Archivos Latinoamericanos de Producción Animal. 2023. 31 (1)
The carbon footprint of beef production from cull cows finished on
sown pastures in the savannas of the Colombian Orinoquía
Received 20210721; Accepted 20220224.
1Corresponding author: c.ramirez@creac.com.au
4Formely CIAT, Tropical Pastures Program, Km 17 CaliPalmira CP 763537, Apartado Aéreo 6713, Cali, Colombia.
5Formely Commonwealth Scientific and Industrial Research Organisation, (CSIRO) Agriculture, Australian Tropical Sciences and Innovation
Precint, James Cook University, Bebegu Yumba Campus, Townsville, QLD 4811, Australia.
1
Raul R. VeraInfanzón2,4
Carlos A. RamírezRestrepo1,4,5
Abstract. Neotropical savannas of the Colombian Orinoquía are largely dedicated to yearround beef production.
There is evidence of sustainable animal production in this savanna environment, but the links among animal
lifetime performance, greenhouse gas emissions, and soil organic carbon (SOC) accumulation at the system level
have been poorly studied to date. The objective of this study was to estimate the carbon (C) footprint of beef
production from Brahman (Bos indicus) cull cows finished on contrasting C4grassbased pastures in the Orinoco
River Basin. Longterm individual variations of liveweights and reproductive performance were used in an Excel®
dynamic model to estimate dry matter intake, methane (CH4) emissions, carcass traits, and the C footprint at the
farm gate. Values from the developed database were computed for cows born and raised on the savanna, bred on
Brachiaria decumbens, and later finished on B. humidicola [Scenario (SCE) 1, SCE 2]; B. decumbens (SCE 3); Andropogon
gayanus + Melinis minutiflora + Stylosanthes capitata (SCE 4); and A. gayanus + S. capitata (SCE 5) pastures. We
estimated the C footprints of SCE 1, SCE 3, and SCE 5 using published values of the rates of emission of CH4 and
nitrous oxide from the soil, feces, and urine; and accumulation of SOC in the soil during the fattening period. Most
of the estimated overall C footprint values at the system level were negative due to expected net SOC accumulation
during the fattening period. Depending on the expected quality of management, systems ranged from near
equilibrium in C balance to net increases in SOC accumulation.
Keywords: beef, methane, scenarios, soils, tropical forages
Huella de carbono en vacas de descarte cebadas en pasturas mejoradas de las
sabanas de la Orinoquia de Colombia
Resumen. Las sabanas Neotropicales de la Orinoquía de Colombia están dedicadas principalmente a la producción
de carne. Hay evidencias de que estos ecosistemas ganaderos pueden ser sostenibles, sin embargo, poco se conoce
acerca de las relaciones entre el desempeño animal, las emisiones de gases invernadero, y la acumulación de
carbono orgánico en el suelo (SOC) a nivel de sistema productivo. El objetivo del presente estudio fue estimar la
huella de C de la producción de carne, utilizando vacas (Bos indicus) de descarte cebadas en praderas contrastantes
de plantas C4. Se utilizaron variaciones reales e individuales y a largo plazo de peso y desempeño reproductivo de
las vacas a través de un modelo dinámico en EXCEL® para estimar el consumo de materia seca, las emisiones
entéricas de metano (CH4), las características productivas y ambientales de las carcasas, y la huella de carbono (C) a
nivel de finca. Las vacas nacidas, criadas y levantadas en sabana nativa, mantenidas en Brachiaria decumbens durante
su vida reproductiva fueron finalizadas en los siguientes escenarios (SCEs) modelados: Braquiaria humidicola (SCE 1
y SCE 2), B. decumbens (SCE 3), pastura mixta de Andropogon gayanus + Melinis minutiflora + Stylosanthes capitata (SCE
4) y pastura mixta de A. gayanus + S. capitata (SCE 5). La modelación se complementó con datos publicados de tasas
de emisión de CH4 y óxido nitroso del suelo, muestras de heces y orina, así como la acumulación de SOC en el
suelo durante la ceba. Las estimaciones de las huellas de C de los sistemas fueron mayoritariamente negativas
debido a la acumulación neta de SOC durante el período de finalización productiva. Dependiendo de la calidad del
manejo, los sistemas variaron entre equilibrio en el balance de C a aumentos netos de acumulación del SOC.
Palabras clave: carne, metano, escenarios, suelos, forrajes tropicales
www.doi.org/10.53588/alpa.310101
1CR Ecoefficient Agriculture Consultancy (CREAC®), Research and Education, 46 Bilbao Place, Bushland Beach, QLD 4818,
Australia. 2Consultant, 2 Norte 443, Viña del Mar, Chile. 3International Center for Tropical Agriculture (CIAT), Km 17 Cali
Palmira CP 763537, Apartado Aéreo 6713, Cali, Colombia.
Idupulapati M. Rao3
2
The world is facing challenges of population growth,
increased hunger, malnutrition, food and feed
insecurity, extreme events, and inequalities at multiple
scales [Food and Agriculture Organization of the
United Nations (FAO), 2009, 2015; Mora et al., 2018].
These challenges apply to Colombia with its growing
population, its allocation of resources, and the
vulnerability of its ecosystem services to climate
variability (Córdoba et al., 2019). The Neotropical
savannas environment of the Orinoco River Basin
represents 30.4 % of continental Colombia (Andrade et
al., 2009), and it is generally considered a major,
underutilized but fragile and biodiverse land resource
(RomeroRuiz et al., 2012). This socioculturalrich
region (Navas os, 1999; RomeroRuiz et al., 2012) is
also believed to have considerable potential to satisfy
increased beef demand, stock turnover efficiency, and
farm profitability (RamírezRestrepo and Vera, 2019;
RamírezRestrepo et al., 2023) based on yearround
grazing on tropical pastures.
In this context, the reliability of evaluating pastoral
systems’ productivity and environmental impact lies in
the reproducibility and replicability of raw data to
refine concepts, hypotheses, and theories and assist in
comparing different development views (Eckard et al.,
2014; Colquhoun, 2017; RamírezRestrepo et al., 2019b,
2019c). On this basis, the preservation, reuse, and
repurposing of existing databases can address new
demands on science (Griffin, 2015; Murdy et al., 2015;
Wyborn et al., 2015). This also applies to environmental
metrics of cattle and their relationship with soil organic
C (SOC) accumulation at the system level (Rao et al.,
2015; RamírezRestrepo et al., 2019a; RamírezRestrepo
et al., 2023). To our knowledge, there are no studies
using longterm individual legacy data of cull cows
between birth and slaughter regarding land use, dry
matter intake (DMI), methane (CH4) emissions, and
animal performance to estimate the C footprint of cull
cows at the system level. Recent C footprint
estimations (RamírezRestrepo et al., 2020a) with year
round grazing of Brahman (Bos indicus) breeding herds
suggest that cull cows fattening phase may assist in
maintaining and even enhancing SOC stocks in
conservatively managed improved sown pastures.
We hypothesized that the fattening of cull cows
derived from tropical beef herds monitored over
several reproductive cycles would allow the estimation
of production efficiency and the C footprints of beef
produced from cull cows grazing on contrasting
tropical pastures. The objective of this study was to
estimate the C footprint of beef production from
Brahman cull cows finished on contrasting C4grass
based tropical pastures in the Orinoco River Basin of
Colombia.
Introduction
Pegada de carbono em vacas de descarte engordadas em pastagens melhoradas
nas savanas da região de Orinoquia da Colômbia
Resumo. As savanas Neotropicais da Orinoquia Colombiana são principalmente dedicadas à produção de carne. Há
evidências de que os ecossistemas pecuários podem ser sustentáveis, no entanto, pouco se sabe sobre as relações
entre o desempenho animal, as emissões de gases de efeito estufa e o acúmulo de carbono orgânico en el suelo
(SOC) no nível do sistema de produção. O objetivo do presente estudo foi estimar a pegada C da produção de carne,
utilizando vacas de descarte (Bos indicus) terminadas em pastagens melhoradas (C4) da região de Orinoquia.
Variações reais, individuais e de longo prazo para peso e desempenho reprodutivo dessas vacas foram usadas em
um modelo dinâmico do EXCEL® para estimar o consumo de matéria seca, emissões de metano entérico (CH4),
características produtivas e ambientais de vacas, carcaças e pegada de carbono (C) ao nível de fazenda. Os valores
da base de dados foram calculados para vacas nascidas e criadas em pastagens de Brachiaria decumbens e
posteriormente as vacas foram terminadas em B. humidicola (Cenário (SCE) 1, SCE 2); B. decumbens (SCE 3);
Andropogon gayanus + Melinis minutiflora + Stylosanthes capitata (SCE 4); e A. gayanus + S. capitata (SCE 5). Nós
estimamos a pegada de C nos cenários SCE 1, SCE 3 e SCE 5 utilizando valores de literatura para níveis de emissão
de CH4 e óxido nitroso do solo, fezes e urina; e acumulação de SOC no solo durante o período de engorda. A
maioria dos valores globais estimados para a pegada C ao nível do sistema foram negativos devido à acumulação
líquida esperada de SOC durante o período de engorda. A maioria dos valores globais estimados para a pegada C
ao nível do sistema foram negativos devido à acumulação líquida esperada de SOC durante o período de engorda.
Dependendo da qualidade do manejo, os sistemas variaram entre equilíbrio no balanço de C a aumentos líquidos
no acúmulo de SOC.
Palavraschave: carne, metano, cenários, solos, forrageiras tropicais
RamírezRestrepo et al.
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Material and Methods
The database used for modeling
Heritage files generated from longterm grazing
experiments carried out at Carimagua Research Centre
(CRC: latitude 4o36’44.6” N, longitude 74o08’42.2” W)
in the Neotropical savannas (Llanos) environment of
Colombia were used as a source of animal data. The
constructed large database included individual
liveweight (LW), LW gains (LWG), and reproductive
performance records from birth to the end of cow’s
reproductive performance in Brahman breeding herds
over six continued reproductive cycles (RCs;
conceptionweaning; Vera et al., 2002; Ramírez
Restrepo et al., 2020a) lasting up to 10 ± 0.13 years.
Females had two provenances, the first one consisting
of animals born and raised on native savannas up to
272.6 ± 2.94 kg LW (25.87 ± 0.635 months), and the
second one was born and raised on wellmanaged
Brachiaria decumbens Stapf (syn. Urochloa decumbens)
cultivar (cv) Basilisk pastures with higher LWG up to
comparable initial breeding LW. In this paper, only
cows initially raised on savanna were included as the
production system represents the more
environmentally costly path and the most common
practice. Comparable shorterterm observations in
several commercial ranches of the region (Vera and
Seré, 1989; Astigarraga and Ingrand, 2011), although
not used in the present study, further support the
magnitudes and trends of the assembled dataset.
Soil properties and environmental conditions
Mean values for soil physi
cal and chemical charac
teristics (020 cm depth) before pasture establishment
(details given below) were 19.8 % sand, 40.2 % clay, 2.9
% organic matter, 4.78 pH, 90.1 % Al saturation, 1.46
µg/g available P (BrayII), and 0.17, 0.08 and 0.06 cmol/
kg of exchangeable Ca, Mg and K, respectively. These
values are similar to the subgroup of farms located on
relatively heavier soils monitored by Vera and Hoyos
(2019) and for which animal outputs were available.
Mean values of 2.202 mm of annual rainfall and 26.5
°C of ambient temperature were recorded at CRC
between 1979 and 1991. The wet season occurred
during the April to November period, while across the
years, January and June were the driest and wettest
months; and July and March were the coldest and
hottest months, respectively (Figure 1).
