Ruminant Contribution To Enteric Methane Emissions

Mitig Adapt Strateg Glob Change (2022) 27: 47
https://doi.org/10.1007/s11027-022-10026-5
Ruminant contribution to enteric methane emissions

and possible mitigation strategies in the Southern Africa
Development Community region
Mompoloki Seketeme1 · Othusitse R. Madibela1 · Thabo Khumoetsile1 ·

Innocent Rugoho2,3
Received: 5 October 2021 / Accepted: 17 August 2022 / Published online: 31 August 2022
© The Author(s), under exclusive licence to Springer Nature B.V. 2022
Abstract
The Southern Africa Development Community (SADC) region is not a major emitter of
greenhouse gases (GHG). However, Sub-Saharan Africa is considered a potential future
hotspot for GHG emissions because of its large livestock population dispersed across large
arid lands, coupled with the inherent low digestible feeds in the region and consequently
low productivity of livestock. In SADC, climate change is predicted to increase tempera-
tures further reducing agricultural productivity. Therefore, there is incentive to reduce
agriculture’s contribution to GHG emissions in the SADC region. Ruminant production,
a mainstay of rural economy, is predicted to decrease because of diminished grazing due
to reduced rainfall and feed quality. However, ruminants’ enteric methane (CH4) produc-
tion contributes to GHG emissions. This review explores strategies for the SADC region to
reduce CH4 by ruminants. As methanogenesis is an outcome of microbial activity, potential
opportunities include selecting animals with inherent low C H4 production; altering rumi-
nal microbial populations to those that do not yield CH4; enhancing feed digestibility by
feeding additives which improve diet quality and alter the ruminal microbiome and using
specific forages such as seaweed or duckweed that contain plant secondary metabolites that
may decrease methanogen populations or methanogenesis. These strategies are considered
in terms of their potential magnitude of CH4 mitigation, the practicality for their imple-
mentation in the SADC region and their suitability to be included in the grazing-based
livestock industries in the SADC region.
Keywords Grazed forage · Greenhouse gas emissions · Methane · Mitigation · Ruminants
* Innocent Rugoho
[email protected]
1
Department of Animal Sciences, Faculty of Animal and Veterinary Sciences, Botswana University
of Agriculture & Natural Resources, Gaborone, Botswana
2
Lely Australia Pty Ltd, Truganina, Melbourne, Vic 3029, Australia
3
YourFarmGenetics Pvt Ltd, Nharira View, Norton, Zimbabwe
13
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47 Page 2 of 26 Mitig Adapt Strateg Glob Change (2022) 27: 47
1 Introduction
Livestock production is an important socio-economic activity for most people living in

