(Sub)tropical Grasslands and

Rangelands
Mediterranean sylvopastoral
systems and savannas
Mediterranean biome
• Subtropical climate:
• annual rainfall about 500 mm
• Hot dry summer x mild rainy
winter

chapparal Mediterranean
basin

matorral
phynbos
Mallee shrublands (eucalyptus)
Kwongan heathland
• SCRUBLANDS : the driest areas, near the seacoast where wind and salt spray
are frequent.
• SHRUBLANDS MACCHIA, MATORRAL, dense thickets of evergreen
sclerophyll shrubs and small trees, the most common plant community,
mature vegetation type, result of degradation of former forest or woodland
by logging or overgrazing, or disturbance by major fires
• WOODLANDS dominated by oak and pine, mixed with sclerophyll and
conifers
• FORESTS highest rainfall and along rivers (summer water), evergreen trees
(oak, pine)
Grazing as driver of deforestation
In more marginal, stress-prone regions and
mountains, where agriculture was hardly
viable, extensive pastoralism has been the
most common and traditional land use.
SILVOPASTORAL SYSTEMS
a combination of trees, pastures or crops
and livestock on the same land

Grazed
Fruit Orchards
Silvopastoral systems
• a combination of trees, pastures or crops and/or
livestock on the same land (Mosquera-Losada et al 2012).
• Trees and shrubs: provide environmental externalities, useful
products, although their full potential is still unknown. Wood
vegetation is compatible with pasture production or act as a
forage complement in SPs.
• provide efficient feed conversion, better animal welfare and
source of environmental services and public goods (e.g.,
more positive carbon balance, highly appreciated landscapes,
and higher biodiversity
• cultural heritages, the basis for unique natural commercial
products, included high-quality foods
Silvopastoral systems
• a combination of trees, pastures or crops and/or
livestock on the same land (Mosquera-Losada et al 2012).

The effect of trees on pasture is controversial …

• Positive effect of trees on pasture yield and

nutritional quality, length of growing season, and
stability against climatic variability,
• Negative effects of trees due to complex
interactions that occur among trees and pastures,
with a dominant role of water competition in dry
areas and for light in high latitude regions.
TREE – GRASS – SOIL INTERACTIONS

• Structural complexity
• low-light, cooler microhabitats, forage, shade for
animals, roosting sites for birds, increase of
biodiversity
• Heterogeneity
• higher nutrient concentrations under crown
• Productivity of the grass understorey
• Improved soil fertility and structure beneath the
crown
• Improved water relations of the shaded grasses
• Competition for light, water and nutrients
TREE – GRASS – SOIL INTERACTIONS

Soil nutrient enrichment

Altered species composition
Enhanced understorey productivity and tissue nutrient
concentrations
Reduced soil and plant temperatures
Reduced solar radiation
 Multipurpose ekosystem: Goods
provided by silvopastoral systems
Forage: Herbaceous Pasture
Browse
Fooder trees
Fruits: Acorn, Chestnuts …
Firewood, charcoal
Materials: Cork
Timber
Fruits

French pre-vergers
Dutch boguards
Spanish pomaradas
Centroeuropean streuobstwiesen
 Goods Provided by silvopastoral systems: Forage
FODDER TREE-FRUIT: Acorn,
Chesnut…

The specific diversity of livestock use efficiently the diversity of pasture

Goods Provided by silvopastoral systems: Forage
Acorns Browse Grass
100
Seasonal use of
80
forage resources
% of distinct fooders

60
(San Miguel et al, 1996)

40

20

0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Goods provided by silvopastoral
systems: Bioenergy

Fodder trees:
prunned branches Firewood

Charcoal
The important contribution of agroforestry
systems to multiple ecosystem services:
Microclimate amelioration
Soil fertilization
C sequestration and reduction of GHGs emission
Water quality
Biodiversity
Reduction of fire risk
Ecosystem services: Microclimate
amelioration
• Trees reduce radiation for pasture understorey but they improve
microclimate
• Although shade usually has detrimental effects for pasture
production in high latitude under dry conditions the reduction of
excess of radiation could become an advantage, and pasture
production be higher under certain level of shade
• Agroforestry systems may be more resilient to climate change, as
the tree cover may reduce some concerning events such as early
heat and drought stresses on herbaceous understory
• Shelter provided by trees is a key element for animal welfare (cows
kept cool produce more milk under better animal welfare
conditions)