Pastures and forage quality characteristics
The pastures used to fatten cull cows, and their
botanical composition (%) on offer were as follows: B.
humidicola cv Llanero (Rendle) Schweick (syn. Urochloa
humidicola B. decumbens; A. gayanus Kunth cv
Carimagua + M. minutiflora P. Beauv + S. capitata
Vogel cv Capica (~ 70: 20: 10); and A. gayanus + S.
capitata (~ 95: 5). The pastures were established with
the recommended level of fertilizer application (kg/
ha) of 20 P, 20 K, 48 Ca, 14 Mg and 10 S, while a third
of that rate was used as maintenance fertilization
every third year (RamírezRestrepo and Vera, 2019;
RamírezRestrepo et al., 2020a; VeraInfanzón and
RamírezRestrepo, 2020). At the beginning of the
experiments, pastures were 34 years old, thus largely
excluding the rapid decline in animal productivity
reported by Vera and Hoyos (2019) in the initial years
following pasture establishment.
Pasture management during the fattening period
consisted of continuous grazing during the rainy
season. Cull cows were sold towards the end of the
rainy season (Figure 1). All pastures were managed to
conserve adequate soil cover and aboveground
biomass as described by Vera and RamírezRestrepo
(2017).
Measurements of the forage biomass and its botani
cal composition in the field during pregrazing and
postgrazing herbage periods were carried out
approximately every two months according to the
BOTANAL procedure (Jones and Tothill, 1985). Forage
on offer in the A. gayanus pastures exceeded 6500 kg
DM/ha, and at the end of the rainy season standing
forage always exceeded 3000 kg DM/ha. Equivalent
forage biomass values for B. humidicola and B.
decumbens pastures were 3500 and 1200 kg DM/ha,
respectively.
Total crude protein (CP) and neutral detergent fiber
(NDF) concentrations (g/kg DM) during the wet
period were derived from the CRC grazingforage
database constructed by RamírezRestrepo and Vera
(2019). Nutritive values (CP vs NDF) were 7337 vs
825785, 75110 vs 710650, 8590 vs 740746, and 101
66 vs 757760 for the B. humidicola, B. decumbens, A
gayanus + M. minutiflora + S. capitata, and A. gayanus +
S. capitata pastures, respectively
3
Sustainable fattening of cull beef cows
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
4
Modeling approach
The LWs of cull cows with different past repro
ductive performances were used to derive longterm
trends in LWs, CH4 emissions, and DMI, and these
were expanded by using recorded cumulative LWGs
(kg) over varied fattening periods (RamírezRestrepo
and Vera, 2019). In the original field experiment, cows
served as the experimental units. However, when
cows were grouped in a paddock for fattening
scenarios, those fattening scenarios represent the
experimental units rather than individual cows
themselves.
Initial and final cold carcass weights (CCW) of all
animals were estimated at the start and the end of the
fattening phase using a 0.4772 ratio of CW to LW
(Velásquez and Ríos, 2010) to determine CCW gains
(CCWG). These specifications allowed the model to
estimate final LWs, carcass traits, the full cycle of
production, production efficiency, and the C footprint
at the system level. This study does not include any
quantification from calves’ data that were analyzed in
detail by RamírezRestrepo et al. (2020a).
Developing scenarios
A similar stocking rate (SR; 1.3 UA/ha, 585 kg LW/
ha) was used to compare five fattening scenarios
(SCEs). Confidence intervals across all SCs and RCs
were calculated for initial LWs (360.6403.4 kg); daily
LWGs (0.3840.573 kg target LWs (439493 kg); and
age (98.49109.08 months) at slaughter. Following the
methodology reported by Velásquez and Ríos (2010),
carcass attributes of cull cows were calculated to
estimate the interaction among LW, age and DMI
dynamics, partial and lifetime CH4 emissions, and
slaughter data (RamírezRestrepo and Vera, 2019). To
derive carcass CO2eq efficiency indices, for CH4
estimation, the century horizon global warming
potential (GWP) of 34 with the inclusion of climateC
feedback was used (Gasser et al., 2017; Mueller and
Mueller, 2017; Allen et al., 2018) to allow for
compatibility with authors’ previously published work
(RamírezRestrepo and VeraInfanzón, 2019; Ramírez
Restrepo et al., 2019a, 2020a, 2023).
The first and second scenarios (SCE 1, SCE 2)
represent the performance of cull cows on an
overgrazed versus a wellmanaged B. humidicola
pasture to yield low and high daily LWGs,
respectively as reported by VeraInfanzón and
RamírezRestrepo (2020). Scenario 3 refers to the
performance of cull cows on B. decumbens pasture that
serves as local control and was initially reported by
RamírezRestrepo et al. (2020a). Based on Ramírez
Restrepo and Vera (2019), SCE 4 and SCE 5
represented the performance of cull cows grazed on
grasslegume associations of A. gayanus + M.
minutiflora + S. capitata, and A. gayanus + S. capitata
pastures, respectively. All sown pastures were 5 to 9
years old when the cows were culled.
Figure 1. Average monthly precipitation (∆) and ambient air temperature (●) recorded over 12 years by the meteorological
station at Carimagua Research Centre in the eastern plains of Colombia.
RamírezRestrepo et al.
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
5
Animals
As indicated before, females were monitored from
birth (26.2 ± 0.12 kg) and were weaned (157 ± 2.82 kg;
Rivera, 1988), and raised on savanna until mating (241
± 9.76 g/day; Vera and Hoyos, 2019). Cows were
offered ad libitum access to fresh water and a standard
mineral formulation of 175 Na, 269 Cl, 137 Ca, 80 P, 20
S, 1.04 Cu, 3.5 Zn, and mg of 10 Co and 76 I g/kg of a
commercial product. Registered Colombian Doctors of
Veterinary Medicine ensured animal welfare
regulations across all livestock manipulations in the
field (Law 73, MINEDUCATION, 1985).
Calculating intakes
Dry matter intake was calculated as an LW function
of feeding daily ad libitum on a DMI basis (i.e., 2.1 % of
total LW; Fisher et al., 1987) from measurements in
opencircuit chambers in tropical Australia (Ramírez
Restrepo and Vera, 2019) using a leastsquares
interceptslope regression (Equation 1) specified as:
DMI kg/day = 2.216 1.315) + 0.014 0.003) LW
(kg)
r2 = 0.491, P < 0.01; CV = 18.94; r.s.d = 1.34; r = 0.701,
P < 0.01
Predictions of DMI with Equation 1 fall within the
range of field measurements by Hess (1995), and
Pereira et al. (2009) for B. humidicola.
Estimating ruminant CH4 emissions
Using the structured daily DMI regression form,
individual estimations of CH4 emissions were based
on the simple relationship between LW per animal and
CH4 emissions per day in calibrated (pure CH4 gas 0.99
recoveries) opencircuit chambers (Equation 2). This
was recorded from wellcared and trained Brahman
(RamírezRestrepo et al., 2016b) and Belmont Red
Composite [Brahman x Africander (African Sanga) x
HerefordShorthorn (3/4 B. taurus RamírezRestrepo
et al., 2014, 2016a] steers fed ad libitum.
CH4 g/day = 16.176 21.087) + 0.324 0.057) LW
(kg)
r2 = 0.663, P < 0.0001; CV = 16.78; r.s.d = 30.82;
r = 0.814, P < 0.0001
Thus, predicted CH4 emissions are consistent with
the estimations of Cottle and Eckard (2018) and are
also consistent with recently published reports
(RamírezRestrepo and Vera, 2019; RamírezRestrepo
et al., 2019a, 2020a, 2023). As expected, this allows the
present results also to be comparable with other
tropical cattle system values (KuVera et al., 2018).
Lastly, we deliberately ignored issues related to CO2
eq costs involving transportation, supply chains, the
slaughterhouse, and the retail meat market.
Estimating CO2eq CH4 footprint, C stocks, rate of
SOC accumulation, and overall C footprint
Greenhouse gas emissions and C footprints were
calculated for three contrasting scenarios (SCE 1, SCE
3, and SCE 5) that were chosen to represent a wide
range of cattle fattening systems currently in use.
Some required parameters were obtained from
adjoining, mostly contemporary, paddocks (Rao, 1998;
Fisher et al., 1998; Rondón et al., 2006). These were
complemented when required with data obtained
under comparable Neotropical savannas, including
abundant data from the Venezuelan savannas
reviewed by Castaldi et al. (2006). Emissions from soils,
feces, and urine were estimated using the published
values on emission factors from the literature, and in
all cases, we used the maximum positive values
reported. Soil emission factors used for CH4 and
nitrous oxide (N2O) were adopted from Castaldi et al.
(2006) of the Venezuelan savannas. Although acid
savanna soils have been reported to act as a seasonal
sink, rather than a source, particularly for CH4, this
was not considered in the present estimations. Values
on N intake by animals were calculated from the
predicted DMI as explained above, and forage N
contents were reported in several grazing experiments
by Lascano and Thomas (1990), Lascano and Euclides
(1996), and from our data. Nitrogen digestibility was
calculated as in Glover et al. (1957), and urinary N
output as per Equation 1 reported by Waldrip et al.
(2013). The resulting animal N balance was checked
for compatibility with the observed LWG against the
requirements suggested by Rotta et al. (2016) for
tropical cattle in Brazil. Maximal fecal emissions were
estimated from the emission factors derived by Zhu et
al (2018), whereas those for fecal and urinary N2O
emissions were derived from Lessa et al. (2014).
Conversion of native savanna to sown pastures
slightly modifies soil emissions as reviewed by
Castaldi et al. (2006) for the seasonally dry Neotropical
savannas. Data from GarcíaMontiel et al. (2001) and
Castaldi et al. (2006) showed an average value of soil
CH4 emissions during the wet season as 1.18 mg/m2
day, but it is noted that soils under these conditions
may also be a net CH4 sink (Sanhueza et al., 1994a,
Sustainable fattening of cull beef cows
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
6
Animal performance and CH4 emission metrics
The effects of SCEs on calculated DMI, LWGs,
slaughter age, carcass traits, daily CH4, CH4 yield (CH4
g/kg DMI), periodCH4 emissions, and CO2eq CH4
concentration in carcass yields are shown in Table 1.
Compared to cull cows on the mixed A. gayanus + S.
capitata pasture (SCE 5), cows on the first B. humidicola
pasture (SCE 1) had the lowest DMI, however, cows
grazed on the second B. humidicola (SCE 2), the B.
decumbens (SCE 3), and the A. gayanus + M. minutiflora
+ S. capitata pasture (SCE 4) performed similarly to cull
cows in the SCE 5. Accordingly, the model yielded
total DMIs of 1.20, 1.51, 1.46, 1.39, and 1.65 ± 0.026 t of
DMI from the first to the fifth fattening SCEs.
Relative to the B. humidicola low LWG scenario
(SCE 1), the remaining SCEs had higher numerical
LWGs and CCWGs of 19, 27, 46, and 49 %, and 46, 50,
63, and 95 %, respectively. This trend was repeated in
terms of lean meat and edible protein carcass gain
(LMCG, EPCG, Table 1).
Daily CH4 and CH4 yield emissions per animal for
the B. humidicola (SCE 1) were significantly (P 0.05)
lower than those of the A. gayanus + S. capitata (SCE 5),
but enteric emissions were similar among the other
SCEs (Table 1). Furthermore, the emission of CH4/kg
LWG decreased linearly from SCE 1 to SCE 5 (r2 =
0.88, P = 0.02, r = 0.93), amounting to a difference of
28 % between the two extreme SCEs. Hence, the
magnitude of CH4 emissions for the finishing period in
terms of kg CO2eq/kg CCWG, LMCG, and EPCG
decreased (P 0.0001) between SCE 1 and SCE 5 by 29
% (Table 1).
The effect of successive RCs on systems performan
ce is given in Table 2. There were significant
differences in DMI values between some RCs (P
0.0001). Lifetime DMIs (t DM) differed (P 0.0001)
among the consecutive RCs evaluated (12.19 ± 0.884 vs
17.65 ± 0.395 vs 23.38 ± 0.316 vs 25.10 ± 0.334 vs 27.39 ±
0.266). Emissions between RCs during the fattening
practice differed when expressed in kg CO2eq/kg
CCWG, with the largest difference amounting to 18 %
between RC 4 and RC 6 (P 0.0001). However, over
the lifetime of the animals, there was at most an 11 %
difference (P 0.0001) between RC 1 and RC 6, such
that the ranking of the RCs depends on the carcass
parameter used for the comparisons.