arid environments of the Southern Africa Development Community (SADC). An estimated
75% of livestock are kept under smallholder traditional farming systems (Southern Afri-
can Development Community 2012). Although ruminant animals (beef cattle, sheep and
goats) are the preferred livestock kept in this region, chicken, pigs, horses and donkey are
also farmed. Livestock populations in SADC are estimated at 64 million cattle, 39 million
sheep, 38 million goats, 7 million pigs, 1 million horses and 380 million poultry (Southern
African Development Community 2012). There is a relatively large population of ruminant
animals in the region mainly because they are fed on forages that are low-cost sources of
nutrients, and therefore, ruminants generally do not compete with humans for food. These
animals can convert low-quality forages into high-quality products such as milk and meat
which are valuable food resources for many people in the region. These ruminant products
supply essential nutrients that play a major role in proper physical and cognitive devel-
opment of infants, children and adolescents (Leroy et al. 2022) and help promote main-
tenance of physical function with ageing (Leroy et al. 2022). Furthermore, in the SADC
region, ruminants such as cattle provide non-monetised contributions (e.g. draught power
and manure), reflecting the importance of integrated crop-livestock farming systems in this
region (Otte and Chilonda 2002).
Forages contain variable nutrient composition, namely crude protein (CP), water-
soluble carbohydrates (WSC), neutral detergent fibre (NDF), acid detergent fibre (ADF),
fat and starch (Rugoho et al. 2016), and as they mature, they become lignified (Cherney
et al. 1991). Forages also contain anti-nutritional factors such as tannins (Cowan 1999).
The ability of ruminant animals to utilise forage is made possible by the ruminal microbes
that are responsible for fermentation processes (Castillo-González et al. 2014). Ruminal
microbes that carry out fermentation processes include bacteria, archaea, fungi and pro-
tozoa, and they break down feed anaerobically to produce fermentation products namely
microbial protein, volatile fatty acid (VFA), methane (CH4) and carbon dioxide (CO2)
(McDonald et al. 2011). The nutrients produced are also beneficial to both the microbes,
and to the host animal, and thus the rumen microbes are in a symbiotic relationship with
the host animal. However, the importance of animals as an efficient and economic means
of food production has been challenged due to their environmental impacts and the fact
that they may compete with humans for food, labour and alternative land uses for arable
farming (Mlambo and Mnisi 2019). The shortage of forage is worsened by the increas-
ing human and animal populations. As human and animal populations gradually increase,
the demand for food and feed also increases. For instance, the estimated population of the
SADC region increased from 327.5 million in 2016 to 337.1 million in 2017 representing
a 2.9% annual population growth rate (Food and Agriculture Organisation 2021). As the
human population increases, land formerly used for grazing is required for crop produc-
tion and developments (housing and infrastructure). Therefore, there is a need to optimise
livestock feed efficiency through growing and harvesting more forage and increasing its
utilization, thereby increasing animal productivity.
Besides increasing animal performance, the need to improve metabolism of ruminants
is driven by the fact that ruminants contribute to emissions of greenhouse gases (GHG)
into the atmosphere especially CO2 and CH4. These gases are produced during fermenta-
tion of feed in the rumen. Of the two gases, C H4 is the most potent GHG as it has a global
warming potential 25 times that of CO2 (Benchaar and Greathead 2011). However, CH4
13
Mitig Adapt Strateg Glob Change (2022) 27: 47 Page 3 of 26 47
is short-lived as its atmospheric life-time is only 12 years compared to 100 to 200 years
for CO2 (Lynch et al. 2020; Allen et al. 2018; Balcombe et al. 2018). From the perspec-
tive of national GHG emission targets, agriculture and livestock are an attractive target for
CH4 reduction campaigns as small changes in agricultural emissions could result in large
changes in total national GHG emissions (du Toit et al. 2013a). To the best of our knowl-
edge at a national level, South Africa is the only country in the SADC region with data
available and livestock contributed 27% to the national emissions (du Toit et al. (2013a),
in which beef and dairy cattle accounted for 73%, with beef in extensive system being the
highest emitter (83%) compared to dairy (14%) and feedlot (3.2%) in 2010. On the other
hand, small ruminants in South Africa contributed 16% of total national livestock enteric
CH4 emissions in 2010, with commercial sheep industry contributing an estimated 91%
of sheep emissions, whereas 56% of goat C H4 emissions originated from the emerging/
communal sector (du Toit et al. (2013b). It is possible that these figures are representa-
tive of the region since production systems are nearly similar. Therefore, the call for tar-
geted interventions cannot be ignored by SADC countries because in addition to the above
ruminant CH4 contributions, it is predicted that the region will experience drier and hotter
conditions as global temperatures increase (Maúre et al. 2018). Some of the worst impacts
on sustainable development are expected to be felt amongst agricultural and coastal sec-
tors. For example, it is predicted that global warming of 1.5 °C would lead to an average
temperature rise above the pre-industrial baseline in Botswana of 2.2 °C and of 2.0 °C in
Namibia (Maúre et al. 2018). These increases in temperatures are predicted to reduce for-
age yield and quality and may correspondingly increase CH4 production by 0.9% with each
1 °C temperature rise (Lee 2017). Therefore, there is a need to reduce the environmental
footprint of ruminant farming in the SADC region. In addition to its detrimental effects on
global warming, enteric CH4 emissions constitute 2 to 12% of gross energy intake (Mitsu-
mori and Sun 2008), and hence there is a need to reduce C H4 emissions to improve produc-
tion efficiency. Notwithstanding the Livestock’s Long Shadow (Steinfeld et al. 2006) nar-
rative in which ruminants are purported to produce 18% of global GHG, a re-calculation of
GHG emissions by Scholtz et al. (2012) showed that their actual contribution is 4%. The
overestimated contribution of ruminants to environment pollution makes them a scapegoat
for GHG emissions, whilst their strategic value in the supply of human nutrition and con-
tribution to biodiversity has often been ignored (Scholtz et al. 2020, 2012). In general, the
contribution of developing countries to GHG emissions is deemed high due to the legacy of
past emissions of long-lived C O2 and N2O which strengthens developing countries’ contri-
bution to global warming (Ward and Mahowald 2014). However, such C O2 and N
2O emis-
sions cannot be used to represent the magnitude of ruminal C H4 production. According to
Ward and Mahowald (2014), CH4 plays a greater role for non-Annex 1 countries, but as has
been pointed out by a number of researchers because of the short atmospheric lifetime of
CH4 relative to CO2, it is erroneous to weight distant past emissions equally with current
emissions by the CO2-eq metric (e.g. Blignaut et al. 2022; Leroy et al. 2022; Scholtz et al.
2020; Lynch et al. 2020). Indeed, it has been suggested that the current emissions weight-
ing scheme leads to unfair punitive polices on ruminant production systems, especially in
the SADC region that relies on agriculture and in particular animal agriculture for their
rural communities’ sustenance (USAID 2015). Agriculture has an important role to play
in an effort to reduce GHG emissions and for the maximisation of carbon sequestration
(Blignaut et al. 2022). This is despite that SADC’s emissions are below the world average
per capita emissions, with the exception of Botswana, Angola, Namibia, Zambia, South
Africa and the Seychelles who all have world average per capita emissions (USAID 2015).
This paper recognised the different pathways that ruminant production contributes to GHG
13
as outlined by Steinfeld et al. (2006) and Blignaut et al. (2022) but would only focus on
enteric CH4 emissions to avoid an expansive scope. Therefore, the objective of this review
H4 production con-
is to assess to what extent ruminant (cattle, sheep and goats) enteric C
H4 and its effects on the SADC region and possible mitiga-
tributes to global emissions of C
tion strategies.
2 Methodology
Journal articles were identified by using the following search engines; Scopus (https://
www.scopus.com/search/), ResearchGate (https://www.researchgate.net/), Google Scholar
(https://scholar.google.com/) and Academia (https://www.academia.edu/) specifying the
following keywords; agriculture, methane emissions, ruminants, rumen fermentation,
rangelands, tannins, phytochemicals, greenhouse gases emissions, Southern Africa Devel-
opment Community and their various combinations. Gray literature, dissertation or theses
were searched from Botswana University of Agriculture and Natural Resources library and
included, as well as reports from authors’ personal collections. Search was not restricted to
year of publication. Old, but classical references (e.g. Hegarty and Gerdes 1999) that lay
fundamental concepts to the review were included. However, for most references, where
newer articles with similar information or data was identified, the latter were selected.
3 The rumen
To better understand how ruminants contribute to global warming and are able to discuss
possible mitigation strategies, an understanding of rumen physiology and microbiology
is necessary. It is well documented that the rumen is a rich microbial ecosystem that is
colonized by billions of microbes including bacteria, archaea (methanogens), fungi, proto-
zoa, mycoplasma and bacteriophages. There are 1 010 to 1 011 bacteria, 1 08 to 1 09 archaea,
approximately 106 ciliated protozoa and approximately 1 06 fungi/mL of rumen fluid (Cie-
slak et al. 2013; Kumar et al. 2009). These microbes degrade cellulose, lipids and protein
to yield VFA, microbial cells and gases (Morgavi et al. 2010) as illustrated by Fig. 1 from
the study by Buddle et al. (2011).
The most important products of ruminal fermentation are VFA and microbial cells.
Volatile fatty acids are the primary source of metabolizable energy (ME); for instance,
propionate absorbed into the animal is used as the main precursor for glucose synthesis
through gluconeogenesis (Rugoho et al. 2019), and microbial cells which are digested
further along the digestive tract are the primary source of metabolizable amino acids for
maintenance, growth and milk synthesis (Fellner 2009). The VFAs are absorbed across the
rumen wall and act as a major source of carbon and energy for the ruminant animal. H2 is
used by methanogens to generate CH4 which is eructated by the animal and released into
the atmosphere (Buddle et al. 2011). Methane is the most potent GHG, and it has a heat of
combustion of 55.7 MJ/kg DM (McDonald et al. 2011), and its release to the environment
robs the ruminant animal of potential energy. A recent study by Blignaut et al. (2022) esti-
mated CH4 production as 6.3% of GEI, resulting in a mean relationship of ME equal to 0.9
digestible energy (DE).
13
Fig. 1 Diagrammatic representation of some rumen processes, highlighting the microbial fermentation
(thick arrows) of the ingested feed to volatile fatty acids (VFAs; mainly acetic, propionic and butyric acids)
and to hydrogen and carbon dioxide (H2 and CO2). Source: Buddle et al. (2011)
3.1 Methane production in the rumen and its importance
Some of the dominant methanogenes in the rumen include Methanobrevibacter ruminan-