Moreno et al 2007. Agroforestry systems

Video
• https://www.youtube.com/watch?v=kqdYl5QfEio
Ecosystem services: Microclimate amelioration
Animal Welfare

50
ST1m ST20m
40
Temperature, ºC
30

20

10

0
ENE
0 FEB
500 MAR1000ABR MAY
1500 JUN
2000 JUL 2500

Moreno et al 2007. Agroforestry systems

Ecosystem services: Carbon
sequestration and reduction of GHGs
emission
• Methane production by ruminants, ammonia and nitrous
oxide production by all forms of livestock production and
emissions related to feed production
• Europe’s meat and dairy consumption = 14% of the total
CO2 emissions in the EU
• The integration of trees with livestock = method to
mitigate ammonia emissions, and to store carbon as an
offset for methane and nitrous oxide production
• C pool is significantly higher in the sylvopastures than
pastures
 Ecosystem services: Carbon sequestration and
reduction of GHGs emission
• Spain: soil C was almost doubled
beneath tree canopies compared
to open pastures. The higher inputs
of residues generated by trees in
SPs than in tree-less systems may
cause high soil C sequestration
potential, but soil C increase
depends both on the incorporation
and mineralization of C in the soil.
The latter is affected by understory
crop management practices
• Long-term effects of improved
pasture establishment in 67
Portuguese montados caused
higher organic-C than those under 94
unmanaged pasture, and the
effects were stronger in the
presence of oak trees
137 g C kg-1 soil
Gómez-Rey et al 2012

(Howlett et al 2011. J. Env. Monit)

 Ecosystem services: Soil fertilization

• Tree roots bring up nutrients from deep in the soil profile that is inaccessible
to herbaceous vegetation so more than 50% of the nutrients could be
annually recycled beneath the canopy despite the canopy cover could be as
low as 20%
• The turnover rate of nutrients on the soil surface of SPs is unusually high
• For instance, dehesa litterfall decomposes up to 24 times faster than in dense
forest due to the herbivores action
• Trees also play a prominent role in the process, as net mineralization is higher
beneath than beyond the canopy cover
• As a result of the nutrient dynamics in SPs, soils are more in soil organic
matter (SOM) and nutrients beneath the tree canopy than beyond the
canopy
• Trees frequently have a positive effect on pasture production especially in
unfertilized oligotrophic soils
Ecosystem services: Soil fertilization
x
25 N Ca Mg P K 250 2,5
Nutrient content (N, Ca, Mg), g / kg

20 200 2,0

x
K content g / kg

P content g / kg
15 150 1,5

10 100 1,0

5 50 0,5

0 0 0,0
0 5 10 15 20 25
Distance to the tree trunk, m

And trees increase the pool of available nutrients

Ecosystem services: Water quality

• The increased losses of nutrients, farm effluents (e.g. livestock wastes),

pesticides such as sheep-dipping chemicals, bacterial and protozoan
contamination of soil and water are some of the main concerns
regarding water quality degradation
• There has been a general uncoupling of nutrient cycles, with frequent
problems of unacceptable high nutrient deposition or uneven spatial
distribution
• Different studies have showed the capacity of deep rooting trees to
capture nutrients at deep soil layers, where the herbaceous plants do
not reach, and thus reduce nutrient leaching in SPs
• For example: integrating pigs in a willow/miscanthus plantation results
in a much smaller risk for N leaching than typically seen on grassland
with the same stocking rate and that the pigs only caused limited
damage to the trees

M. L. López-Díaz, V. Rolo, and G. Moreno

Journal Environmental Quality 40: 853–859 (2011)
Ecosystem services: Water quality