Estimated C footprint of beef production of cull
cows
Enteric emissions increased 30 % from SCE 1 to
SCE 5 (Table 3), but the magnitude of the changes in
1994b). Similarly, soil N2O emissions averaged 0.32 mg/
m2 per day, but the range of values includes negative
values as well. However, we followed a conservative
approach and used relatively high values for soil
emissions. Pasture establishment and maintenance
incur in GHG emission costs related to machinery use,
and the use of fertilizers, and these were calculated as
in RamírezRestrepo et al. (2020a). Pasture
establishment costs were prorated over the assumed
persistence of wellmanaged pastures for 15 years, as it
was documented for onfarm pastures by Vera and
Hoyos (2019) for similarly managed onfarm sown
grasslands.
The aboveground and belowground C stocks and SOC
accumulation rates from improved pastures were
estimated for the three contrasting SCEs to reflect
differences that are not usually captured by animal
performance variables and static characteristics.
Values on SOC accumulation derived from
contemporaneous experiments at CRC and nearby to
the current study area were used (Fisher et al., 1994;
Rao, 1998; Trujillo et al., 2006; Costa et al., 2022; Hyman
et al., 2022). The methods used to determine SOC
stocks in different pastures were described by Fisher et
al. (1994). Values used to show the range of pasture
biomass production of both aboveground (Fisher et al.,
1998; Rao, 1998; Grace et al., 2006) and belowground
(Fisher et al., 1998; Rao, 1998; Trujillo et al., 2006) are
summarized in Table 3.
Statistical analyses
Data were analyzed with the GLM procedure of
the Statistical Analysis System (SAS, 2016; version 3.5)
including the upper and lower confidence limits
(CLM) for each mean observation and CL for the
parameter estimates (CLPARM) methods. Models for
LW and age dynamics, DMI, CH4 emissions, carcass
features, and carcassrelated indices considered the
fixed effects of breedingherd RCs and animal
fattening SCEs and their interaction. The relationships
between total LWG and CH4 emissions per kg LWG
during the fattening process were performed using the
CORR and REG procedures. Significant differences
between leastsquares means ± standard errors of the
mean were declared when P ≤ 0.05.
Results
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fecal CH4 production was less. On the contrary and
associated with the increased N intake of SCE 5,
urinary N2O emission rose considerably. Gains in the
carcass, lean meat, and edible protein weights (Table 1)
differed between SCEs, with SCE 5 nearly doubling the
gains in SCE 1, and emissions per kg of gain in carcass
parameters diminished from SCE 1 to SCE 5 by 29 %.
Over the lifetime of the animals, differences between
SCEs in emissions per kg of the final carcass, lean meat
and edible protein weights were less pronounced, with
at most 12 % higher emissions in SCE 1 than in SCE 5
(Table 2). As before, the remaining SCEs showed
intermediate values between the two extremes. Thus,
the magnitude of the differences between SCEs
depended very much on the denominator used for the
comparisons, while LW determined the final point for
sales of cows.
The estimated CO2eq CH4 emission values from the
feces of SCE 3 were 9.2 % higher while SCE 5 was 13.7
% larger compared to SCE 1. Estimated CO2eq N2O
emission values from the feces of SCE 3 were 30 %
higher, while SCE 5 was 44 % higher compared to SCE
1. The CO2eq N2O emission values estimated from the
urine of SCE 3 were 2.11fold greater, while SCE 5 was
2.87fold greater compared to SCE 1. The contribution
of the mineral supplement consumed by cows to CO2
eq GHG emission was also greater for SCE 5 than SCE
3 and SCE 1. The estimated values of total CO2eq
GHG emissions (kg/ha) during the grazing period,
including CH4 from animals, CH4 and N2O from feces,
and N2O from urine together with the mineral
supplement consumed were 1.232; 1.483 and 1.640 for
SCE 1, SCE 3, and SCE 5, respectively. Total values of
CO2eq GHG emissions (kg/ha) during the grazing
period at the system level were 1.345; 1.613 and 1.781
for SCE 1, SCE 3, and SCE 5, respectively (Table 3).
Nevertheless, differences between SCEs are con
founded with the different final LWs (Table 1) and
consequently different lengths of the fattening period,
and these differences disappear if expressed per day,
with an average for total GHG emissions of 8.4 kg
CO2eq/ha day.
The stocks of SOC to 1 m soil depth for the medium
textured soils of all three scenarios (SCE 1, SCE 3, and
SCE 5) were estimated to be in the range of 130 to 160
Mg/ha based on the published reports cited in Table 3.
The range of values of the standing aboveground
biomass estimated from published reports of SCE 1
was 1.2 to 3.5 Mg/ha and this was markedly lower
than those of SCE 3 (5.15 to 9.07) and SCE 5 (3.0 to 6.5).
Standing belowground biomass values (Mg/ha)
estimated from published reports ranged from 2.8 to
9.3, 8.3 to 10.5, and 6.0 to 9.0 for SCE 1, SCE 3, and SCE
5, respectively. We assumed the same values for CO2
eq CH4 (3.2 x 34 kg/ha/year) and N2O (1.05 x 298 kg/
ha/year) emission from the soil for all three SCEs to
estimate differences among the three SCEs based on
the number of days of the grazing period.
The estimated emission values from fertilizers and
tillage were the same among the three SCEs since
pasture establishment methods were the same. Using
the published values of above and belowground
biomass and their contribution to the SOC
accumulation, we estimated a range of rate of SOC
accumulation from 1.0 to 3.0 Mg/ha/year (Table 3).
The estimated overall C footprint values at the
system level in CO2eq in Mg/ha were 0.140 to 3.110;
0.154 to 3.688; and 0.167 to 4.059 for SCE 1, SCE 3,
and SCE 5, respectively. These negative values imply
that all three systems showed small to high values of
net SOC accumulation after correcting for GHG
emissions from animals, soil, and external inputs to
the system.
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Table 2. Influence of computed finishing scenarios (SCEs)† on dry matter intake (DMI), methane (CH4) emissions, and carbon
dioxide equivalent (CO2eq)CH4 carcass efficiency indices of consecutive reproductive cycles (RCs), and on CO2eqCH4 lifetime
emissions of carcass traits in multiparous commercial Brahman (Bos indicus) cull cows
2RC 3RC 4RC 5RC 6RC
Fattening phase
Number of animals 30 29 24 18 11
DMI, kg/head/day 8.469 ± 0.345ab 8.727 ± 0.154a 7.717 ± 0.140c 7.981 ± 0.130bc 8.848 ± 0.104a
CH4, g/head/day 159.1 ± 7.88ab 165.0 ± 3.52a 141.9 ± 3.21c 147.9 ± 2.97bc 167.2 ± 2.37a
CH4, g/kg DMI 18.78 ± 0.173ab 18.89 ± 0.077a 18.33 ± 0.070c 18.50 ± 0.065b 18.91 ± 0.052a
Period CH4, kg/head 27.59 ± 1.374ab 28.61 ± 0.614a 24.61 ± 0.561c 25.66 ± 0.519bc 29.09 ± 0.414a
kg CO2eq/kg CCWG 23.48 ± 1.185ab 24.32 ± 0.530a 20.91 ± 0.483c 21.80 ± 0.447bc 24.73 ± 0.357a
kg CO2eq/kg LMCG 38.60 ± 1.950ab 40.03 ± 0.872a 34.41 ± 0.796c 35.88 ± 0.737bc 40.70 ± 0.588a
kg CO2eq/kg EPCG 148.46 ± 7.500ab 153.98 ± 3.354a 132.37 ± 3.062c 138.02 ± 2.835bc 156.57 ± 2.261a
SCE 1 SCE 2 SCE 3 SCE 4 SCE 5
Lifetime emissions
Number of animals 30 30 30 30 30
kg CO2eq/kg CCW 65.06 ±1.851a 61.82 ± 1.851ab 61.30 ± 1.851ab 60.01 ± 1.851b 58.22 ± 1.851b
kg CO2eq/kg LM 107.05 ± 3.046a 101.73 ± 3.046ab 100.87 ± 3.046ab 98.75 ± 3.046b 95.81 ± 3.046b
kg CO2eq/kg EP 411.76 ± 11.715a 391.29 ± 11.715ab 387.99 ± 11.715ab 379.82 ± 11.715b 368.50 ± 11.715b
Table 1. Dry matter intake (DMI), liveweight (LW) gain, slaughter age, carcass attributes, methane (CH4) emissions, and carbon
dioxide equivalent (CO2eq)CH4 carcass efficiency indices of commercial Brahman (Bos indicus) cull cows continuously mated
on B. decumbens pastures and fattened on grass alone Brachiaria humidicola (SCE 1, SCE 2), B. decumbens (SCE 3), and grass
legume association pastures of Andropogon gayanus + Melinis minutiflora + Stylosanthes capitata (SCE 4) and A. gayanus + S.
capitata (SCE 5)
SCE 1 SCE 2 SCE 3 SCE 4 SCE 5 Pooled s.e.m.
Number of animals 30 30 30 30 30
DMI, kg/head day 8.093b 8.337ab 8.342ab 8.419ab 8.551a 0.153
LW gain, kg/day 0.384 0.457 0.486 0.561 0.573 0.014
Final LW, kg 439c 465b 467b 475a 493a 10.8
Age at slaughter, months 103.0 103.0 103.9 103.6 104.5 2.30
Cold carcass weight gain (CCWG, kg)T 27.20 39.60 40.84 44.37 52.96 4.164
Lean meat carcass gain (LMCG, kg) 16.52 24.06 24.82 26.96 32.18 2.885
Edible protein carcass gain (EPCG, kg) 4.29 6.25 6.45 7.01 8.36 0.658
CH4 emission metrics
CH4, g/head day 150.6b 156.0ab 156.2ab 157.9ab 160.9a 3.50
CH4, g/kg DMI 18.57b 18.68ab 18.69ab 18.72ab 18.78a 0.077
Period CH4, kg/head 22.34e 28.35bc 27.50c 26.17d 31.20a 0.610
kg CO2eq/kg CCWG 27.98a 24.33b 22.88c 20.04de 19.99e 0.526
kg CO2eq/kg LMCG 46.05a 40.04b 37.66c 32.98de 32.90e 0.886
kg CO2eq/kg EPCG 177.12a 154.03b 144.85c 126.85de 126.55e 3.332
Adapted from Velásquez and Ríos (2010),RamírezRestrepo and Vera (2019), and Vera et al. (2002) and RamírezRestrepo et al. (2020a).
The leastsquares means values in the same row bearing different letters between SCEs are significantly different (a,e: P ≤ 0.05). Standard error of the mean (s.e.m.).
Brachiaria humidicola (SCE 1, SCE 2), B. decumbens (SCE 3), Andropogon gayanus + Melinis minutiflora + Stylosanthes capitata (SCE 4) and A. gayanus + S. capitata (SCE 5)
pastures.
Adapted from RamírezRestrepo and Vera (2019), and Vera et al. (2002) and RamírezRestrepo et al. (2020a). EPCG: Edible protein carcass gain. CCWG: Cold
carcass weight gain. LMCG: lean meat carcass gain. The leastsquares means (LSM) ± standard error of the mean in the same row bearing different letters are
significantly different (ad: P ≤ 0.05).