tium, Methanosarcina barkiri, Methanobrevibacter formicium and Methanomicrobium
mobile (Kumar et al. 2009). Methane is produced as a result of fermentation of hydrolysed
dietary carbohydrates such as cellulose, hemi-cellulose, pectin and starch (Beauchemin
et al. 2020; Bhatta and Enishi 2007). Other gases produced inside the rumen are H2 (dur-
ing the conversion of hexose to acetate or butyrate) and carbon dioxide (Bhatta and Enishi
2007). Ninety-five percent of these gases are released through eructation during rumina-
tion (Lee et al. 2017). Methane synthesis is a necessary step for removing H 2, thus pro-
pelling the reduction of protons to H 2. Conforming to historical paradigms (Hegarty and
Gerdes 1999), a study by Greening et al. (2019) found that hydrogenotrophic methanogen-
esis appears to be the largest sink of H2. However, some H2 sinks are not net H2 sinks in
the sense that their production involves greater H2 production than incorporation (Unger-
feld 2020). For instance, recent studies (e.g. Roque et al. 2021) have shown elevation of
H2 from inclusion of anti-methanogenic feed additives such as red seaweed (Asparagop-
sis taxiformis) in total mixed ration (TMR) diets, which resulted in decrease in CH4, and
feed intake but improved carcass quality. This work by Roque et al. (2021) is consistent
with a metatranscriptomic study by Greening et al. (2019) which reported that whereas
methanogenesis-related transcripts predominated in sheep with high C H4 yield, alternative
uptake pathways were significantly upregulated in low C H4 yield sheep. Methanosarcina
mazei synthesises CH4 from acetate, methanol and methylamines, whilst M. barkeri uti-
lises H2/CO2, acetate, methanol and methylamines for C H4 synthesis (Jarvis et al. 2000).
2 for use in the synthesis of C
Thus, in order to reduce the availability of H H4, H2 should be
converted into propionate via lactate or fumarate (Asanuma et al. 1999) as shown in Fig. 2
(Morgavi et al. 2010). The recent work by Roque et al. (2021) thus proposes an alternative
theory in dealing with high H2 concentration when anti-methanogenic feed additives such
as seaweeds are added; the redirection of H 2 molecules that would otherwise be utilized to
13
Fig. 2 Schematic microbial fermentation of feed polysaccharides and H2 reduction pathways in the rumen.
Source: Morgavi et al. (2010)
form CH4 and into different pathways that could be beneficial to the animal. This theory
is supported by Greening et al. (2019) to the effect that whereas the enzymes mediating
fermentative H2 production are expressed at similar levels, those supporting H2 uptake
through acetogenesis, fumarate reduction and nitrate ammonification pathways are highly
upregulated in sheep with low C H4 yield. Wang et al. (2018) also noted that when com-
pared with control (TMR with high forage NDF), a non-forage fibre diet increased the pro-
duction of H2 by 8.5%, but the production of C H4 decreased by 14%. This was attributed to
shifting of H2 flow toward propionate, and part of it to H2 (Wang et al. 2018).
Thus, practical methods that reduce CH4 emissions should come from multiple path-
ways. Some of these strategies include feeding legumes, incorporating forages with sec-
ondary compounds, providing supplements which improve animal nutritional status and
efficiency of feed energy and improved genomic selection (Blignaut et al. 2022). These
and other strategies will be discussed later. Formate is also used to produce CH4, and it
accounts for approximately 15–20% of the total CH4 production in the rumen (Asanuma
et al. 1999). Studies by Bhatta and Enishi (2007) and Kumar et al. (2009) have shown
that M. ruminantium, M. formicicum and M. mobile utilise H2, CO2 and formate to pro-
duce CH4. These methanogenic archaea are attached to the outer surface of ciliate protozoa
(Vogels et al. 1980), and thus the removal of protozoa has been reported to decrease C H4
production (Ushida et al. 1997).
Carbon dioxide is more concentrated than C H4 in the atmosphere; however, C H4 is
more harmful to the environment than C O2, because CH4 is a more potent GHG than car-
bon dioxide and constitute energy loss for ruminants (Eckard et al 2010). Despite this, the
reduction of CH4 emissions in order to decrease global warming appears to be the most
effective in the short term than C O2 because C H4 has been estimated to have a chemical
lifespan of 12 years unlike C O2 that has a lifespan of 50–200 years (Moss et al. 2000).
Trends in C H4 emissions and its oxidation can produce long-term trends in stratospheric
water vapour. The required temperature for halogen activation allowing heterogene-
ous ozone-depleting reactions on polar stratospheric cloud (PSC) particles is a function
of water vapour concentration (Rosenlof 2018) that facilitate CH4 emissions. This allows
13
more heat to reach the earth resulting in increased global temperature. The rise in tem-
peratures alters precipitation patterns, causing extreme weather conditions such as floods
and high temperatures (Moss et al. 2000) with severe impact on agricultural land, which
ultimately affects food security (Masipa 2017). The SADC region is already experiencing
fluctuating weather patterns, with delayed rainfall but heavy floods when it rains. In addi-
H4 emissions from ruminants con-
tion to its contribution to greenhouse gasses, enteric C
stitute a loss of 2–15% of ingested energy (Eckard et al. 2010; Johnson and Johnson 1995;
Moss et al. 2000). This reduces feed efficiency and utilization by the animal. Thus, there is
a need to find ways in which C H4 production can be reduced. Several studies (e.g. Eckard
H4 production.
et al. 2010 and Moate et al. 2018) have indicated that feed type influences C
Therefore, a feed conceptual framework has been proposed (see Fig. 3) based on several
studies reviewed. The agents, when introduced, result in the blockage or minimisation of
the red pathways (symbolised by X) but promote the green pathways (Fig. 3), thus reduc-
ing CH4 emissions and increased ruminant productivity respectively.
4 Strategies to reduce methane emissions in the SADC region
Within SADC, countries with the highest agriculture emissions are Angola, South Africa,
Madagascar, Zambia, Mozambique, Botswana and Zimbabwe and their agriculture emis-
sions account for 90% of the region’s GHGs from agriculture (USAID 2015). This has led
Sub-Saharan African (SSA) governments to commit through the Gaborone Declaration for
Sustainability in Africa to uphold the United Nations’ Sustainable Development Goals and
the Paris Accord (Gaborone Declaration 2017). Of the seven SADC countries with highest
Fig. 3 Conceptual framework on feed manipulation to reduce methane yield in the rumen and improve ani-
mal productivity. Green pathway minimises methane production and promotes animal productivity whilst
the red pathway yield methane and needs to be blocked
13
agriculture emissions, enteric fermentation is amongst the top three emitting agriculture
subsectors; hence, a range of needs has been identified, including improved livestock, pro-
moting feedlots and using improved feed; promoting livestock breeding and fighting pas-
tureland fires; reduction in emissions from livestock and manure management and biogas
captures (USAID 2015) as well as pledges to reduce emissions by 19% by 2030. Therefore,
this review is restricted to identifying strategies that are consistent with the above policy
commitments by improving the efficiency of ruminant production and hence reduce CH4
emissions and C H4 intensities.
4.1 Feeding management and methane production
Feed type, quality and DM intake have been found to affect C H4 production, e.g. high
concentrate diets, leguminous forages, diets containing tannin and diets with high lipid
concentration have been found to decrease CH4 production (Clark et al. 2010). Feeding
ruminants more concentrates, e.g. grains, will result in more production of propionate and
reduced rumen pH which inhibits the growth of protozoa and methanogens in the rumen
resulting in reduced C H4 production. Grain type is also associated with reduction in C H4
production as demonstrated by a recent study by Moate et al. (2018) who noted that lactat-
ing dairy cows fed wheat (Triticum) produced substantially lower C H4 yield and intensity
than when fed rolled corn (Zea mays) grain.
However, for the SADC region, feeding ruminants with grains is not a suitable or a sus-
tainable strategy. When reared as nature intended, ruminants should offer very little compe-
tition to humans for food because they prefer to consume grass and other fibre-rich forages
(Mlambo and Mnisi 2019). Ruminants in developing countries are predominantly main-
tained on a high-roughage diet with little or no concentrate resulting in increased ruminal
methanogenesis (Goel and Makkar 2012) and reduced productivity. Therefore, a challenge
for SADC countries will be to make forage more efficient in three ways: (1) increasing fibre
digestibility and (2) reducing detrimental effects of tannins on digestibility and (3) taking
advantage of tannins to reduce CH4 production in the rumen. On the other hand, more
concentrate, i.e. low fibre intake or excessive intake of rapidly fermentable carbohydrates,
causes sub-acute rumen acidosis (Chiba 2009; Rugoho et al. 2019) and should be avoided.
Consequently, concentrates should form less than 60% of the animal diet and/or should be
even less in the SADC countries where grain for ruminant reaches a prohibitive price. We
propose that when grains are weather damaged, or of such poor quality that they are unsuit-
able for milling and therefore unsuitable for human consumption, then under these circum-
stances limited amounts of grains may be included in the diets of ruminants in the SADC
region. Earlier research (Lichtenwalner et al. 1979) documented higher ash and in situ
digestibility of weathered sorghum (Sorghum bicolor) grain than nonweathered grains;
however, intake was depressed. PennState Extension (2016) cautioned about presence
of aflatoxins and mycotoxin in wet grains and provided classification of risks at various
mould spore counts in which grains with 3 to 5 million spore count per gram could still be
used provided they are diluted with other feeds. An increase in sorghum grain production
by farmers, which is agronomically suitable for the SADC region, would fulfil this purpose
(Madibela and Lekgari 2005), and inclusion of second grade sorghum grains in ruminant
diets would supply tannins that could reduce C H4 yield. This was demonstrated by Mavasa
et al. (2022) who observed reduction in C H4 yield without reduction in performance of
Pedi goats in South Africa when sorghum was used to partially replace maize grain. A
study by Beechem (2010) found that the protozoal populations fell by 83% and 76% in
13
cattle fed high and low concentrate diets respectively when treated with ionophores. This
indicates that a small amount of concentrate may still be effective, especially to make the
rumen more efficient. Hence, a combination of concentrates and ionophores can be used as
H4 production. However, the use of ionophores and other growth hor-
a strategy to reduce C
mones is prohibited in some SADC countries. For example, Botswana (Van Engelen et al.
2013) and Namibia were the first African nations to ban the routine use of antibiotics in the
beef industry 26 years ago (World Health Organisation 2017). There is a global concern
regarding food safety and environmental burden due to use of chemicals (Kobayashi 2010).
Therefore, phytochemicals as alternatives to antibiotics to promote growth (Lillehoj et al.
2018) and novel carbohydrate sources to substitute grains should be explored.
Forage quantity, quality and intake are associated with CH4 production, i.e. the more
fibre the forage contains, the more CH4 is produced in the rumen (Drehmel 2017). There-
fore, CH4 production can be reduced by decreasing fibre content of forages. Alternatively
non-forage fibre sources (wheat bran and soyabean (Glycine max) hulls) could be used
to reduce forage’s contribution of NDF (Wang et al. 2018) which was found to reduced
CH4 yield in vitro. Non-forage fibre sources which are derived from crop processing and
hence by-products like cereal bran, hulls and husks from post harvesting at farm levels
in the SADC region would fit this concept. Furthermore, steers grazing an early season
pasture produced approximately 45% less C H4 than those fed on mid or late season pas-
ture (Boadi and Wittenberg 2002). Hence, forages should be fed or harvested at an early
bloom when they have a low fibre concentration. Therefore, mechanisation in fodder pro-
duction in SADC is important to cut and conserve forages whilst still in a high-quality con-
dition. Manufacture of suitable machinery for small-scale livestock farmers is imperative
as demonstrated by Rural Industry Innovation Centre in the 1980s and 1990s in Botswana
(Chanda 2000) who manufactured small-scale forage harvesters. Farmers can use appropri-
ate machinery to top up the grass before it lignifies to keep it in a vegetative state thereby
improving quality. The starting point would be to encourage livestock farmers to set land
aside for fodder production and the use of organic fertiliser (manure) from livestock. Also,
in areas with high volumes of water, irrigation might be used to maintain animal forages in
an actively growing vegetative (high-quality) state. However, the scarcity of water for irri-
gation and aridity of much of the SADC region does not make irrigation a viable option for
many smallholder farmers. Alternatively, green pasture belts can be developed along rivers
and water bodies in the drier parts of the SADC region.
Methane production was 9% higher in cows fed on pasture than on alfalfa (Medicago
sativa) due to pasture having more fibre in the study by McCaughey et al. (1999). Fur-
thermore, Kurihara et al. (1999) reported that C H4 energy loss in cattle fed tropical forage
diets was higher than in those fed on temperate forage diets due to higher levels of fibre and
lignin in the former than the later. For instance, ruminants fed C4 grass species produced
17% more C H4 as L/kg OM intake compared to those fed C3 grass species (Archimède
et al. 2011). A recent modelling study (Lee et al. 2017) predicted that increasing global
temperatures will reduce forage quality and correspondingly increase CH4 production by
0.9% with each 1 °C temperature rise. Considering the already low digestibility of rumi-
nant feed resources from rangelands in the SADC region, this will constitute a serious chal-
lenge. Another modelling study by Herrero et al. (2013) expressed non-CO2 eq emissions
per consumed protein from livestock, and their comparisons found SSA as a hotspot for
GHG emissions generally because of low animal productivity due to the use of low-quality
feeds and feed scarcity. As a result, there is a need for strategies for modifying fibre in for-
ages to reduce CH4 production. Strategies include the use of alternative grazing species,
e.g. adding the foliage of the browse tree legumes like leucaena (Leucaena leucocephala)
13
has been shown to improve the quality of low-quality tropical grasses (Santana et al., 2019)
and reduce enteric CH4 emissions (Davison et al. 2020; Piñeiro-Vázquez et al. 2018; Soltan
et al. 2017). For example, the inclusion of 80% of leucaena in the diet of heifers fed low-
quality tropical forages reduced enteric C H4 emission by up to 61.3% without affecting
DMI, OMI and ruminal protozoal populations in an in vivo study (Piñeiro-Vázquez et al.
2018). The effects of leucaena on reducing enteric CH4 emission in vitro studies are shown
in Table 1.
Other strategies, like reducing the fibre content by adding fibre degrading enzymes dur-
ing silage production (Khota et al. 2018; Nkosi et al. 2011), would be worth exploring.
Under smallholder farming conditions, enzymes may not be accessible, and hence locally
available feed additives such as malt may be appropriate (Matshaba 2014). Malted sor-
ghum, for instance, used in small quantities, can be used as an inoculant in silage making.
The malting process is incorporated in indigenous knowledge of making traditional beer
in the SADC region. Other options entail the use of white rot fungi (Pleurotus spp) to dis-
rupt and solubilise lignin of lignocellulosic biomass (Akinfemi et al. 2009; Ntokome et al.
2018) and hence improve digestibility and metabolic efficiency in the rumen. Including
millet (Pennisetum glaucum), sorghum and maize (Zea mays) stover treated with white rot
fungi increased CP, in vitro organic matter digestibility (OMD), soluble fraction (a) and
effective degradability but reduced ADF and ADL (Ntokome et al. 2018). Type of forage
fed to ruminants could play a major role in reducing C H4 yield with feed management in
SADC. For example, legumes from warm climates produce less CH4 than legumes from
cold climates, and that legumes produce less CH4 than grasses (Archimède et al. 2011).
Therefore, tropical legumes with secondary metabolites would be a suitable option within
smallholder farms. In addition, tropical legumes with their high CP content can be used in
intercropping systems with grasses to improve the overall sward quality and reduce its CH4
yield potential. Moses et al. (2018) in Botswana observed that when Macroptilium atropur-
pureum (siratro) was used in a pasture mixture including buffel grass (Cenchrus ciliaris),
not only was the CP content improved, but the degradation lag time in vitro was short-
ened, and the effective degradability and in vitro dry matter digestibility (IVDMD) were
increased. Further research to quantify C H4 yields from mixed tropical legumes and C4
grass species is needed. The extension message from the above argument would be to pro-
vide nutritional supplements that improve the nutritional status of the animal, and the effi-
ciency of feed energy use in order to reduce C H4 emissions from grazing cattle (Thompson
and Rowntree 2022).
4.2 Use of monensin as a feed additive
In the past, major interest for producers, especially in feedlot systems, was the use of
monensin for the re-partitioning energy away from C H4 production to growth for efficient
energy utilization. However, in recent times, interest in ionophores use has focused on its
ability to reduce enteric CH4 production. The addition of monensin to ruminant diets has
been associated with a reduction of C H4 production (McGinn et al. 2004; Tedeschi et al.
2003), albeit to a lesser extent than that which occurs in response to feeding sunflower
(Helianthus) oil (McGinn et al. 2004). In a study by Odongo et al. (2007), CH4 production
was reduced by 7 to 9% when monensin was added to the diet of lactating dairy cows with
no effect on milk quantity. However, the long-term use of monensin might lead to rumen
microbes developing resistance to antibiotics (Sahoo et al. 2010). Hook et al. (2009) con-
ducted a study to find out if monensin has a significant effect on diversity of methanogens
13
Table 1 Effects of tannins: source, type, state and inclusion rate on methane reduction both in vitro and in vivo
Plant Plant/crude extract/ Measurement system Feed for animals Inclusion rate CH4 reduction (%) Reference
purified
a
Acacia mearnsii Tannin extract In vivo in sheep Eragrostis + Lucerne 40 g/kg feed 30% of CH4 g/kgDM Adejoro et al. (2019)
hay:concentrate intake
(50:50)
Silvafeed bypro Commercial tannin In vivo in sheep Eragrostis + Lucerne 10 g/kg feed 12% of CH4 g/kgDM Adejoro et al. (2019)
additive hay:concentrate intake
(50:50)
b
Acacia nilotica Plant leaves In vitro gas production Natural pasture based 25,50, 75,100% + 8.2, 14.8, 31.4, 64% Rira et al. (2019)
on Dichanthium spp (400 mg feed
sample)
c
Acacia nilotica Plant pods In vitro gas production Natural pasture based 25, 50, 75, 100% + 4.1, 27, 37.7, 55.1% Rira et al. (2019)
on Dichanthium spp (400 mg feed
sample)
d
Moringa oleifera Leaf meal In vitro gas production Replacing soyabean 10, 20, 30, 40, 50, (% reduction of mL Elghandour et al.
using goat rumen meal in TMR 60, 70, 80, 90, 100 CH4/g incubated (2017)
liquor at 48 h of (g/100 g DM) DM) 31.4, 32.4,
incubation 49.7, 19.7, 16.9,
19.0, 21.3, 19.7,
16.9, 15.9%
d
Moringa oleifera Leaf meal In vitro gas production Replacing soyabean 10, 20, 30, 40, 50, (% reduction of mL Elghandour et al.
using steer rumen meal in TMR in 60, 70, 80, 90, 100 CH4/g incubated (2017)
liquor at 48 h of Steer (g/100 g DM) DM) 20.0, 21.5,
incubation 21.5, 17.4, 24.0,
20.0, 10.7, 9.6, 16.0,
11.0%
e
Leucaena leuco- Leaf meal In vitro gas production Incubating Leucaena 30.7% of mL/g DM Soltan et al. (Soltan
cephala using rumen liquor (1000 mg) with low et al. 2017)
from Santa Ines mimosine (2.3 mg/kg
sheep DM) after addition of
PEG (MW6000)
Page 11 of 26 47
13
Table 1 (continued)
Plant Plant/crude extract/ Measurement system Feed for animals Inclusion rate CH4 reduction (%) Reference
purified
13
e
Leucaena leuco- Leaf meal In vitro gas production Incubating Leucaena 19.4% of mL/g DM Soltan et al. (2017)
47 Page 12 of 26
cephala using rumen liquor (1000 mg) with low