= Bare soil;
= MonoPasture;
= Tree;
= Wood-Pasture

M. L. López-Díaz, V. Rolo, and G. Moreno

Journal Environmental Quality 40: 853–859 (2011)
 Ecosystem services: Biodiversity

• Noticeably high diversity of vascular plants, birds, mammals, lizards and butterflies
for Iberian dehesas compared to other adjacent land uses
• Similar results for vascular plants, birds, snails and beetles for other European SPs
• Increase in nvertebrate species and numbers reported when moving from open
grassland to agroforestry conditions for carbid beetles in Northern Ireland
• The tree-cover provides multiple tree-based gradients, in terms of light, soil
nutrients and moisture, food availability, refuge, even certain low level of
disturbance caused by uneven use of space by livestock = key factor for species
and niche diversity of SPs
• Trees are essential sources of food and refuge (For example: bat activity increased
by a factor of 100 with the presence of 3–5 trees).
• Habitat heterogeneity at multiple spatial scales has been revealed as key for
biodiversity conservation
Diaz (2008) and Marañon et al (2009)
Ecosystem services: Biodiversity TREE AS SOURCE of FOOD
for a PLETHORA of ANIMALS

MAMALS & INSECTS: TREE

REPRODUCTION
Hervibores
INSECTS: Leaf feeder

SAPLINGS
FEMALE
FLOWERS

SHRUBS

SEEDLINGS POLINIZED
FRUIT

MAMALS, BIRDS INSECTS: Furit feeder

DISPERSED MATURE
ACORNS ACORNS
BIRDS RODENTS
 WOOD vs OPEN PASTURES Ecosystem services: Biodiversity

Moreno
Moreno in prep.
et al et al. In preparation
Ecosystem services: Reduction of
fire risk
• Mediterranean rangelands and forests are prone to wildfire. However,
periodical wildfires may also occurs in some continental and atlantic
mountainous regions across Europe.
• Livestock feces, trampling and browsing can kill shrubs at a medium
term, although anti-nutritional components of woody vegetation may
difficult the efficiency of livestock as fireguard
• SPs more profitable than single timber production systems when fire
risk was included in the analyses.
• Forest grazing the best cost-effective treatment (6–30 € ha−1 per year
depending on animal type and management system adopted) x
investments (water supplies, fences…)
• Using combination of manual and mechanical treatments alone costs
200–300 € ha−1 per year, with a return interval of 3-to-5 years. Fuel
reduction technique by prescribed fires costs 600–1,000 € ha−1 and
cannot be used repeatedly without jeopardizing the nitrogen fertility of
the ecosystem Casals et al 2004).
Ecosystem services: Reduction of fire risk
RAPCA:
Grazed Fuelbreak Network in Andalusia
Silvopastures dynamics: Patterns
and Processes
• Different management of SP across the Europe
• Woody vegetation hardly managed and/or management did not
follow underwent ecological and economical changes
• Declining processes as lack of regeneration and exhaustion of
resources
• Mountain areas abandoned, low value of the traditional products in
the actual markets
• Extensive SPs result from a simplification, in structure and species
richness, of native forests, and are attained by tree clearance,
eliminating of shrubs, and favoring grasses by means of grazing
and occasionally forage sown
Silvopastures dynamics: Patterns
and Processes
• The landscape formed is maintained by a balance between
divergent ecological processes such as grazing pressure and tree
regeneration
• The coexistence of patches of pastures, woodlands and/or isolated
trees is a result of an unstable equilibrium leading either to closed
forests or to open pastures
• Adequate grazing pressure for the maintenance of sufficient but
not excessive tree/shrub regeneration is crucial for the persistence
of SPs
• Intensification = gradual deforestation due to the lack of tree
regeneration, as has been well documented for Iberian dehesas
• Undergrazing = loss of the characteristic open two-layer structure
of SPs, which tends to become dense forests or scrublands
Silvopastures dynamics: Patterns and Processes
EXTENSIFICATION

