These conclusions suggest that the indigenous herbivores of the
Highlands are likely to keep the landscape open. It is a situation we
are unfamiliar with in Europe as a whole because we have made
most such species extinct or manage them to very low population
levels. For example, the environmental historian Kaplan says that
removal of large animal species by humans has had effects on the
landscape that are apparent almost everywhere:
“A lot of land would be semi-open, kept partly open by these big
herds of grazers and browsers and predators. It is important to keep
in mind that landscape is also shaped by animals. These giant herds
of bison would be trampling down little trees and keeping the
landscape open (Kaplan 2013).”
Perhaps within Europe it is only the Scottish Highlands where free-
ranging herds of indigenous large herbivores have been present
throughout the postglacial period that we can observe their role in
Hence the Scottish hills are not treeless through ‘overgrazing’ which
is the current opinion of many conservation organisations: the
grazing level of 4-8 deer km
necessary to maintain woodland in
the landscape is a low grazing level according to the trophic level
model. To quote Milner et al. (2002) in reference to the Highlands:
“Overgrazing is a controversial term ... Its precise definition is
dependent on management objectives ... Overgrazing is not
generally applied to natural ecosystems, even under heavy grazing
pressure, because wild herbivores are regulated by their food supply
during the unfavourable season ... For example, there is no evidence
of habitat degradation on St Kilda or the North Block of Rum where
herbivore populations are naturally regulated.”
2) What is the expected natural distribution of woodland at
this stage of the postglacial cycle?
In northwest Europe during interglacial periods there is recognised
plant succession from an initial protocratic phase following ice
retreat, through a mesocratic to an oligocratic as illustrated in
Figure 3 (Birks & Birks 2004). This succession has taken place in
previous interglacials in the absence of humans.
The middle mesocratic phase is optimal for woodland with the
highest percentage of the landscape covered with trees, a
percentage which thereafter declines.
The succession is brought about through long-term changes in
both soil and climate, with the availability of soil nutrients,
particularly phosphorus and nitrogen, reducing over time through
leaching. In the past, the successional changes finished with a
return to glacial conditions (the cryocratic phase), although
anthropogenic global warming means that this is unlikely to be the
case in the future.
Scenarios of woodland survival
For woodland to persist in the light of these long-term changes,
conditions have to be permanently favourable for the survival of
seedlings and saplings. Given below is a list of conditions which
would enable woodland to persist in the European landscape (i.e.
young trees can survive browsing) assuming indigenous large
herbivores are present (and/or free-ranging domestic analogues) in
accordance with the theory of trophic levels discussed above.
Looked at logically from first principles, woodland will remain
or become common in the landscape if:
Optimal soils/climate allows numerous seedlings to
germinate and grow so fast that the probability of some
surviving browsing is high: e.g. mesocratic phase (and also
likely to be the case in the tropics).
Young trees are protected from browsing by thorny shrubs:
temperate forest – the Vera hypothesis (Vera 2000).
Young trees are protected by snow in winter and early
spring, the period when seedlings and saplings are most
vulnerable: e.g. boreal & sub-alpine forests, sub-alpine &
Tree morphology or toxicity reduces browsing: e.g. spiky
needles of spruce, Rhododendron ponticum.
The anthropogenic activities of removal or reduction of
grazing: e.g. compartmentalised broadleaved woodland
(browsing animals significantly reduced or excluded).
Woodland may remain localised in the landscape:
Where young trees are protected from grazing by natural
features such as cliffs, boulder fields, gullies, or,
occasionally, dense vegetation such as tall Calluna vulgaris.
Episodically, owing to temporary reduction in grazing
through extreme weather or disease events.
Locations commonly visited by predators.
Each of these eight scenarios is now discussed in more detail.
Woodland dominant or common in the landscape
A) Optimal soils/climate
In conditions favourable for tree growth with abundant production
of viable seeds and seedlings, then there is a greater probability of
woodland persisting in the landscape (Figure 4(A)). It should be
noted, though, that optimal conditions for trees are also likely to
result in optimal conditions for the associated herbivores so it may
not always be the case that optimal conditions for tree growth
result in closed high forest without the help of the other factors (as
discussed below). But with optimal conditions, there will be a
higher probability of at least some woods surviving in the
landscape whatever the grazing level.
As the current interglacial progressed in the wet climate of the
Scottish Highlands, with its generally base-poor bedrock, continual
leaching resulted in soils too acid for earthworms causing the
formation of unmixed, stratified soils, often underlain by an
impermeable ironpan. The presence of a thick humus layer
together with the ironpan makes the soils liable to waterlogging
and has resultedin the large-scale development of blanket peat on
land with a gentle gradient. This characterises the natural
succession from the mesocratic to the oligocratic phase (Figure 3).
Thompson (2004) states that such ‘a combination of very low soil
nutrient availability and high soil moisture provides very
unfavourable conditions for colonisation of birch (Betula), rowan
(Sorbus aucuparia) and Scots pine (Pinus sylvestris)’. Hence the
Birks HJB, Birks HH. 2004. The rise and fall of forests. Science
Kaplan JO. 2013 November 16. New Scientist. p.37.
Milner J, Alexander J, Griffin C. 2002. A Highland Deer Herd and its
Habitat. London: Red Lion House.
Thompson R. 2004. Predicting site suitability for natural
colonisation: upland birchwoods and native pinewoods in northern
Scotland. Edinburgh: Forestry Commission Information Note.
Vera FWM. 2000. Grazing Ecology and Forest History. Wallingford:
WOODLAND OR OPEN GROUND? Scenarios for the persistence of woodland in the presence of grazing
Red deer grazing on Scottish moorland
Figure 3. The interglacial cycle in northern latitudes. The
mesocratic phase is optimal for woodland, which thereafter
declines in the oligocratic. Anthropogenic warming may mean
that there is no return to a cryocratic phase.
From Birks and Birks (2004)
https://doi.org/10.1126/science.1101357. Reprinted with
permission from AAAS.
Figure 4. Diagrammatic examples of seedling and sapling
establishment and growth, showing why tree regeneration is
more susceptible to browsing in the oligocratic phase of
interglacials, and hence the probability of woodland surviving
in the landscape decreases. Growth rates are indicative only.
(A) In the mesocratic phase optimum soils and climate results
in numerous seedlings becoming established which
subsequently grow fast and have a higher probability of
(B) In the oligocratic phase conditions are less suitable for
seedling establishment and any that do establish have a
higher probability of being browsed owing to their slow