from Santa Ines mimosine (11.1 mg/
sheep kg DM) after
addition of PEG
(MW6000)
e
Biophytum peter- Leaf meal In vitro gas produc- Basal grass of Pennise- 20% 25% Hariadi and Santoso
sianum tion using rumen tum purpurem (80%) 20% 29% (2010)
f
Acacia mangium fluid from Ongole supplemented with
crossbred cattle B. petersianum or A
mangium
a
Acacia mearnsii contained 0.651 g/g DM total phenol, 0.58 g/g DM total tannin (as tannic acid equivalent) and 0.35 g/g DM of CT (as leucocynidin equivalent)
b
Acacia nilotica leaves contained 80 g/kg DM CT, 31 g/kg DM gallotannins and 147 g/kg DM ellagitannins
c
Acacia nilotica pods contained 157 g/kg DM CT, 84 g/kg DM gallotannins and 266 g/kg DM ellagitannins
d
Cannulated Creole Goats and Holstein steers were fed ad lib with a diet consisting of oat hay and concentrate at 60:40 ratio
e
Cannulated Santa Ines sheep (n = 6) grazed Brachiaria decumbens, Pennisetum purpureum and Leucaena leucocephala and supplemented with graze maize and soyabean
(0.7 kg/100 g LW) and free access to mineral premix