Two divergent trends

INTENSIFICATION
Challenges and New
Opportunities
Tree regeneration and Nursery plants

Ecological intensification

High Nature Value farming network

Carbon sequestration and Life Cycle

Assessments

Branding high quality products of SPs

Challenges and New Opportunities: Ecological
intensification
Animal feeding:
comercial supplementary fodder: high quality
versus
in-farm forage: diversification of forage, of environment,
mitigate seasonal shortage of commercial supply
 Challenges and New Opportunities: Ecological
intensification
The need of producing in-farm forage
• Woody Forage Bank = summer /winter diet of animals
• Highly water-use efficient fodder shrubs
• Intercroping by species: Medicago arborea, Atriplex sp., Acacia
spp., Morus spp., Ceratonia siliqua
Medicago arborea Morus Carob
Challenges and New Opportunities: Ecological
intensification
The need of producing in-farm forage
Livestock breeders are interested in
increasing the quality of the feed as
well as to diversify forage offer
(mitigating seasonal shortages)

Permanent
pastures rich
in legumes
Challenges and New Opportunities:
Ecological intensification
• Genetic base of currently sown pastures is very narrow
• More than three-quarters of the grass cultivars registered in the
European Union are of just six species
• Lolium perenne and L. multiflorum more than 80 percent of the
forage grass seed sold in the EU
• Availability in the seed market of pasture species suitable for dry
environments (e.g. semiarid Mediterranean areas) is still scarce
• Need of widening the choice of available high-value grass and
legume cultivars by exploring, evaluating and selecting from a wide
range of species of several genera
Challenges and New Opportunities: Ecological
intensification

The need of selecting shade-adapted forage crop

Results of ten years of screening forage species for shade tolerance that clearly
demonstrate that many cool-season forages benefit from 40% to 60% shade
when grown in Missouri. Grazing trials have proven to be successful at least in the
short-term (Adapted from Garret et al 2004).
Challenges and New Opportunities: Ecological
intensification
Planting trees
Challenges and New Opportunities: C
sequestration and Life Cycle Assessments

• The increasing intensification of many SPs, especially those in

Mediterranean regions, has produced a worrying loss of soil quality
• Identification of ecologically complex organic materials, and
assessment specific management requirements, is a vital issue for
ecosystem conservation or restoration
• Ramial wood and biomass charcoal: a “source” ecosystem (where
the organic material is produced) and a “sink” ecosystem (where
the organic material is applied)
• In Europe demand for the use of biomass crops dedicated to
bioenergy
• The opportunity to use (the excess of) wood biomass of SPs as
local or in-farm source of energy
• Carbon (C) sequestration and the specific humus quality in the soil
of arable agroforestry systems.
Challenges and New Opportunities: C sequestration and
Life Cycle Assessments
Challenges and New Opportunities:
High Nature Value farming network

• Agricultural intensification = decline in farmland biodiversity across

many different taxa
• Farmland also hosts many species that depend on appropriate
agricultural management for their survival
• At least 7% of the utilized agricultural area of each farm should be
allocated to ecological focus areas, which could include landscape
features, buffer strips or afforested areas
• Biodiversity conservation is more likely to be effective on farmlands
that are already managed at low intensity and that retain a certain
amount of seminatural vegetation
• These criteria mostly accomplished by extensive pasturelands that
consist of grassland, scrub or woodland or a combination of different
types, used for raising livestock
• Extensive pasturelands are dominant in the last European map of High
Nature Value farming
• (Oppermann et al 2012)
Challenges and New Opportunities:
High Nature Value farming network