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Table 3. Estimated greenhouse gas (GHG) emissions and overall carbon (C) footprint of commercial Brahman (Bos indicus) cull
cows fattened on B. humidicola (SCE 1), B. decumbens (SCE 3), and Andropogon gayanus + Stylosanthes capitata (SCE 5) pastures
Scenarios Observations, data sources, and
emission factors
Parameters SCE 1 SCE 3 SCE 5
Days on a grazing system 148 176 194
Cows 30 30 30
SR, cows/ha 1.42 1.38 1.34 Average onranch and onstation SR values on
MTAC
Mean final LW, kg/head 439 467 493 Table 1 of this paper
CO2eq enteric CH4, kg/ha 1082 1289 1418
CO2eq CH4 from feces, kg/ha
113.8
124
.3 129.4 Zhu et al. (2018): EF dry season = 0.001, rainy =
0.003
CO2eq N2O from feces, kg/ha 6.17 8.02 8.87 Lessa et al. (2014): EF = 0.0014
CO2eq N2O from urine, kg/ha 28.44
59.96 81.71 Chirinda et al. (2019): EF = 0.016; Lessa et al.
(2014): EF = 0.0193
CO2eq of mineral supplement consu
med, kg/ha 1.52 1.75 1.87 Cardoso et al. (2010) cited by Cerri et al. (2016)
Total CO2eq GHG emissions, kg/ha 1232 1483 1640
Estimation of overall C balance
SOC to 1 m depth, medium texture
soil, Mg/ha 130  160 130  160 130  160 Fisher et al. (1994); Rondón et al. (2006); Ramírez
Restrepo et al. (2019a)
Standing aboveground (shoot) biomass,
DM Mg/ha 1.2  3.5 5.1  9.1 3.0  7.0 Fisher et al. (1998); Kanno et al. (1999); Rao et al.
(2001a); Grace et al. (2006)
Standing belowground (root) biomass,
DM Mg/ha 2.8  9.3 8.3  10.5 6.0  9.0 Fisher et al. (1998); Rao (1998); Kanno et al.
(1999); Rao et al. (2001b); Trujillo et al. (2006)
Total aboveground and belowground
biomass, DM Mg/ha 4.0  12.8 13.4  19.6 9.0  15.0
CO2eq CH4 emission from the soil, kg/ha 2.74 3.20 3.59 Castaldi et al. (2006)
CO2eq N2O emission from the soil, kg/ha 91.23 108.49 119.59 Castaldi et al. (2006)
CO2eq emission from fertilizer inputs and
tillage, kg/ha 24.64 24.64 24.64 EdwardsJones et al. (2009); Kim et al. (2011);
University of Arkansas (2019)
CO2eq total GHG emissions at the system
level, kg/ha 1345 1613 1781
SOC accumulation rate, Mg/ha/year
1.0 to 3.0
1.0 to 3.0 1.0 to 3.0 Fisher et al. (1994); Rao et al. (1998); Fisher et al.
(1998); Rondón et al. (2006); Fisher et al. (2007);
Costa et al. (2022); Hyman et al. (2022)
SOC accumulation, kg/ha 405 to
1215
482 to 1446 531 to 1593 SOC accumulation during the grazing period
CO2eq soil C accumulation, kg/ha
1485 to 4445
1767 to 530
1
1948 to 5840
SOC accumulation in CO2eq during the
grazing period
Overall C footprint at the system level in
0.140 to 3110
0.154 to 3688
0.167 to 4059
Estimated from the difference between GHG
CO2eq, Mg/ha emissions and the soil C accumulation during
the grazing period
CH4: Methane. CO2eq: Carbon dioxide equivalent. EF: Emission factor. Mg: megagram. LW: Liveweight. N2O: Nitrous oxide. MTAC: Medium texture acid soils.
SOC: Soil organic carbon. SR: Stocking rate. Negative CO2eq values for soil C for overall C footprint imply SOC accumulation. CO2eq emission values are
expressed for the total duration of the grazing period.
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Discussion
The use of legacy data in the present paper aligns
well with the views of many authors that reviewed the
issue by taking into consideration of reduced funding
and the need to take a fresher, and more
comprehensive view of longterm results from studies
on grazing systems (Rouquette et al., 2009; Black 2014,
2018). Some authors have even questioned the
occasional excess of collecting “futile data” (Tedeschi,
2019), rather than making fuller and more innovative
use of existing information. The present inclusion of
cull cows’ emissions adds to the need to consider full
cattle production cycles as discussed by Eckard et al.
(2014). Lastly, current cattle systems in the region
continue to be dominated by breeding herds grazed on
traditional lowinput Brachiaria spp. pastures [Romero
et al., 2018; AGROSAVIA, 2019; Encuesta Nacional
Agropecuaria (ENA), 2019].
Grazing systems and animal production
Early onfarm studies (Vera and Seré, 1989) noted
that the fattening of cull cows in extensive ranches
constitutes an opportunistic use of recently established
sown pastures and provides a quick and profitable
return on investments. Fat cull cows constitute 32 % of
the cattle market in the study region (Romero et al.,
2018), whereas males account for 53 % of the total, and
all of them are mostly raised and fattened on sown
tropical grass pastures. A recent survey (Romero et al.,
2018) revealed that the most important grasses during
the study were B. dictyoneura, B. humidicola, and B.
decumbens. Another survey (Díaz et al., 2018) carried
out in both the Orinoco River Basin and the Atlantic
coast of Colombia, verified the continued popularity of
those three grass species. The present study, therefore,
includes pastures that represent a range of current
alternatives for grass pastures differing in quality and
management demands. B. humidicola (SCE 1)
constitutes the lowest limit of that range, while the A.
gayanus + S. capitata (SCE 5) represents a higher quality
pasture that is admittedly more demanding of
management, particularly under conditions of
extensive production.
In this study, we used a combination of onstation
mid to longterm grazing data to estimate the C
footprint of beef production from cull cows finished on
contrasting C4grassbased pastures. To our
knowledge, this is the first such integrated analysis in
the Orinoco River Basin. In this context, the results
presented in Tables 2 and 3 show evidence for the
varying links between CH4 emissions, LWs, carcass
metrics, and SOC accumulation. Scenario 1 relied on
an intensively grazed B. humidicola pasture, a
management technique that is most frequently
observed on commercial ranches, whereas SCE 3
represents a traditional approach to fattening, and SCE
5 indicates the potential of a stable grasslegume
pasture which could be feasible under muchimproved
management and production conditions.
Animal performance, CH4 emissions, and carcass
measures
The cumulative emissions (Table 2) would justify
culling cows after the third weaning as emissions
subsequently increase again. The effects of this practice
might be an increased number of heifers (Vera
Infanzón and RamírezRestrepo, 2020) for earlier
breeder replacement and specialized farming systems
(RamírezRestrepo, et al., 2023) resulting in more stable
temporal emissions. Culling of older mature breeding
dams should result in more efficient cowcalf
operations with lower environmental impact. In
contrast, Roberts et al. (2015) suggested the opposite
view when applied to temperate, more intensive,
breeding systems. Nevertheless, given the diversity of
pastoralbeef farming systems in the Llanos
(CORPOICA, 2010; Vera and Hoyos, 2019), further
data and ruminant modelmediated predictions would
be desirable. A small further refinement of the carcass
analyses could be made if additional estimates of the
carcass: LW ratio, as opposed to the fixed 0.4772 value,
become available for cull cows in the study region.
Our results suggest a need to strategically switch
fertile heifers from grazing on native savanna or low
quality B. humidicola pastures to superior nutritive
value B. decumbens pastures and other types of grass
pastures in the mid to longterm (Vera et al., 2002;
RamírezRestrepo et al., 2020a). Previous Australian
studies (Marshall, 2010; Marshall and Smajgl, 2013;
Marshall et al., 2014) indicated that delaying emissions
would facilitate adaptation to climatedriven changes.
Furthermore, there is evidence from consumer
research in Colombia (Charry et al., 2019) that buyers
have preferences for beef welfare and ecofriendly
certified beef production. This means that some
consumer segments would pay about 52 % more when
in addition to welfare and ecofriendly labels, certified
reduction in GHG emissions of the marketed beef is
available (Charry et al., 2019).
The analysis of the present C balance at the system
level is conservative, and we avoided dealing with
current controversies on the biogenic C cycle in
grazing systems that have been previously addressed
by Adegosan et al. (2015) and Wiloso et al. (2016). This
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also includes the possible shortlife effect of emitted
enteric CH4 (GWP 020year average vs GWP 0100
year average; Mueller and Mueller, 2017; Allen et al.,
2018) from animals. Consistent with these views, we
have reported the actual amounts of CH4 and
discussed our calculations based on the CO2eq values
at 100 years’ average horizon that we consider to be a
conservative approach to emissions.
Cull cows in the present study, raised on savanna as
heifers, and bred and fattened on B. decumbens showed
average carcass CO2eq stores 1.1 times the amount of
CO2eq accumulated in their counterparts grazing
lifetime on only B. decumbens pastures (detailed data
not shown). These estimated values indicated that
although the same beef product is produced, the CO2
eq overhead of cull cows across their lives varies
considerably according to management in early life.
This differs from FAO (2013)’s beef production data
that showed a global life cycle analysis (LCA) value of
67.8 kg CO2eq/kg CW), or the LCA annual
estimations of Cardoso et al. (2016) on degraded
Brachiaria pastures (51.66 kg CO2eq/kg CW). These
contrasting differences suggest the need to evaluate
beef systems relative to culling strategies since the
present results showed that culling over the second or
third weaning is a practical farming management
response to mitigate climate change. Opencircuit
respiration chamber studies (Grandl et al., 2016) with
Brown Swiss cattle (B. taurus) have shown that CH4
production in heifers increases with age, while in
multiparous cows the highest emissions are linked to
their second and third lactations (4 to 6.5 years). This
was followed by slightly lower CH4 production in
older cows. In parallel, the highest magnitude of that
CH4 emission response from B. indicus cows in
lactation (1 to 6) was found by RamírezRestrepo et al.
(2020a) during the fourth lactation (9 years).
Estimated C footprints
Extensive grazing systems can accumulate SOC
under different soil, climate, and management
systems, as shown by metaanalysis and synthesis of
published results (Conant et al., 2017; Viglizzo et al.,
2019). Several practices, including proper grazing
management, fertilization, sowing of improved grass
and/or legume species, irrigation, and conversion
from cultivation, could contribute to an increase in
SOC accumulation to a range of 0.105 to more than 1
Mg C/ha/year (Conant et al., 2017) over the longer
time but as yet the undefined length of years. Results
from longterm grazing experiments conducted in
wellwatered tropical areas, using wellmanaged
Brachiaria grassbased pastures indicated that it is
possible to not only improve animal production but
also increase SOC stocks (da Silva et al., 2017; Segnini et
al., 2017; dos Santos et al., 2019) over the medium
term.
Tropical pastures with sown grasses with deep root
systems, such as B. humidicola and A. gayanus when
wellmanaged (Fisher et al., 2007) with proper grazing
and maintenance fertilizer application (Braz et al.,
2013; Saravia et al., 2014) can increase SOC stocks in
deep soil layers up to 1 m depth, while soils under
poorly managed or degraded pastures may lose SOC
over time (Boddey et al., 2010). In this study, our
assumed values of SOC accumulation rates of 1 to 3
Mg/ha/year were based on the published reports on
above and belowground net primary productivity
(NPP; Fisher et al., 1998; Kanno et al., 1999; Rao et al.,
2001a; Trujillo et al., 2006) and SOC accumulation in
longterm pastures in Colombia and Brazil (Fisher et
al., 1994, 2007; Bustamante et al., 2006; Braz et al., 2013;
Saravia et al., 2014; Baptistella et al., 2020; Hyman et al.,
2022).