and found that it does not have a direct effect on the methanogen population. It has been
reported that ruminal ciliate protozoa are reduced by initial ionophore exposure, but that
long-term exposure to monensin led to recovery of the ruminal population (Guan et al.
2006). This is also an issue for microbes developing resistance to antibiotics. Moreover, a
study by Rogers et al. (1997) showed that the extent of reductions in total ruminal protozoa
numbers as a result of monensin supplementation decreased with time. Many methanogens
are found in association with ciliate protozoa, and there is evidence they live on the outer
surface of the protozoa (Krumholz et al. 1983; Newbold et al. 2015).
Although monensin is effective in reducing C H4 production, it is believed that the con-
tinuous use of antibiotics will end up with their residues reaching the human food chain
(Odongo et al. 2007). This should be avoided because antibiotic residues may induce path-
ogenic microbes that inhabit the human gastrointestinal tract to develop resistance to anti-
biotics. In countries such as Botswana and Namibia, the ban on chemicals like monensin
(Van Engelen et al. 2013; World Health Organisation 2017) means that other alternatives
should be explored (Lillehoj et al. 2018). For example, a possible alternative is tannins as
they have been shown to reduce enteric CH4 production (Greathead 2003; Rira et al. 2015,
2019). Feeding forages containing plant secondary metabolites (i.e. saponins, tannins and
essential oils) to ruminant animals could be used as a strategy to reduce CH4 production
by ruminant animals. In addition, in contrast to ruminant agriculture in developed west-
ern countries where cattle are often fed TMR, ruminant production in the SADC region
is based on grazed pasture where monensin has been found to be ineffective at reducing
methanogenesis. Therefore, besides the ban on production enhancing chemical, the use of
monensin in ruminants grazing natural pasture would be irrelevant in the SADC region due
to very little or no effect on CH4 emissions (Grainger et al. 2010).
4.3 Use of tannins as feed additives
Tannins are plant secondary compounds that are not involved in the primary biochemical
process of plants, but act as plant protection mechanisms (Silanikove et al. 2001). They
have antimicrobial activity (Cowan 1999), they protect plants against insect attack (Ørskov
1998), and as tannins are astringent and unpalatable, they dissuade animals from grazing
tanniniferous plants (Barbehenn and Constabel 2011). Tannins are divided into two groups,
namely condensed tannins (CT) and hydrolysable tannins (HT) (Lee 2017). Condensed
tannins are found in gymnosperms and angiosperms whilst HT are found only in dicoty-
ledons (Silanikove et al. 2001). Local forages and fodder trees in the SADC region are
endowed with polyphenolic compounds including both condensed and hydrolysable tan-
nins (Madibela et al. 2018; Phale and Madibela 2006). However, there is limited evidence
(Theart et al. 2015) on the use of African forages containing secondary metabolites to
reduce CH4 yield apart from few studies done outside the SADC region (see Table 1). The
effects of African forages containing secondary metabolites on CH4 yield need more study
in the SADC region.
Nevertheless, based on an extensive findings in the scientific literature, tanninifer-
ous plants should be incorporated in ruminant diets in the SADC region as they are well
adapted to the semi-arid conditions and tend to produce nutritious browse products well
into the dry season (Mlambo and Mapiye 2015). However, diets with high CT concen-
trations (> 55 g CT/kg DM) generally reduce voluntary feed intake and digestibility and
are associated with low rates of body and wool growth in grazing ruminants (Min et al.
2003). On the other hand, Tiemann et al. (2008) reported that CT reduces CH4 production
13
by reducing fibre digestibility, perhaps because C H4 production is associated with highly