Farm types and landscapes valuable for

biodiversity, always low-input farms,
managed at low intensity and that
retains semi-natural vegetation.
MOSTLY SILVOPASTURES.
Challenges and New Opportunities:
Branding high quality products of SPs
• Controlled certification processes
• Consumers can buy products with high environmental
added-value
• Pig meat production in the Iberian dehesas with local Iberian
pig
• Parkland systems in the UK of particularly high cultural value
• New needs for natural and high quality products derived
from extensive SPs: acorn-derived products (tannins for tan
leather and for antioxidant uses, gluten-free flours,
unsaturated fat …).
• Need for decision-making support tools, for joint
participatory actions supporting decision
Challenges and New Opportunities: Branding high quality
products of SPs
Video
• https://www.youtube.com/watch?v=vhprxJP7fhs
Challenges and New Opportunities: Branding high quality
products of SPs
Savannas
• Tropical seasonal ecosystems with a
continuous herbaceous layer, usually
dominated by grasses or sedges, and a
discontinuous layer of trees and/or shrubs
 Savannas
• Tropical hot climates: annual rainfall about 500
– 1270 mm
• SEASONALITY Wet season (summer) x dry
season (winter)
 Savannas
• The soil savanna is porous, with rapid drainage of water
• Thin layer of humus which provides vegetation with nutrients
• Sometimes classified as forests
• Different grasses due to disparities in rainfall and soil condition
• Usually only one or a few kinds of grass more successful in a
particular area
• Deciduous trees and shrubs are scattered across the open
landscape
• Southwestern Kenya, Tanzania, and Uganda, „grouped-tree
grassland“, trees growing only on termite mounds
• Frequent fires and large grazing mammals kill seedlings, thus
keeping the density of trees and shrubs low
Savannas
Ecological determinants
• Soils: mostly infertile, lateritic, lithosols, clays, porous,
rapid drainage of water, thin humus layer (organic
matter created by partial decomposition of plant or
animal matter)
Savannas in South America
• 45% of area
• Llanos: tall grass savanna with
evergreen broad-leaved trees .
• Cerrado: a mosaic of grasslands, tree
savannas and woodlands, with
patches of semi-deciduous forest.
• Pampa: subtropical deciduous
thicket, thorn woodland, subdesert
shrubland .
• Few deer, rodents (capybara), few
predators, 521 bird species
Savannas in Australia
• Around one-quarter of mainland Australia's land
area.
• The most common tree eucalyptus
• Oil, gum, and timber, tanine
• The foreign pests and weeds: wetland weed
Mimosa, threaten the survival of the region's
ecosystems and native species
• The Australian government programs to clear
away brush for agricultural reasons
• The World Wildlife Fund (WWF) in Australia is
reacting to this destruction by working with the
communities to stop the clearing programs
Savannas in Australia

• Eucalyptus spp.
• Dense grass and scattered trees and
shrubs
• Strong effect of fires
• Both seasons are very extreme
• 227 bird species
• Large herbivores = marsupials
 Savannas in Africa
• Covers 65% of area
• 708 bird species,
Sahelian • The most spectacular
Sudanese mammalian fauna – more than
90 species of ungulates, wide
range of large predators

Guinean

mopane, miombo, karoo, …

 Savanna:INTERACTIONS
Plant available moisture
• The main limiting factor for vegetation development
• Grass- tree interactions: two soil layers with independent water
budgets x grass and tree roots – strong competitors in the surface
soil
BUSH ENCROACHMENT
• If grasses are destroyed,
e.g. by heavy grazing,
more water becomes
available for trees and
shrubs, leading to
changes in the
interactive pattern and
to an increase in woody
vegetation
 BUSH
ENCROACHMENT
OF WETLANDS IN
SAVANNAS
Savanna:INTERACTIONS
Plant available nutrients
• Changes in soil nutrient content on the intermediate
and larger scale: SLOW PROCESSES (weathering,
erosion, rainfall regime, fire, herbivory)
• On small scale, e.g. the size of a buffalo urine patch,
changes in nutrient content: QUICK and can cause
disturbance
• Higher soil nutrient content is generally associated with
a better developed woody layer → the role of trees in
savanna
• Large herbivores speed up the nutrient turn-over rate in
savanna. Extinction / exclusion → a rapid loss of
nutrients from the surface soil.
Trees in savanna

