exercise. GPS collars tracked them travelling much longer distances below 5°C and resting more often when the heat rose, especially above 27.5°C, which appeared to be a threshold for strenuous activity.

The study also found that, as the temperature changed, so did the animals’ preferred habitat – they sought out the shade of deciduous forest, with this making up ~20% of the range at 22-26°C, but ~60% of their range from 37-41°C. In colder temperatures, animals spent the majority of their time in dry grassland, where exposure to sunlight could provide a warming effect.

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