Pasture utilization by grazing cattle under tropical
conditions is in the range of 20 to 30 % and 4050 % of
the consumed DM is returned as feces, so that the
annual amount of litter and feces returned to the soil
surface is in the range of 33.5 to 40.5 Mg/ha/year
(Fisher et al., 1998, 2007). This C return to soil depends
on the level of pasture utilization in the range of 13 to
16 Mg C/ha/year, assuming a value of 40 % C in DM.
The NPP of the aboveground biomass of A. gayanus
pastures in the eastern plains was estimated to be as
much as 43 Mg/ha/year (Fisher et al., 1998). Trujillo et
al. (2006) found that the NPP of belowground biomass
of wellmanaged grass alone (B. dictyoneura) pasture
was 30.0 Mg/ha/year, while the grasslegume (B.
dictyoneura + S. capitata) association was 31.34 Mg/ha
year. Brachiaria dictyoneura grass has been recently
classified as B. humidicola and it is similar in growth
habits to the B. humidicola grass used in this study.
Kanno et al. (1999) compared five different tropical
grasses in their differences in root distribution under
continuous grazing with low amounts of fertilizer
application in the Cerrados of Brazil. Although they
measured the total root biomass only up to 40 cm soil
depth, the values of different grasses ranged between
6 to 16 Mg/ha. Urquiaga et al. (1998) estimated that 54
% and 62 % of root C from A. gayanus and B. decumbens
pastures, respectively will be nondecomposable
thereby contributing to an increase in SOC
accumulation. The belowground root biomass and its
functional activity in tropical pastures result from root
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production, growth, mortality, and decomposition
processes (i.e., root turnover), which occur
simultaneously and fluctuate widely under grazing
conditions and the age of the pasture (Rao, 1998; Fisher
et al., 2007; Siqueira da Silva et al., 2019).
Root turnover in the grass was estimated to be 2.2
times in both grass alone (B. dictyoneura) and grass +
legume (B. dictyoneura + Arachis pintoi) pastures (Rao et
al., 2001b). If one assumes that the NPP of
belowground DM is similar to aboveground DM, the
total C inputs to a grazed pasture are estimated to be
in the range of 26 to 32 Mg/ha/year (Fisher et al.,
1998). Improved grasses used in this study differ in
their growth habits and values for both above and
belowground DM production as well as in litter
decomposition. Andropogon gayanus is an erect,
tussockforming tall grass, B. decumbens is low
growing, erect or decumbent, rhizomatous and
stoloniferous grass while B. humidicola is a strongly
stoloniferous and rhizomatous grass. The rate of litter
decomposition reported was lower for B. humidicola
and A. gayanus compared to B. decumbens. The legume
litter of S. capitata decomposes much faster than that of
the three grasses (Thomas and Asakawa, 1993).
Boddey et al. (2020) indicated that the amount of N
being recycled through aboveground plant litter could
be over 100 kg N/ha/year for the grasslegume
association; and they also pointed out that
belowground N recycling may be of the same
magnitude as the aboveground N (Trujillo et al., 2006).
Root tissue of higher C: N ratios in tropical grasses
(109224) could lead to slower decomposition and
formation of fewer microbial byproducts (Rao, 1998,
Trujillo et al., 2006; Dietzel et al., 2017). Root C: N ratios
could play a key role in contributing to a larger
proportion of SOC that is found in deeper soil layers
below 20 cm soil depth (Fisher et al., 2007). Under
pasture (B. brizantha) land use treatment, de Figueiredo
et al. (2010) reported that 48 % of the total SOC was in
the form of particulate organic carbon (POC), and a
high value of C: N ratio of 47.8 was observed with this
POC by these authors. Clearly, all of the above factors
may influence the range of the assumed values of SOC
accumulation.
The amount of SOC accumulation in longterm (up
to 9 years old) pure grass pastures in the eastern plains
of Colombia was about 3.0 Mg/ha/year but it was
markedly increased with a legume component in the
pasture (Fisher et al., 1994). The use of legume
components in association with grasses can not only
improve N supply to the grass but also considerably
enhance soil biological activity. This is through an
increase not only in terms of earthworm biomass and
soil aggregation (Lavelle et al., 2014) but also in the
rate of N mineralization (Rao et al., 1994). Proper
grazing management together with maintenance
fertilizer application every two to three years could
sustain both aboveground and belowground NPP
leading to significant amounts of SOC accumulation in
improved pastures in the eastern plains of Colombia.
We do not assume that SOC accumulation in
introduced pastures will either be constant or be
continued indefinitely. But two recent studies from the
Brazilian agroecosystems presented some interesting
observations. Durrer et al. (2021) used a 100year
observational chronosequence spanning primary
foresttopasture conversion and subsequent
secondary forest succession in the Amazon region and
observed a surprising increase in topsoil SOC
concentrations in pastures at 60 years following
conversion. To predict longterm changes in SOC
stocks after pasture intensification and diversification,
Damian et al. (2021) used the DayCent model and
simulated the effects of converting poorly managed
pastures (PMP) to more intensive and diversified
systems of pasture management, including fertilized
pasture (FP). They used field data collected from three
regions with contrasting climatic conditions. The
DayCent model estimated that the conversion of PMP
to FP increases the soil C stocks by 0.95 Mg/ha/year.
The model also estimated that the fertilization of the
pasture every year (FP) could result into higher SOC
stocks.
The differences in overall C footprint estimates
among SCE1, SCE 3, and SCE 5 were not that marked
at the lower range, but at the higher range, those were
larger due to a greater number of grazing days and the
resultant SOC accumulation. The overall C footprint
values shown in Table 3 include an interval of possible
values that range from near neutrality to high net soil
C accumulation (negative values). They reflect the
natural variability encountered due to soils,
management strategies, betweenyears variability, and
variation in the potential growth of different species.
The three different tropical grasses included in the
SCEs generally exhibit high belowground
productivity, thus accumulating large amounts of
organic C in depths up to 2 meters, a contribution
frequently underestimated in the past due to only
superficial soil measurements (Qi et al., 2019). The
values estimated for B. humidicola in SCE 1 are
particularly interesting since this germplasm accession
CIAT 679 has been consistently demonstrated to have
RamírezRestrepo et al.
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
Conflicts of interest: The authors declare that there has been no interest(s) that might raise the question of bias in
the reported research or the manuscript's inferences, opinions, or conclusions.
13
a high root growth rate, and very slow decay rate of
stored biomass, even in visually degraded, overgrazed
pastures (Chirinda et al., 2019). This greater ability of
B. humidicola pastures with lower values of
digestibility and N content may be associated with the
ability of biological nitrification inhibition (BNI), a trait
that even extends to inhibition of N release from urine
patches (Byrnes et al., 2017). These BNI effects were not
accounted for in our calculations, so the corresponding
overall C footprint values may be a conservative
estimate. On the contrary, generalizations of results for
SCE 5 based on A. gayanus grass are narrower, as there
is limited information on its environmental impact,
this species has a smaller niche, and it is more
demanding of adequate grazing management than
generally practiced in extensive systems. Nevertheless,
SCE 5 constitutes a prototype of a productive grass
legume pasture that has persisted well under good
management in onstation trials and a small number of
commercial ranches.
Conclusions
This study elaborates on realistic simulated SCEs of
beef production by culling cows in the Llanos of the
Colombian Orinoquía. It was made possible by the
intensive use of longterm onstation legacy data of
animal production and studies related to animal GHG
emissions and dynamics of plant biomass, both above
and belowground, complemented with data on soil
emissions and SOC accumulation. Large and
practically important differences were found in
emissions and footprints between the tropical pasture
systems studied. These differences were particularly
notable during the fattening phase of cull cows on the
three systems chosen to represent two widely used
production practices with grassalone pastures,
compared to one promising grasslegume pasture. The
results clearly demonstrated that given a fixed
fattening period during the wet season, system
performance depends very much on the daily weight
gains allowed for by the pastures used. Furthermore,
the present estimates of emissions and footprints
suggest that cows’ age at culling may significantly
impact system performance, an issue that deserves
further longterm research. The estimates of C
footprints based on substantiated rates of SOC
accumulation over the mediumterm show that the
tested system may compensate for animal and soil
GHG emissions. However, the net values may vary
over a relatively wide range and demand above
average grazing management strategies relative to
current practices.
Ethics statement: The research did not require ethical review and approval because the animal data that were used
in the study were repurposed from legacy files recorded ethically at the Carimagua Research Station (CRC).
Author contributions: CARR: Investigation, Data curation, Conceptualization, Methodology, Model development,
Writing, Review, and Editing. RRVI: Investigation, Data curation, Conceptualization, Methodology, Model
development, Writing, Review, and Editing. IMR: Methodology, Writing, Review, and Editing.
Funding: The original pastoral research conducted at the CRC was financially supported between 1979 and 1991 by
a core budget from the International Center for Tropical Agriculture (CIAT). This manuscript and model develop
ment were funded from 2017 to 2021 by CR Ecoefficient Agriculture Consultancy (CREAC©) and R. R. Vera
Infanzón Private Consultant Services.
Acknowledgments: We would like to thank the technical assistance given by many staff at the CRC and CIAT years
ago. Special thanks are extended to the Commonwealth Scientific and Industrial Research Organisation (CSIRO) for
allowing C.A. RamírezRestrepo to review, collate, and analyze the original dataset between 2016 and 2017.
Edited by Dr. Ana María Herrera Angulo and Dr. Claudia Faverin.
Sustainable fattening of cull beef cows
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
14
Literature Cited
Adesogan, A. T., J. C. Dubeux, and L. E. Sollenberger.
2015. Nutrient movements through ruminant
livestock production systems. In: M. M. Roy, D. R.
Malaviya, V. K. Yadav, T. Singh, R. P. Sah, D. Vijay,
A. Radhakrishna (Eds.). Proceedings of 23rd
International Grassland Congress. Range
Management Society of India. New Delhi. India pp.
79–94.
Boddey, R. M., D. R. Casagrande, B. G. C. Homem, and
B. J. R. Alves. 2020. Forage legumes in grass
pastures in tropical Brazil and likely impacts on
greenhouse gas emissions: A review. Grass Forage
Sci. 75(4): 357–371.
https://doi.org/10.1111/gfs.12498
AGROSAVIA. 2019. Adopción e impacto de los
sistemas agropecuarios introducidos en la
altillanura plana del Meta. Corporación Colombiana
de Investigación Agropecuaria (AGROSAVIA),
Mosquera, Colombia.
https://repository.agrosavia.co/handle/
20.500.12324/35451
Allen, M. R., K. P. Shile, J. S. Fuglestvedt, R. J. Millar,
M. Cain, D. J. Frame, and A. H. Macey. 2018. A
solution to the misrepresentation of CO2equivalent
emissions of shortlived climate pollutants under
ambitious mitigation. Clim. Atmos. Sci. 16: 1–8.
https://doi.org/10.1038/s4161201800268
Andrade, G. I. P., L. G. G. Castro, A. D. Durán, M. B.
Rodríguez, G. L. Rudas, E. B. Uribe, and E. H. Wills.
2009. La mejor Orinoquía que podemos construir.
Elementos para la sostenibilidad ambiental del
desarrollo. Universidad de los Andes, Bogotá,
Colombia.
Astigarraga, L., and S. Ingrand. 2011. Production
flexibility in extensive beef farming systems. Ecol.
Soc. 16(1): 1–7.
http://www.ecologyandsociety.org/vol16/iss1/
art7/
Baptistella, J. L. C., S. A. L. Andrade, J. L. Favarin, and
P. Mazzafera. 2020. Urochloa in tropical agroecosys
tems. Front. Sustain. Food Syst. 4: 119.
https://doi.org/10.3389/fsufs.2020.00119
Black, J. L. 2014. Brief history and future of animal
simulation models for science and application. Aust.
J. Agric. Res. 54: 1883–1895.
https://doi.org/10.1071/AN14650
Black, J. L. 2018. Perspectives on animal research and
its application. Anim. Prod. Sci. 56: 756–766.
https://doi.org/10.1071/AN15793
Boddey, R. M., C. P. Jantalia, P. C. Conceicao, J. A.
Zanatta, C. Bayer, J. Mielniczuk, J. Dieckown, H. P.
Santos, J. E. Denardin, C. Aita, S. J. Giacomini, B. J.