fibrous feed material. However, the study by Carulla et al. (2005) found that black wattle
(Acacia mearnsii) containing 0.6% CT reduced CH4 production without decreasing fibre
digestibility. The challenge, therefore, is to identify cost-effective components (of plants
and their type) (Bodas et al. 2012) that favourably modify ruminal fermentation to reduce
CH4 production without reducing production by decreasing digestibility. For instance, in an
in vitro study conducted with fodder material from the Thorny Kalahari Dune Bush veld of
South Africa, Theart et al. (2015) found that samples of kalahari-sand acacia (Acacia lue-
deritzii) and blue bush (Monechma incanum) with total tannin content of 299 and 283 g/kg,
respectively, substantially reduced CH4 yield without adversely affecting organic matter
digestibility (OMD). The effects of tannins in reducing CH4 production depend on its con-
centration in plants, composition of the diet, animal physiological state and animal species
(Makkar 2003) and whether it occurs as HT or CT (Aboagye and Beauchemin 2019; Rira
et al. 2019). Condensed tannins have other benefits to animals such as promoting increased
liveweight gain and decreased severity of infestations by gastrointestinal parasites (nema-
todes) (Mata-Padrino et al. 2019; Min et al. 2003). Because of this inverse relationship
between CT concentrations, type and structure with voluntary feed intake and fibre digest-
ibility in ruminants, forages that contain high levels of tannins need to be mixed with those
that do not contain any tannins or contain less CT to reduce their adverse effect on feed
intake and digestibility. For example, farmers in the SADC region have adopted feeding
tanniniferous plants as part of strategy for suppling essential limited nutrients and improv-
ing liveweight gains as demonstrated by Brown et al. (2018) in South Africa. This strategy
involves harvesting tree foliage and collection of pods to offer as supplementary ration,
especially during the dry season (Smith et al. 2005) reducing mortality of twin kids and
improving goat performance in Zimbabwe. Other than tannins, fatty acids (FA) are also
believed to reduce CH4 production when incorporated in animal diets (Beauchemin et al.
2008, 2009; Soltan et al. 2018).
4.4 Use of fatty acids as feed additive
Polyunsaturated FA (PUFA), e.g. linoleic and linolenic acids, and medium chain FA
(MCFAs), e.g. lauric and myristic acids, have been reported to reduce CH4 production
when included in animal diets (Eugène et al. 2008). This is mainly due to their toxic effect
on bacteria, protozoa and methanogens (McAllister et al. 1996). Sunflower seed, linseed
(Linum usitatissimum), coconut (Cocos nucifera) and rape (Brassica napus) seed are the
main source of PUFA and MCFAs (Rasmussen and Harrison 2011). Beauchemin et al.
(2009) using lactating dairy cows noted a 10% decrease in CH4 production in cows fed on
diets containing 3.3% DM of sunflower seed. In the study by (Beauchemin et al. 2009), the
addition of sunflower seeds to the diet of dairy cows resulted in reduced CH4 production
with no change in rumen pH and milk production. Just like sunflower oil, linseed oil also
contains a high concentration of PUFA (18:3) (Rasmussen and Harrison 2011), and addi-
tion of linseed oil (3.3% DM) to the diets of dairy cows reduced CH4 production by 18%
with no effect on rumen pH (Beauchemin et al. 2009).
Canola (Brassica napus) seeds contain high concentrations of medium unsaturated FA
(18:1), and when it was included in the diets of dairy cows (3.3% DM) it reduced CH4 pro-
duction by 16% (Beauchemin et al. 2009). The variability in CH4 production resulting from
dietary addition of these oil sources indicates that the types of FA and their concentrations
determine the amount of C H4 reduced. However, fat should not exceed 6% of the diet of
13
ruminants because too much of it will cause digestion disorders (Machmüller et al. 2000).
The sources of FA outlined above are not produced in large quantities in SADC, except for
sunflower. Recently canola has been introduced in South Africa and its adaption and adop-
tion will contribute to a variety of ingredients that can be used in ruminant’s diets. This
calls for investigations into other sources of lipids and essential oils that are locally avail-
able, especially from indigenous plants, as ruminal fermentation modifiers. For instance,
Ramathudi (2016) estimated in vitro CH4 reduction potential of 51.2, 43.4 and 40.8% by
coating maize stover with lipids from Ximenia caffra, Racinus communis and Citrullus vul-
garis respectively, at 5% inclusion. Studies outside the SADC region have also reported
that dietary inclusion of essential oils reduces CH4 emissions (see Table 2).
Dietary inclusion of essential oils is believed to reduce C H4 emissions by shifting the
microbial fermentation in the rumen to decrease the acetate to propionate ratio (Castillejos
et al. 2007; Ebeid et al. 2020). However, some oils such as Moringa (Moringa oleifera) seed
oil, in addition to their broad-spectrum antimicrobial activity, are rich in sterols and anti-
oxidants such as polyphenolic compounds, α-tocopherol (134 mg/kg) and vitamin E which
can potentially affect microbial diversity and fermentation process in the rumen (Ebeid
et al. 2020). Even though positive results on CH4 reduction by essential oils have been
observed in vitro, this was accompanied by negative effects on DMD by some essential oils
unless included as a combination of oils to provide a synergistic associative positive effect
(Cobellis et al. 2016) (Table 2). Therefore, the challenge remains to identify essential oils
that selectively inhibit rumen methanogenesis at practical feeding rates, with lasting effects
and without depressing feed digestion and animal productivity (Benchaar and Greathead
2011). One strategy is the use of essential oil or oil from plants from small-scale cottage
industries which process seed for oil either for food, biofuel or cosmetics. Essential oil and/
or cake from such processing would be made available as feed additives or feed cake to
supplement ruminants supplying the needed FA to reduce CH4 yields. For example, cottage
industries in Botswana and Namibia in which morula (Sclerocarya birrea) fruits are har-
vested by women groups and processed for cosmetic oil have resulted in a seed cake used
as a source of energy and protein which Baleseng (2022) found to improve growth rates
of local sheep. Due to exports sanitary compliance to European Union markets, feeding
material considered as natural products such as seaweeds to ruminants will be attractive
to SADC farmers. Seaweed (known as marine algae) has a tradition of being part of the
animal feed in the coastal areas, from ancient times (Morais et al. 2020). Feeding livestock
seaweed has been shown to reduce CH4 production (Vijn et al. 2020) and will be suit-
able for circumventing effects of feed shortage in the SADC region. Angola, Mozambique,
Namibia and South Africa have access to a long coast from the Atlantic to India oceans,
and hence there exists an opportunity for harvesting and/or farming of seaweed.
4.5 Potential use of red seaweed and other aquatic plants in reducing methane
yield
Recent research has shown the potential of seaweed in reducing CH4 yield in ruminants.
Studies by Machado et al. (2014) and Maia et al. (2016) noted high reduction of CH4
in vitro, with inhibition of more than 90%. In support, feeding studies (Roque et al. 2019,
2021) with dairy cows and beef steers respectively fed TMR-based diets observed CH4
reduction of 26.4 vs 67.2% and 69 vs 80% and as a result of inclusion of seaweed (Aspara-
gopsis spp) at rate of 0.5% vs 1.0% and 0.25 vs 0.5% respectively. Other aquatic plants with
feeding possibilities are Azolla (i.e. Azolla filiculoides), Salvinia (i.e. Salvinia molesta) and
13
Table 2 Effects of essential oil and inclusion rate on methane reduction both in vitro and in vivo
Plant Plant/crude extract/ Measurement system Feed for animals Inclusion rate CH4 reduction (%) References
purified
13
a
Pequi Essential oil In vitro gas produc- TMR of corn silage, 5% v/v of inoculation 86.7% CH4 Freitas et al. (2018)
47 Page 16 of 26
tion with rumen Tifton 85 concen- media (24 h mL/g DM