• Structural complexity
• low-light, cooler microhabitats, forage, shade for
animals, roosting sites for birds, increase of
biodiversity
• Heterogeneity
• higher nutrient concentrations under crown
• Productivity of the grass understorey
• Improved soil fertility and structure beneath the
crown
• Improved water relations of the shaded grasses
• Competition for light, water and nutrients
 Soil nutrient enrichment
Altered species composition
Enhanced understorey productivity and tissue nutrient
concentrations
Reduced soil and plant temperatures
Reduced solar radiation
 FIRES
• Natural fires from lightning around the rainy season
• Man-made fires during the dry season (since millenia)
• After fire: resprouting – fresh resource for herbivores
• Frequent fires
• may lead to replacement of perennial grasses by ephemerals
• controlling woody seedlings and young trees
FIRES
LARGE HERBIVORES – ECOSYSTEM ENGINEERS

• Arid/ semi-arid savannas – low rainfall – herbivores

abundant only in rainy season, then migrate: dependence
upon water availibility
• Increasing humidity - interactive processes, e.g.
COMPETITION AND PREDATION – effect on dynamics of
populations and systems
• High degree of responsiveness to the vegetation: selectivity
on all scales, regular and irregular migrations and
demographic changes
• Competitive patterns of vegetation
• preferentially grazed species loose competitive
power and decrease
• Palatable and grazing tolerant species
become dominant
LARGE HERBIVORES
PLANT - ANIMAL INTERACTIONS
• ELEPHANT‘s engineering

• Elephant damage may suppress recruitment in preferred tree

species in woodlands
Haynes 2012
 Hayes et al. 2012

Large-mammal trampling reduces soil

macropore space (impeding water
infiltration and increasing surface
runoff)
Lowers root biomass and whole plant
biomass for many species, seriously
decreasing the fertility index of land
surfaces
• Large-mammal trampling reduces soil macropore space (impeding
water infiltration and increasing surface runoff)
• Lowers root biomass and whole plant biomass for many species,
seriously decreasing the fertility index of land surfaces
Coverdale et al. 2016
• Kenya: experiment
• Neutral effects on most 40 (83-89%) species, with
a similar frequency of positive and negative
responses among the 41 remainder.
• Elephants create associational refuges for
understory plants by damaging tree canopies in
ways that physically inhibit feeding by other large
herbivores
• Understory biomass and species richness beneath
elephant damaged trees were 55% and 21%
greater, respectively, than under undamaged
trees
• Experimentally removing elephant damaged
branches decreased understory biomass by 39%
and richness by 30% relative to no manipulated
trees
• Camera-trap surveys revealed that elephant
damage reduced the frequency of herbivory by
71%
LARGE HERBIVORES
PLANT - ANIMAL INTERACTIONS
• Grasses have about the same competitive power and tolerance
to defoliation and that they differ mainly in palatability
• Grazing lawns: preferentially and intensely grazed swards with
more palatable and nutritious grasses than in the less grazed
surroundings, grasses both palatable and competitive

• African savanna: many grasses fairly tolerant to grazing,

intensive livestock grazing - species turnover - reduction in
palatability of the sward

• South American savanna: indigenous dominant grasses are

often unpalatable and intolerant to grazing, more palatable and
grazing resistant species are confined to disturbed habitats
LARGE HERBIVORES

NUTRIENTS
LIVESTOCK GRAZING IN SAVANNAS

• The change in vegetation under heavy livestock grazing, i.e.

from dominance of perennial palatable grasses to annuals,
unpalatable dicots or dwarf shrubs
• Dietary overlap with wildlife species (zebra, wildebeest, buffalo)
• livestock and wild herbivores have not shared a long common
evolutionary history
• Wild: 5 million yrs ago
• Cattle: 3000 BP

• STRONG COMPETITION
in critical periods
Videos
• Savannas:
• https://www.youtube.com/watch?v=Mle5gmEpYys
• Shrublands:
• https://www.youtube.com/watch?v=wcYWS-5iUcc
• Grasslands:
• https://www.youtube.com/watch?v=Yy191KVBNP0
THANK YOU
FOR
ATTENTION!