R. Alves, and S. Urquiaga. 2010. Carbon
accumulation at depth in Ferralsols under zerotill
subtropical agriculture. Glob. Chang. Biol. 16: 784–
795.
https://doi.org/10.1111/j.13652486.2009.02020.x
Braz, S. P., S. Urquiaga, B. J. R. Alves, C. P. Jantalia, A.
P. Guimaraes, C. A. Santos, S. C. Santos, E. F. M.
Pinheiro, and R. M. Boddey. 2013. Soil C stocks
under productive and degraded Brachiaria pastures
in the Brazilian Cerrado. Soil Sci. Soc. Am. J. 77: 914–
928. https://doi.org/10.2136/sssaj2012.0269
Bustamante, M. M. C., M. Corbeels, E. Scopel, and R.
Roscoe. 2006. Soil carbon and sequestration
potential in Cerrado region in Brazil. In: R. Lal, C. C.
Cerri, M. Bernoux, J. Etcherves, C. E. P (Eds.).
Carbon Sequestration in Soils of Latin America,
CRC Press, Boca Raton, USA, pp. 285–304.
Byrnes, R. C., J. Nùñez, L. Arenas, I. Rao, C. Trujillo, C.
Alvarez, J. Arango, F. Rasche, and N. Chirinda.
2017. Biological nitrification inhibition by Brachiaria
grasses mitigates soil nitrous oxide emissions from
bovine urine patches. Soil Biol. Biochem. 107: 156–
163. https://doi.org/10.1016/j.soilbio.2016.12.029
Cardoso, A. S., A. Berndt, A. Leytem, B. J. R. Alves, and
I. N. O., de Carvalho, L. H. d. B. Soares, S. Urquiaga,
and R. M. Bodde. 2016. Impact of the intensification
of beef production in Brazil on greenhouse gas
emissions and land use. Agr. Syst. 143: 86–96.
https://doi.org/10.1016/j.agsy.2015.12.007
Castaldi, S., A. Ermice, and S. Strumia. 2006. Fluxes of
N2O and CH4 from soils of savannas and seasonally
dry ecosystems. J. Biogeogr. 33: 401–415. https://
doi.org/10.1111/j.13652699.2005.01447.x
Cerri, C. C., C. S. Moreira, P. A. Alves, G. S. Raucci, B.
de A. Castigioni, F. F. C. Mello, D. G. P Cerri, and C.
E. P. Cerri. 2016. Assessing the carbon footprint of
beef cattle in Brazil: a case study with 22 farms in
the state of Mato Grosso. J. Clean. Prod. 112: 2593–
2600. https://doi.org/10.1016/j.jclepro.2016.02.032
Charry, A., M. Narjes, K. Enciso, M. Peters, and S.
Burkart. 2019. Sustainable intensification of bee
production in Colombia Chances for product
differentiation and price premium. Agric. Food
Econ. 7(22): 1–18.
https://doi.org/10.1186/s4010001901437
Chirinda, N., S. Loaiza, L. Arenas, V. Ruiz, C. Faverín,
C. Alvarez, J. V. Savian, R. Belfon, K. Zuniga, A.
MoralesRincon, C. Trujillo, M. Arango, I. Rao, J.
Arango, M. Peters, R. Barahona, C. Costa Jr., T. S.
Rosenstock, M. Richards, D. MartinezBaron, and L.
RamírezRestrepo et al.
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
15
Colquhoun, D. 2017. The reproducibility of research
and the misinterpretation of pvalues. Royal Society
Open Science. 4: 171085.
https://royalsocietypublishing.org/doi/pdf/
10.1098/rsos.171085
De Figueiredo, C. C., R. D. V. Siqueira, and C. M. A.
Carbone. 2010. Labile and stable fractions of soil
organic matter under management systems and
native Cerrado. Rev. Bras. Ciênc. Solo. 34: 907–916.
https://doi.org/10.1590/S010006832010000300032
Cardenas, 2019. Adequate vegetative cover
decreases nitrous oxide emissions from cattle urine
deposited in grazed pastures under seasonal
conditions. Sci. Rep. 9: 908.
https://doi.org/10.1038/s41598018374532
Conant, R. T., C. E. P. Cerri, B. B. Osborne, and K.
Paustian. 2017. Grassland management impacts on
soil carbon stocks: a new synthesis. Ecol. Appl. 27:
662–668. https://doi.org/10.1002/eap.1473
Córdoba, C. A. V., S. R. Hortúa, and T. LeónSicard.
2019. Resilience to climate variability: the role of
perceptions and traditional knowledge in the
Colombian Andes. Agroecol. Sustain. Food Syst.
44(4): 419–445.
https://doi.org/10.1080/21683565.2019.1649782
CORPOICA. 2010. Evaluación de crecimiento, calidad
de la canal y cortes de carne en cinco grupos raciales
bovinos de la Orinoquia Colombiana. Informe
Técnico Final Proyecto. Corporación Colombiana de
Investigación Agropecuaria (CORPOICA),
Ministerio de Agricultura y Desarrollo Rural
(MADR), Federación Colombiana de Ganaderos
(FEDEGAN), Corporación Comité de Ganaderos del
Meta, Villavicencio, Colombia.
Costa, C. Jr., D. M. Villegas, M. Bastidas, N. M. Rubio,
I. Rao, and J. Arango. 2022. Soil carbon stocks and
nitrous oxide emissions of pasture systems in
Orinoquía region of Colombia: Potential for
developing landbased greenhouse gas removal
projects. Front. Clim. 4: 916068.
https://doi.org/10.3389/fclim.2022.916068
Cottle, D. J., and R. J. Eckard. 2018. Global beef cattle
methane emissions: yield prediction by cluster and
metaanalyses. Anim. Prod. Sci. 58(12): 2167–2177.
https://doi.org/10.1071/AN17832
Da Silva, A. C., L. B. de Figueiredo, E. R. Janusckiewicz,
E. M. da Silva, R. B. Pavezzi, J. B. K. Werner, R. R.
Andrade, and A. C. Ruggieri. 2017. Impact of
grazing intensity and seasons on greenhouse gas
emissions in tropical grassland. Ecosystems. 20(4):
845–859.
https://doi.org/10.1007/s1002101600650
Damian, J. M., E. S. Matos, B. C. Pedreira, F. C. F.
Carvalho, L. M. Premazzi, S. Williams, K. Paustian,
and C. E. P. Cerri. 2021. Predicting soil C changes
after pasture intensification and diversification in
Brazil. Catena. 202: 105238. https://doi.org/10.1016/
j.catena.2021.105238
Díaz, M., D. Vergara, V. Castiblanco, and S. Burkart.
2018. Colombian cattle producers' preferences for
improved forage technologies: chances for forage
breeding and selection. Poster, TROPENTAG,
Ghent. https://cgspace.cgiar.org/bitstream/handle/
10568/97078/
Diaz_MF_et_al._2018_Colombian_Cattle_producers
_Preferences_for_Improved_Forage_Technologies_
web.pdf?sequence=1
Dietzel, R., M. Liebman, and S. Archontoulis. 2017. A
deeper look at the relationship between root carbon
pools and the vertical distribution of the soil carbon
pool. Soil. 3: 139–152.
https://doi.org/10.5194/soil31392017
Dos Santos, C. A., C. P. Rezende, E. F. Machado
Pinheiro, J. M. Pereira, B. J. R. Alves, S. Urquiaga,
and R. M. Boddet. 2019. Changes in soil carbon
stocks after landuse change from native vegetation
to pastures in the Atlantic Forest region of Brazil.
Geoderma. 337: 394–401.
https://doi.org/10.1016/j.geoderma.2018.09.045
Durrer, A., A. J. Margenot, L. C. R. Silva, B. J. M.
Bohannan, K. Nusslein, J. v. Haren, F. D. Andreote,
S. J. Parikh, and J. L. M. Rodrigues. 2021. Beyond
total carbon: conversion of amazon forest to pasture
alters indicators of soil C cycling. Biogeochemistry.
152: 179–194. https://doi.org/10.1007/s10533020
00743x
Eckard, R. J., V. O. Snow, I. R. Johnson, and A. D.
Moore. 2014. The challenges and opportunities
when integrating animal models into grazing
system models for evaluating productivity and
environmental impact. Anim. Prod. Sci. 54(12): 1896–
1904. https://doi.org/10.1071/AN14551
EdwardsJones, G., K. Plassmann, and I. M. Harris.
2009. Carbon footprint of lamb and beef production
systems: insights from an empirical analysis of
farms in Wales, UK. J. Agric. Sci. 147: 707–719.
https://doi.org/10.1017/S0021859609990165
ENA. 2019. Encuesta nacional agropecuaria (ENA).
Departamento Nacional de Estadística (DANE),
Bogotá, Colombia. https://www.dane.gov.co/
FAO. 2009. Highlevel expert forum How to feed the
world in 2050. Food and Agriculture Organization
of the United Nations (FAO), Rome, Italy.
FAO. 2013. Greenhouse emissions from ruminant
supply chains. A global life cycle assessment. Food
and Agriculture Organization of the United Nations
(FAO), Rome, Italy.
Sustainable fattening of cull beef cows
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
16
FAO. 2015. Climate change and food systems: Global
assessments and implications for food security and
trade. Food and Agriculture Organization of the
United Nations (FAO), Rome. Italy.
Grandl, F., S. L. Ameichanka, M. Furger, M. Clauss, J.
O. Zeitz, M. Kreuzeer, and A. Schwarm. 2016.
Biological implications of longevity in dairy cows: 2.
Changes in methane emissions and efficiency with
age. J. Dairy Sci. 99: 3475–3485.
https://doi.org/10.3168/jds.201510262
Fisher, M. J., S. P. Braz, R. S. M. dos Santos, S.
Urquiaga, B. J. R. Alves, and R. M. Boddey. 2007.
Another dimension to grazing systems: Soil carbon.
Trop. Grassl. 41: 65–83.
http://www.tropicalgrasslands.asn.au/
Tropical%20Grasslands%20Journal%20archive/
PDFs/Vol_41_2007/Vol_41_02_2007_pp65_83.pdf
Fisher, D., J. Burns, and K. Pond. 1987. Modeling ad
libitum dry matter intake by ruminants as regulated
by distension and chemostatic feedbacks. J Theor
Biol. 126: 407–408. https://doi.org/10.1016/S0022
5193(87)801480
Fisher, M. J., I. M. Rao, M. A. Ayarza, C. E. Lascano, J. I.
Sanz, R. J. Thomas, and R. R. Vera. 1994. Carbon
storage by introduced deeprooted grasses in the
South American savannas. Nature. 371: 236–238.
https://doi.org/10.1038/371236a0
Fisher, M. J., R. J. Thomas, and I. M. Rao. 1998.
Management of tropical pastures in acidsoil
savannas of South America for carbon sequestration
in the soil. In: R. Lal, J. M. Kimble, R. F. Follett, B. A.
Stewart (Eds.). Management of carbon sequestration
in soil (Advances in soil science). CRC Press, Boca
Raton, USA, pp. 405–420.
https://doi.org/10.1201/9781351074254
GarciaMontiel, D. C., P. A. Steudler, M. Piccolo, J. M.
Melillo, C. Neill, and C. E. Cerri. 2001. Controls of
soil nitrogen emissions from forest and pastures in
the Brazilian Amazon. Global Biogeochem. Cycles.
15(4): 1021–1030.
https://doi.org/10.1029/2000GB001349
Gasser, T., G. P. Peters, J. S. Fuglestvedt, W. J. Collins,
D. T. Shindell, and P. Ciais. 2017. Accounting for the
climatecarbon feedback in emission metrics. Earth
Syst. Dyn. 8: 235–253.
https://doi.org/10.5194/esd82352017
Glover, J., D. W. Duthie, and M. H. French. 1957. The
The apparent digestibility of crude protein by the
ruminant: I. A synthesis of the results of digestibility
trials with herbage and mixed feeds. J. Agric. Sci.