fluid from Holstein trate and minerals incubated)
crossbred
b
Pequi Essential oil In vivo in Dorper TMR of corn silage, 75 mL oil/animal 17.7% of CH4 g/kgDM Freitas et al. (2018)
sheep in respiration Tifton 85 concen- intake
chamber trate and minerals
c
Agolin Ruminant A blend of essential Four multiparous Fed mixture of grass 200 mg essential oils/ 15% (g/d) and 14% (g/ Castro-Montoya et al.
oils (coriander oil, lactating Holstein silage, maize silage animal kgDMI) (2015)
geranyl acetate and cows in open circuit and soybean meal
eugenol) chambers and supplemented
with concentrate
d
Agolin Ruminant A blend of essential Four Belgian Blue Fed maize silage 200 mg essential oils/ Ranged 13% and 20% Castro-Montoya et al.
oils (coriander oil, double muscled ad libitum and animal (g CH4/kg BW) (2015)
geranyl acetate and beef heifers in open supplemented with
eugenol) circuit chambers concentrate
e
Moringa oleifera Seed oil In vitro using rumen TMR of Roughage 4% of sample incu- 48.4% of CH4 L/g DM Ebeid et al. (2020)
fluid from three 70:Concentrate 30 bated
crossbred (Mur- was incubated
rah × Chinese local)
water buffaloes.
Methane measured
using GC system
f
Cinnamon leaves Essential oil In vitro using fluid Alfalfa hay and a 1.125 mL/L culture 69.2% of mL C
H4 Cobellis et al. (2016)
from lactating Jersey concentrate mixture (24-h incubation)
cows (ground corn, soy
Methane measured hulls, soybean meal
with GC and a mineral/vita-
min mixture)