48(3): 373–378.
https://doi.org/10.1017/S0021859600031750
Grace, J., J. San José, P. Meir, H. S. P. Miranda, and R.
A. Montes. 2006. Productivity and carbon fluxes of
tropical savannas. J. Biogeogr. 33: 387–400.
https://doi.org/10.1111/j.13652699.2005.01448.x
Griffin, R. E. 2015. When are old data new data?
GeoResJ. 6: 92–97.
https://doi.org/10.1016/j.grj.2015.02.004
Hess, H. D. 1995. Grazing selectivity and ingestive
behaviour of steers on improved tropical pastures in
the eastern plains of Colombia. (Ph.D. Thesis). Swiss
Federal Institute of Technology Zurich, Zurich,
Switzerland.
Hyman, G., A. Castro, M. Da Silva, M.A. Arango, J.
Bernal, O. Perez, and I. M. Rao. 2022. Soil carbon
storage potential of acid soils of Colombia’s Eastern
High Plains. Front. Sustain. Food Syst. 6: 954017.
https://doi.org/10.3389/fsufs.2022.954017
Jones, R. M., and J. C. Tothill. 1985. BOTANAL  A field
and computing package for assessment of plant
biomass and botanical composition. In: J. C. Tothill,
J. J. Mott (Eds.). Proceedings of the International
Savanna Symposium. Australian Academy of
Science, Canberra, Australia, pp. 318–320.
http://hdl.handle.net/102.100.100/276273?index=1
Kanno, T., M. C. Macedo, V. P. B. Euclides, J. A. Bono,
Jr J. D. G. Santos, M. C. Rocha, and L. G. R. Beretta.
1999. Root biomass of tropical grass pastures under
continuous grazing in Brazilian savannas. Grassl.
Sci. 45: 9–14.
https://agris.fao.org/agrissearch/search.do?
recordID=JP1999005204
Kim, S. C., K. U. Kim, and D. C. Kim. 2011. Prediction
of fuel consumption of agricultural tractors. Appl.
Eng. Agric. 27(5): 705–709.
https://doi.org/10.13031/2013.39565
KuVera, J. C., S. S. ValenciaSalazar, A. T. Piñeiro
Vázquez, I. C. MolinaBotero, J. ArroyaveJaramillo,
M. D. MontoyaFlores, F. J. LazosBalbuena, J. R.
CanulSolís, J. I. ArceoCastillo, L. RamírezCancino,
C. S. EscobarRestrepo, J. A. AlayónGamboa, G.
JiménezFerrer, L. M. ZavalaEscalante, O. A.
CastelánOrtega, P. QuintanaOwen, A. J. Ayala
Burgos, C. F. AguilarPérez, and F. J. Solorio
Sánchez. 2018. Determination of methane yield in
cattle fed tropical grasses as measured in open
circuit respiratory chambers. Agric and For Meteor.
258: 3–7.
https://doi.org/10.1016/j.agrformet.2018.01.008
RamírezRestrepo et al.
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
17
Lascano, C., and V. P. B. Euclides. 1996. Nutritional
quality and animal production of Brachiaria
pastures. In: J. W. Miles, B. L. Maass, C. B. do Valle
(Eds.). Brachiaria: Biology, agronomy, and
improvement. CIAT, Cali, Colombia, pp. 106–123.
https://hdl.handle.net/10568/82028 Mueller, R. A., and E. A. Mueller. 2017. Fugitive
methane and the role of atmospheric halflife.
Geoinformatics & Geostatistics: An Overview. 5(3):
1–7. https://doi.org/10.4172/23274581.1000162
Lascano, C., and D. Thomas. 1990. Quality of
Andropogon gayanus and animal productivity. In: J.
M. Toledo, R. Vera, C. Lascano, J. M. Lenné (Eds.).
Andropogon gayanus Kunth. A grass for tropical acid
soils. CIAT, Cali, Colombia pp. 247–276.
https://hdl.handle.net/10568/54875
Lavelle, P., N. Rodríguez, O. Arguello, J. Bernal, C.
Botero, P. Chaparro, Y. Gómez, A. Gutiérrez, M. P.
Hurtado, S. Loaiza, S. X. Pulido, E. Rodríguez, C.
Sanabria, E. Velásquez, and S. J. Fonte. 2014. Soil
ecosystem services and land use in the rapidly
changing Orinoco River Basin of Colombia. Agric.
Ecosys. Environ. 185: 106–117.
https://doi.org/10.1016/j.agee.2013.12.020
Lessa, A. C. R., B. E. Madari, D. S. Paredes, R. M.
Boddey, S. Urquiaga, C. P. Jantalia, and B. Jr. Alves.
2014. Bovine urine and dung deposited on Brazilian
savannah pastures contribute differently to direct
and indirect soil nitrous oxide emissions. Agric.
Ecosys. Environ. 190: 104–111.
https://doi.org/10.1016/j.agee.2014.01.010
Marshall, N. A. 2010. Understanding social resilience to
climate variability in primary enterprises and
industries. Glob. Env. Change. 20: 36–43.
https://doi.org/10.1016/j.gloenvcha.2009.10.003
Marshall, N. A., A. and Smajgl. 2013. Understanding
variability in adaptative capacity on rangelands.
Rangeland Ecol. Manage. 66: 84–94.
https://doi.org/10.2111/REMD1100176.1
Marshall, N. A., C. J. Stokes, N. P. Webb, P. A.
Marshall, and A. J. Lankester. 2014. Social
vulnerability to climate change in primary
producers: A typology approach. Agric. Ecosys.
Environ. 186: 86–93.
http://dx.doi.org/10.1016/j.agee.2014.01.004
MINEDUCATION. 1985. Ley 0073 de Octubre 8 de
1985. Ministerio de Educación (MINEDUCATION),
Bogotá, Colombia.
https://www.mineducacion.gov.co/1759/w3
article103974.html?_noredirect=1
Mora, C., D. Spirandelli, E. C. Franklin, J. Lynham, M.
B. Kantar, W. Miles, C. Z. Smith, K. Freel, J. Moy, L.
V. Louis, E. W. Barba, K. Bettinger, A. G. Frazier, IX.
J. F. Colburn, N. Hanasaki, E. Hawkins, Y.
Hirabayashi, W. Knorr, C. M. Little, K. Emanuel, J.
Sheffield, J. A. Patz, and C. L. Hunter. 2018. Broad
threat to humanity from cumulative climate hazards
intensified by greenhouse gas emissions. Nat. Clim.
Chang. 8: 1062–1071.
https://doi.org/10.1038/s4155801803156
Murdy, J., J. Orford, and J. Bell. 2015. Maintaining
legacy data: Saving Belfast Harbour (UK) tidegauge
data (1901–2010). GeoResJ. 6: 65–73.
https://doi.org/10.1016/j.grj.2015.02.002
Navas Ríos, C. L. 1999. Caracterización socioeducativa,
evaluativa y comparativa de cuatro comunidades en
los Llanos Orientales de Colombia (Master Thesis).
Universidad de Antioquia, Medellín, Colombia.
Pereira, J. M., R. M. Tarré, R. Macedo, C. d. P. Rezende,
B. J. R. Alves, S. Urquiaga, and R. M. Boddey. 2009.
Productivity of Brachiaria humidicola pastures in the
Atlantic forest region of Brazil as affected by
stocking rate and the presence of a forage legume.
Nutr. Cycl. Agroecosystems. 83: 179–196.
https://doi.org/10.1007/s107050089206y
Qi, Y., W. Wei, C. Chen, and L. Chen. 2019. Plant root
shoot biomass allocation over diverse biomes: A
global synyhesis. Glob. Ecol. Conserv. e00606.
https://doi.org/10.1016/j.gecco.2019.e00606
RamírezRestrepo, C. A., C. J. O’Neill, N. López
Villalobos, J. Padmanabha, and C. McSweeney.
2014. Tropical cattle methane emissions: the role of
natural statins supplementation. Anim. Prod. Sci. 54:
1294–1299. http://dx.doi.org/10.1071/AN14246
RamírezRestrepo, C. A., C. J. O’Neill, N. López
Villalobos, J. Padmanabha, J. K. Wang, and C.
McSweeney. 2016a. Effects of tea seed saponin
supplementation on physiological changes
associated with blood methane concentration in
tropical Brahman cattle. Anim. Prod. Sci. 56: 457–
465. http://dx.doi.org/10.1071/AN15582
RamírezRestrepo, C. A., C. Tan, C. J. O’Neill, N.
LópezVillalobos, J. Padmanabha, J. K. Wang, and C.
McSweeney. 2016b. Methane production,
fermentation characteristics and microbial profiles
in the rumen of tropical cattle fed tea seed saponin
supplement. Anim. Feed. Sci. Tech. 216: 58–67.
http://dx.doi.org/10.1016/j.anifeedsci.2016.03.005
RamírezRestrepo, C. A., and R. R. Vera. 2019. Body
weight performance, estimated carcass traits and
methane emissions of beef cattle categories grazing
Andropogon gayanus, Melinis minutiflora and
Stylosanthes capitata mixed swards and Brachiaria
humidicola pasture. Anim. Prod. Sci. 56(4): 729–750.
https://doi.org/10.1071/AN17624
Sustainable fattening of cull beef cows
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
18
RamírezRestrepo, C. A., and R. R. VeraInfanzón.
2019. Methane emissions of extensive grazing
breeding herds in relation to the weaning and
yearling stages in the Eastern Plains of Colombia.
Rev. Med. Vet. Zoot. 66(2): 111–130.
https://doi.org/10.15446/rfmvz.v66n2.82429
RamírezRestrepo, C. A., R. R. VeraInfanzón, and I. M.
Rao. 2020a Predicting methane emissions, animal
environmental metrics and carbon footprint from
Brahman (Bos indicus) breeding herd systems based
on longterm research on grazing of neotropical
savanna and Brachiaria decumbens pastures. Agric.
Syst. 184: 102892.
https://doi.org/10.1016/j.agsy.2020.102892
RamírezRestrepo, C. A., R. R. VeraInfanzón, and I. M.
Rao. 2023. The carbon footprint of youngbeef cattle
finishing systems in the Eastern Plains of the
Orinoco River Basin of Colombia. Front. Anim. Sci.
4: 1103826.
https://doi.org/10.3389/fanim.2023.1103826
RamírezRestrepo, C. A., R. R. Vera, and I. M. Rao.
2019a. Dynamics of animal performance, and
estimation of carbon footprint of two breeding herds
grazing native neotropical savannas in eastern
Colombia. Agric. Ecosys. Environ. 281: 35–46.
https://doi.org/10.1016/j.agee.2019.05.004
RamírezRestrepo, C. A., R. R. Vera, and I. M. Rao.
2019c. Producción de carne, emisión de metano y
huella de carbono en sistemas de hato de cría en
pasturas de los Llanos Orientales de Colombia.
https://www.engormix.com/ganaderiacarne/
articulos/produccioncarneemisionmetano
t44332.htm
RamírezRestrepo, C. A., R. R. I. Vera, and I. M. Rao.
2019b. Environmental performance of grazing beef
cattle systems in the welldrained neotropical
savannas of Colombia: A review of results from
modelling research. In: A. Das, S. Das, S. Sarkar, A.
K. Patra, G. P. Mandal, S. Soren (Eds.). Nutritional
Strategies for Improving Farm Profitability and
Clean Animal Production. Book of Abstracts of
International Conference on Animal Nutrition.
Animal Society of India, Kolkata, India, p. 413.
https://hdl.handle.net/10568/106716
RamírezRestrepo, C. A., and R. R. VeraInfanzón.
2019. Methane emissions of extensive grazing
breeding herds in relation to the weaning and
yearling stages in the Eastern Plains of Colombia.
Rev. Med. Vet. Zoot. 66(2): 111–130.
https://doi.org/15446/rfmvz.v66n2.82429
Rao, I., J. Arango, M. Ishitani, M. Peters, M., J. Miles, J.
Tohme, A. Castro, J. A. Cardoso, M. Worthington,
M. Selvaraj, R. van der Hoek, R. SchultzeKraft, A.
Rincón, C. Plazas, R. Mendoza, M. Cuchillo, J.
Tapasco, J. Martinez, G. Hyman, D. Moreta, M.
Mena, H. Karwat, J. Nunez, G. Subbarao, and G.
Cadisch. 2015. Strategic management for forage
production and mitigation of environmental effects:
Development of Brachiaria grasses to inhibit
nitrification in soil. In: A. R. Evangelista, C. L. S.
Avila, D. R. Casagrande, M. A. S. Lara, T. F.
Bernardes (Eds.). Proceedings of the 1st
international conference on forages in warm
climates. Universidade Federal de Lavras, Lavras,
Brazil, pp. 85–102.
Rao, I. M. 1998. Root distribution and production in
native and introduced pastures in the south
American savannas. In: J. E. Jr. Box (Ed.). Root
Demographics and Their Efficiencies in Sustainable
Agriculture, Grasslands, and Forest Ecosystems.
Kluwer Academic Publishers, Dordrecht, The
Netherlands, pp. 19–42.
Rao, I. M., M. A. Ayarza, and R. J. Thomas. 1994. The
use of carbon isotope ratios to evaluate legume
contribution to soil enhancement in tropical
pastures. Plant Soil. 162: 177– 182.
https://doi.org/10.1007/BF01347704
Rao, I. M., C. Plazas, and J. Ricaurte. 2001b. Root
turnover and nutrient cycling in native and
introduced pastures in tropical savannas. In: W. J.
Horst, M. K. Schenk, A. Burkert, N. Claassen, H.
Flessa, W. B. Frommer, H. Goldbach, HW. Olfs, V.
Romheld, B. Sattelmacher, U. Schmidhalter, S.
Schubert, N. V. Wiren, L. Wittenmayer (Eds.). Plant
Nutrition: Food security and sustainability of agro
ecosystems through basic and applied research,
Kluwer Academic Publishers, Dordrecht, The
Netherlands, pp. 976–977.
Rao, I. M., G. Rippstein, G. Escobar, and J. Ricaurte.
2001a. Producción de biomasa vegetal epígea e
hipógea en las sabanas nativas. In: G. Rippstein, G.
Escobar, F. Motta (Eds.). Agroecología y
biodiversidad de las sabanas en los llanos orientales
de Colombia. CIAT, Cali, Colombia, pp. 198–222.
Rivera, B. S. 1988. Performance of beef cattle herds
under different pasture and management systems in
the Llanos of Colombia (Doctoral dissertation).
Technische Universitat, Berlin, Germany.
Roberts, A. J., M. K. Petersen, and R. N. Funston. 2015.
2015. Beef Species Symposium: Can we build the
cowherd by increasing longevity of females? J.
Anim. Sci. 93: 4235–4243.
https://doi.org/10.2527/jas.20148811
RomeroRuiz, M. H., S. G. A. Flantua, K. Tansey, and J.
C. Berrio. 2012. Landscape transformations in
savannas of northern South America: Land use/
RamírezRestrepo et al.
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
19
Romero, A. M. M., J. H. A. Cárdenas, M. E. O. Triana,
and L. G. D. Muñoz. 2018. Caracterización y
tipificación de los sistemas productivos de ceba de
ganado bovino en la Orinoquia colombiana. Zootec.
Tropic. 36: 131–143.
http://www.publicaciones.inia.gob.ve/index.php/
zootecniatropical/issue/view/10/30
Segnini, A., A. A. P. Xavier, P. L. OtavianiJunior, P. P.
A. Oliveira, A. d. F. Pedroso, M. F. F. P. Ferreira, P.
H. R. Mazza, and D. B. P. M. Marcondes. 2017. Soil
carbon stock and humification in pastures under
different levels of intensification in Brazil. Sci. Agric.
76(1): 33–40.
https://doi.org/10.1590/1678992X20170131
cover changes since 1987 in the Llanos Orientales of
Colombia. Appl. Geogr. 32: 766–776.
https://doi.org/10.1016/j.apgeog.2011.08.010
Rondón M., D. Acevedo, R. M. Hernández, Y. Rubiano,
M. Rivera, E. Amézquita, M. Romero, L. Sarmiento,
M. A. Ayarza, E. Barrios, and I. M. Rao. 2006.
Carbon sequestration potential of the neotropical
savannas (Llanos) of Colombia and Venezuela. In: R.
Lal, J. Kimble (Eds.). Carbon sequestration in soils of
Latin America. The Haworth Press, Inc.,
Binghampton, USA, pp. 213–243.
Rotta, P. P., A. C. B. M. Menezes, F. C. Costa e Silva, S
de C. Valadares Filho, L. F. Prados, and M. I.
Marcondes. 2016. Protein requirements for beef
cattle. In: S. de C. Valadares Filho, L. F. Costa e
Silva, M. P. Gionbelli, P. P., Rotta, M. I. Marscondes,
M. L. Chizotti, L. F. Prados (Eds), Nutrient
requirements of Zebu and crossbred cattle BR
CORTE, 3rd edition, Universidade Federal de
Viçosa, Viçosa, Brazil, pp.185–212.
https://editorascienza.com.br/pdfs/
Rouquette, F. M. Jr., L. A. Redmon, G. E. Aiken, G. M.
Hill, G. M., L. E. Sollenberger, and J. Andrae. 2009.
ASAS Centennial Paper: Future needs of research
and extension in forage utilization. J. Anim. Sci.
87(1): 438–446. https://doi.org/10.2527/jas.2008
1273
Sanhueza, E., L. Cárdenas, L. Donoso, and M. Santana.
1994a. Effect of plowing on CO2, CO, CH4, N2O, and
NO fluxes from tropical savannah soils. J. Geophys.
Res. 99: 16429–16434.
https://doi.org/10.1029/94JD00265
Sanhueza, E., L. Donoso, D. Scharffe, and P. J. Crutzen.
1994b. Carbon monoxide fluxes from natural,
managed, or cultivated savannah grasslands. J.
Geophys. Res. 99: 16421–16427.
https://doi.org/10.1029/93JD02918
SAS. 2016. Statistical Analysis System. University
Edition version 3.5. Cary, NC, USA: SAS Institute.
https://www.sas.com/en_au/software/university
edition.html
Saravia, F. M., J. C. B. Dubeux Junior, M. de A. Lira, A.
C. L. de Melo, M. V. F. dos Santos, F. de A. Cabral,
and V. I. Teixeira. 2014. Root development and soil
carbon stocks of tropical pastures managed under
different grazing intensities. Trop. GrasslForraj.
Trop. 2: 254–261.
https://doi.org/10.17138/tgft(2)254261
Siqueira da Silva, H. M., J. C. B. Dubeux, M. L. Silveira,
M. V. F. dos Santos, E. V. de Freitas, and M. de A.
Lira. 2019. Root decomposition of grazed
signalgrass in response to stocking and nitrogen
fertilization rates. Crop Sci. 59: 811–818.
https://doi.org/10.2135/cropsci2018.08.0523
Tedeschi, L. O. 2019. ASNASAS SYMPOSIUM:
FUTURE OF DATA ANALYTICS IN NUTRITION:
Mathematical modelling in ruminant nutrition:
approaches and paradigms, extant models, and
thoughts for upcoming predictive analytics. J. Anim.
Sci. 97: 1921–1944.
https://doi.org/10.1093/jas/skz092
Thomas, R. J., and N. M. and Asakawa. 1993.
Decomposition of leaf litter from tropical forage
grasses and legumes. Soil Biol. Biochem. 25: 1351–
1361. https://doi.org/10.1016/00380717(93)90050
L
Trujillo, W., M. J. Fisher, and R. Lal. 2006. Root
dynamics of native savanna and introduced
pastures in the Eastern Plains of Colombia. Soil Till.
Res. 87: 2838.
https://doi.org/10.1016/j.still.2005.02.038
University of Arkansas. 2019. The field capacity
calculator. https://www.uaex.edu/farmranch/
economicsmarketing/docs/FieldCapacity.xls
Urquiaga, S., G. Cadisch, B. J. R. Alves, R. M. Boddey,
and K. E. Giller. 1998. Influence of decomposition of
roots of tropical forage species on the availability of
soil nitrogen. Soil Biol. Biochem. 30: 2099–2106.
https://doi.org/10.1016/S00380717(98)000868
Velásquez, J. C., and M. Ríos. 2010. Evaluación de la
producción de carne a partir de vacas cebú de
descarte. Revista de Ciencia Animal. 3: 23–29.
https://ciencia.lasalle.edu.co/ca/vol1/iss3/2/
Vera, R. R., and RamírezRestrepo, C. A. 2017.
Complementary use of neotropical savanna and
grasslegume pastures for early weaning of beef
calves, and effects on growth, metabolic status and
reproductive performance. Trop. GrasslForrajes
Trop. 5(2): 5065.
https://doi.org/10.17138/tgft(5)5065
Sustainable fattening of cull beef cows
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20
20
Vera, R. R., and F. Hoyos. 2019. Longterm beef
production from pastures established with and
without annual crops compared with native
savanna in the high savannas of Eastern Colombia: a
compilation and analysis of onfarm results 1979
2016. Trop. GrasslForrajes Trop. 7(1): 1–13.
https://doi.org/10.17138/TGFT(7)113
Vera, R. R., C. A. Ramírez, and Velásquez, N. 2002.
Growth patterns and reproductive performance of
grazing cows in a tropical environment. Arch.
Latinoam. Prod. Anim. 10: 14–19. https://
ojs.alpa.uy/index.php/ojs_files/article/view/116
Vera, R. R., and C. Seré. 1989. On farm results with
Andropogon gayanus. In: J. M. Toledo, R. R. Vera, C.
Lascano, J. L. Lenné (Eds.). Andropogon gayanus
Kunth. A grass for tropical acid soils. CIAT, Cali,
Colombia, pp. 323–356.
VeraInfanzón, R. R., and C. A. RamírezRestrepo.
2020. Long term beef production in extensive cow
calf systems in the tropical savannas of eastern
Colombia. Rev. Med. Vet. Zoot. 67(1): 42–59.
https://doi.org/10.15446/rfmvz.v67n1.87678
Viglizzo, E. F., M. F. Ricard, M. Taboada, and G.
VázquezAmábile. 2019. Reassessing the role of
grazing lands in carbonbalance estimations: Meta
analylsis and review. Sci. Total Environ. 661: 531–
542. https://doi.org/10.1016/j.scitotenv.2019.01.130
Waldrip, H. M., R. W. Todd, and N. A. Cole. 2013.
Prediction of nitrogen excretion by beef cattle: A
metaanalysis. J. Anim. Sci. 91: 4290–4302.
https://doi.org/10.2527/jas.20125818
Wiloso, E. I., R. Heijungs, G. Huppes, and K. Fang.
2016. Effect of biogenic carbon inventory on the life
cycle assessment of bioenergy: challenges to the
neutrality assumption. J. Clean. Prod. 125: 78–85.
https://doi.org/10.1016/J.JCLEPRO.2016.03.096
Wyborn, L., L. Hsu, and M. Parsons. 2015. Guest
Editorial: Special issue rescuing legacy data for
future science. GeoResJ. 6: 106–107.
https://doi.org/10.7916/D8H131FD
Zhu, Y., L. Merbold, D. Pelster, E. DiazPines, G. N.
Wanyama, and K. ButterbachBahl. 2018. Effect of
dung quantity and quality on greenhouse gas fluxes
from tropical pastures in Kenya. Global
Biogeochem. Cycles. 32: 1589–1604.
https://doi.org/10.1029/2018GB005949
RamírezRestrepo et al.
ISSNL 10221301. Archivos Latinoamericanos de Producción Animal. 2023. 31 (1): 1  20