Table 2 (continued)
Plant Plant/crude extract/ Measurement system Feed for animals Inclusion rate CH4 reduction (%) References
purified
g
Combination of Essential oils In vitro for using fluid Alfalfa hay and a 0.8 mL/L total 78.5% of mL CH4 Cobellis et al. (2016)
cinnamon leaves, from lactating Jersey concentrate mixture (48-h incubation)
oregano leaves, cows (ground corn, soy
rosemary leaves Methane measured hulls, soybean meal,
with GC and a mineral/vita-
min mixture)
a,b
Pequi contained on g/100 g basis; 0.08 lauric acid, 0.12 myristic acid, 27.1 palmitic acid, 2.57 stearic acid, 44.7 oleic acid, 21.1 linoleic acid, 1.95 linolenic acid and 0.21
arachidic acid. The incubated material was Tifton 85 hay with 3 increasing levels (1, 2 and 5% (v/v)) of oil
c,d
Each animal received a daily dose of 200 mg essential oils, which was homogenously mixed with part of the concentrate
e
The study does not indicate what type of material was fed to the cannulated buffaloes
f
Jersey cows fed on corn silage, mixture of alfalfa hay and grass hay and concentrate mixture
g
Jersey cows fed on corn silage, mixture of alfalfa hay and grass hay and concentrate mixture
Page 17 of 26 47
13
Lemna (i.e. Lemna minor). However, limited literature (Chakrabarti et al. 2018; Goopy
and Murray 2003; Halmemies-Beauchet-Filleau et al. 2018; Leterme et al. 2009; Sońta
et al. 2019; Zakaria and Shammout 2018) report only the production capacity and feeding
value indicating endowment with high protein and amino acids concentration, high trace
minerals and high DMD. However, experimental studies have mainly focused on poultry
and pigs (Leterme et al. 2009; Mwale and Gwaze 2013; Zakaria and Shammout 2018).
An early study by Huque et al. (1996) recorded high mean CP (32.7%) of three duckweed
(Lemnoideae) species with DM and CP digestibility of 57 and 68.2% respectively. To our
knowledge, there is no study which has reported their CH4 reducing potential and hence a
worthy research area. The studies mentioned earlier, on seaweed, both in vitro and in vivo,
demonstrate that seaweeds could play an important role in reducing CH4 in ruminant pro-
duction, an innovation which the SADC region can adopt. For instance, South Africa has a
costal length of 2850 km, Namibia has 1500 km, Angola has 1600 km, whilst Mozambique
has 2600 km and therefore could be a source of seaweeds and/or foundation for commer-
cial farming of seaweed. Notably, seaweed is currently being used to produce commercial
chicken feed products and feed supplements for livestock farms in South Africa (Matshogo
et al. 2021). However, it is important to note that innovative strategies which use seaweed
to reduce enteric C H4 have been developed for feeding to ruminants on TMR based diets
which might be impossible in grazing-based ruminant systems in the SADC region. How-
ever, used as supplements, these can still find a niche in the grazing-based systems where
animals are conditioned to return to a watering point every day or alternate day for water-
ing and mineral licks. As a matter of practice, grazing ruminants in the SADC region,
especially for 70 to 90% of cattle owners who are small-scale resource limited farmers
(Statistics Botswana 2019), are brought to watering points, which consist of pens and han-
dling facilities to be watered or kraaled overnight. With regard to other aquatic plants such
as duckweed and Salvinia which customarily invade water bodies and wetlands in SADC
like the Kariba Dam in Zimbabwe and Okavango Delta in Botswana, physical harvesting
has been found to have limited but effective control (Kurugundla et al. 2016). Therefore,
management with utilization for livestock feed can be mounted in locations where livestock
are reared proximal to these water bodies. Alternatively, local on-farm production in ponds
incorporating nutrient scavenging from field runoff, manure and greywater may offer a via-
ble possibility for ruminant feed production and thus reinforce circular economy practices
at farm level and decrease the environmental footprint of ruminant-based food production
systems (Halmemies-Beauchet-Filleau et al. 2018). The strategies outlined above would
serve as short- and medium-term approaches, and in the long term, selecting animals with
H4 would be sustainable.
inherent ability to produce less C
4.6 Genetic selection of ruminants for low methane emissions
Although the magnitude of emissions of C H4 expected to be reduced by selecting for low

CH4 ruminant animals is small about 7% (Davison et al. 2020), mitigation of enteric CH4
emission by genetic selection has become an important area of research in many regions of
the world, e.g. Richardson et al. (2021) in Australia. One area of research interest is select-
ing for feed efficiency using residual feed intake (Basarab et al. 2013; De Haas et al. 2011)
which was observed to have heritability of about 0.26 to 0.58 (Hendriks et al. 2013). The
use of genetic and phenotypic correlations of C H4 outputs with various production traits
could also be of considerable use for identifying more efficient animals (Pinares-Patiño
et al. 2013). Application of genomics opens the possibility to efficiently select for hard to
13
measure traits (Oddy et al. 2014). For instance, application of genomics to identify rumen
microbiota related to CH4 and their association with cattle feed efficiency (Hernandez-
Sanabria et al. 2012) would not only be fast but an efficient way of formulating feeding
manipulation strategies to reduce CH4 emissions. The development of these proxies with
high correlation with C H4 production would be beneficial for identifying animals with low
CH4 emissions (Negussie et al. 2017). Once these genetic biomarkers for low C H4 emis-
sions have been identified in Western breeds, the same genetic biomarkers might, in the
future, be able to be used to select low CH4 emitting cattle of African breeds.
5 Conclusion
Based on the present review, use of alternative grazing species, e.g. leucaena, on-farm pas-
ture management, use of seaweed, tannins, FA, concentrates and the use of genomics are
possible strategies for reducing rumen CH4 production in the SADC region. Use of cereal
crop residues needs bioprocessing to improve nutritive value and digestibility, hence mak-
ing rumen metabolism more efficient. These will depend on the region’s strategy for SDGs
and the Paris Accord as some of the proposed interventions will compete with humans
for food such as the use of concentrate and edible oil. However, condensed tannins, lipids
and essential oils from indigenous plants (forages, trees and shrubs) including seaweed
could play a critical role in manipulating rumen fermentation to mitigate C H4 production
in this region. When implementing these mitigation strategies, the region should not lose
sight of global distorted CH4 accounting that focuses on isolated overemphasised metrics.
Such overestimating of emissions and inflated effects of C H4 from ruminants may result
in unnecessary costs further impoverishing resource limited communities in the SADC
region.
Acknowledgements We thank Dr Peter Moate (Agriculture Victoria Research, Australia) for his helpful
comments and critiquing the manuscript. Authors also want to thank Mr. Zelin Li (The University of Mel-
bourne, Australia) for help with formatting this paper.
Funding This manuscript forms part of MS’s BSc final year research dissertation and was funded by Bot-
swana University of Agriculture and Natural Resources.
Declarations
Competing interests The authors declare no competing interests.
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Mitig Adapt Strateg Glob Change (2022) 27: 47

Ruminant contribution to enteric methane emissions

Mompoloki Seketeme1 · Othusitse R. Madibela1 · Thabo Khumoetsile1 ·

Keywords Grazed forage · Greenhouse gas emissions · Methane · Mitigation · Ruminants

Livestock production is an important socio-economic activity for most people living in

3.1 Methane production in the rumen and its importance

Some of the dominant methanogenes in the rumen include Methanobrevibacter ruminan-

4 Strategies to reduce methane emissions in the SADC region

4.1 Feeding management and methane production

4.2 Use of monensin as a feed additive

cephala using rumen liquor (1000 mg) with low

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

4.3 Use of tannins as feed additives

by reducing fibre digestibility, perhaps because C ­ H4 production is associated with highly

4.4 Use of fatty acids as feed additive

tion with rumen Tifton 85 concen- media (24 h mL/g DM

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

4.6 Genetic selection of ruminants for low methane emissions

Although the magnitude of emissions of C ­ H4 expected to be reduced by selecting for low

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3.1 Methane production in the rumen and its importance

4 Strategies to reduce methane emissions in the SADC region

4.1 Feeding management and methane production

4.2 Use of monensin as a feed additive

4.3 Use of tannins as feed additives

by reducing fibre digestibility, perhaps because C H4 production is associated with highly

4.4 Use of fatty acids as feed additive

4.6 Genetic selection of ruminants for low methane emissions

Although the magnitude of emissions of C H4 expected to be reduced by selecting for low